tek sia acetatesaseses ipditeldewnte peaagnd.@ dae)? ‘ ‘ sie sh 884 bree ay +9 = y55g 428 178 Vet gta saet orn aqoqedraepae iat OO te ie te th aoe i Dasher poate sata’ a0 and “pau tiagd paae ae ik dy aor jeite 4 4 oii oe has teh te A679; acter aaa A at Vv eateeg "4 as hiiiet Soa eye» tie oot ty 8 HHH Fiat it HR HEATH Hi ms Pt he sti seid ante ath dot if he “4 ae e eee RMT REC EH ager stissaae ana tis Hite nat rae mdae ttiacs i ats titake eh nit ae AR AWypst tat Byer Hideto t aid Spa pss \ j sa ey ; pitt es, +} ite, aye) reine re fer rr 503428) souttaatt Wade * piste Hope ae ste aa: ops sh Vile saaerel pee) Scie qn hry fe a teas hoon sayn ines Vay ie ee He te eperiorees ed iad Arey Bure atte ” ase. te yestta at dit et Wi. hae HOSS pena pein) abs 2 ‘i hie i} faerie sai 889 youre i yeas Ch phe nay ris) a erga” 4 is i nee) ay sepeae in Paphitis Winuad gL) tat satera ee 48 Baie ab igi My eh vhrece 1.95 Fee ge ihe erst pt RIS kt hua bade a5 ph ee 4 Hh Ni teeses ne Maseneteets oo a? is ieee a8 peenerey) 4 a a thisige pial nye bee jes re se, at tis ne) Sine ; ra? a Aaa pe Be Py oy Rata Sida anypeye Bare nit ee We + a Ba a ne __ hioata 4 5 y he eins ae pabaiiay a ate isn He pilsas fae ths rive ou bahe ana eH * ey ‘] vine Ba i ae i idan aught ites 3 thy a beeen Ae \ 4 thes aye ree hay fais Rae ee bbl aay tates agai he sDaled! an Ne Spe ‘aie on bean ts 433 ’ 4 hebpepe net r. ct af woh ae ai VOLS Doe Hie Dake ted isa an gas ” a = mah ees eras a yaa ey iis pada! pa Pd eta vga Nit > irev ieee SOA ne ait ter aetogt 8 th 7 4 yey at i 3 wsdl veh, ete DUE enett os eit iy at fy Cu segiaaes rt ibe Rae A ve pats ; Baath tie SDA F Seay bea Petpieseenad Sie Hel engin! Gio? sk han ty presse) nies PSSSLES D1 GUL 0 Rg nee ere ee ee RR dT “TP RSILIES (C104 (HS 00 Ce eng erie eae eR ae nee Ser ee RSME MSE CRUITE, BODY: NVA TAU (2c 27 cece gaa ANS 220. 2). sen ee 89 Deeper muscles of thigh and shank, lateral.............. 92 Deepest muscles of hip and thigh, lateral................ 95 Muscles ‘of upper shank; lateral: $102. 727 5 30= eee 96 Deeper muscles of upper shank, lateral.................. 98 Third layer of muscles of upper shank, antero-lateral...... 100 Deepest muscles of upper shank, lateral................. 101 Superficial muscles of vertebral column, lateral........... 111 Diagram of details of M. longus colli.................... 113 . Diagram of details of Mm. ascendentes.................. 115 Deep muscles of neck; dorsal. 2... 2.5..5. oe Ly, viii MATERIALS Preserved specimens, other than skins, of Whooping Cranes are even fewer than the living birds. For example, the senior author, in gathering material for an osteological study of the genera Grus, Balearica, and Anthropoides, could not locate a single complete skeleton of Grus ameri- cana and only two that were reasonably complete. Most osteological specimens consist of one or a few bony elements. There is an unverified report that the Senkenberg Museum in Frankfurt has complete skeletons of a male and a female that died in captivity, but we have not been able to check this. It is our belief that the three birds used in this study were the only ones preserved as alcoholic specimens. In the fall of 1948 the first specimen was received from the United States Fish and Wildlife Service through the courtesy of Mr. Robert P. Allen and Dr. John W. Aldrich. It had been skinned, and the head was missing, as were the entire wing and the leg distal to the knee. For ship- ment from Texas to the United States National Museum the carcass was frozen. At the museum it was preserved in 70 per cent alcohol for ship- ment to the University of Illinois. The following information concerning this specimen is taken from a letter of January 18, 1949, to Harvey I. Fisher from Robert P. Allen: It is not known when this individual received the original wound that broke the bones of the left wing. It was first noted in December, 1947, when my notes read: “Dec. 14, 1947—Ground inventory. Note on the ‘Summer Pair’: evidently both of these birds have injured left wing.” This was on the Aransas National Wildlife Refuge near Austwell, Texas, where most of the surviving Whoopers winter. This “pair” (They may be merely companions) had spent the summer, and it was now presumed that failure to migrate was a result of injured wings that prevented flight. A certain percentage of the Texas flock lives outside the refuge area and are subject to illegal shooting by waterfowl hunters in the nearby bays. This pair became accustomed to whole yellow corn bait as the winter ad- vanced and by this means were kept on Long Pond on the refuge and away from the shore of the Intercoastal Waterway, where they often fed on blue crabs (Callinectes sapidus), mud shrimps (Callianassa) and other small in- vertebrates of the region, as well as killifishes, these items comprising their normal winter food. However, on March 14th I left for Louisiana to investi- gate the status of two captive Whooping Cranes we were holding there and did not return until March 20th. During this brief period the two injured Cranes presumably wandered once more to the shore of the Waterway. On March 24th this individual of the pair was found lying down on the edge of Salada Pond. Dried blood (at first thought to be dried mud) was noted on its neck and upper breast. When I tried to approach, it walked in labored manner towards Mustang Slough. The next morning, with the help of refuge personnel, the bird was captured. Left wing was hanging heavily, breath was labored, bird thin and weak. We carried it to headquarters, phoned for a vet but the bird died at 11:30 a.m. 1S) THE MYOLOGY OF THE WHOOPING CRANE With Russel Clapper’s assistance the bird was skinned and it was discovered that the neck had been pierced by a bullet, apparently from a .22 calibre rifle. Trachea was shattered and wound clotted. Examination also showed that the old wound had broken humerus completely, the ends being separated by weight of wing so that healing was impossible. The ulna was shattered. The following measurements were made before skinning: Wreightiattimeof deaths 002505 ae aa ee 10 Ibs. 5 ounces. Gulmienta. (bles ax ene el nee eee Bs eae tte 133 mm. ANAT OVERS oh Meco eee tod Selec Sar 1 AO Mee Nad ee 619 mm Bhs Fete caee see e aaye- Le ts Sa Ration ene Pee 329 mm RATS W Segue eyecare ee tae se eo ee eR ee ees 278 mm. Midztoe (tor nice tee re eee ae ee cd 138 mm. Far EEOC s Rearrange eto ee 30 mm. Outsideitoen Fees alee eee oe er ee 110 mm. MM SIGS tO CS wok coi secon wed eee eT RSE 87 mm. Palmation. (relaxecl) oy bace 2 Sap us code wees ae 24.5 mm. (depth) Ralmationn (extended) rise tare ee 38 mm. Spread =. so... see eben ee ee sn. 22209 mmiee Commences) HEPES. << CCOTMLO UT) PAE es ERE eh: Laces ae 1289 mm. dee Th: 3(eh@rds) Pt) aes aoa ha a ee 1257 mm. Basecporstansusm (CincUIns) a ciante ae ae ee 1 & % inches. Extreme. waclthe Ofetracksws ssote seein) see 7 inches Brains weighed 314 grams. Bird lice, eyeballs and oil gland were sent to Dr. Aldrich in separate con- tainers. The skeletal remains of this specimen are now No. 347355 in the series of the United States National Museum. Dissection showed it to be a female. The second specimen (KU No. 31198) was found October 31, 1952, by Mr. Thane S. Robinson. The location was 8.5 miles south of Sharon, Barber Co., Kansas. Mr. Robinson telephoned Dr. E. Raymond Hall who notified Dr. Clarence Cottam of the United States Fish and Wildlife Service. Employees of this service picked up the crippled crane that same day and transported it by truck to Mineral Wells, Texas, where it died November 1. The carcass was then taken to San Antonio. On November 6 the now frozen specimen was flown by pilot Hanson to Topeka, where it was secured by Drs. Tordoff and Hall. Measurements of this female, taken before skinning, include: Weight—13.25 Ibs. Middle toe with claw—132 mm. Chord of wingspan—7 ft. Exposed culmen—152 mm. Chord of wing—612 mm. Bill from ant. end nostril—90 mm. Arc of wing—632 mm. Angle of gonys to tip of mandible Tarsus—309 mm. (= length of gonys in Baldwin, etc.)—66 mm. No physical injuries could be found when the bird was skinned by Tordoff and Fisher, or even when it was dissected. There was no evi- dence that the bird had been shot. MATERIALS 3 In late November, 1952, we received a report that a bird had been shot in Canada and transported to Texas. It subsequently died, was frozen, and shipped to us December 1, 1952. We skinned this female and sent the skin to the National Museum of Canada. The skeleton is preserved at Southern [linois University. The history of the bird is pieced together in the following excerpts from letters to Harvey I. Fisher: From Mr. Fred Bard—March 6, 1953: To review the events that took place when we were notified of the capture of a wounded Whooping Crane at Griffin, Saskatchewan. The bird was appar- ently seen a week or two earlier by Mr. Orville Blosser, a nephew of Mr. P. E. Barlow of Griffin, a farmer, who saw and caught the Crane on October 30th. The Crane was apparently walking around in a stubble when he spotted it quite early in the morning. He pursued it with the truck, chasing it into a willow bluff in the field. He threw his coat over the bird and picked it up. The bird was placed in the back of his truck and he drove to Weyburn, where he contacted Mr. Jim Mahoney of the Weyburn Branch of the Fish and Game League. Mr. Mahoney ’phoned me reporting the capture of a bird and that it had been injured. I instructed him to call in a veterinarian and do whatever they could for the bird. When Mr. Lahrman and I lifted the bird from the crate and placed it on the straw, we noticed one leg badly twisted and on examination found the socket joint to be completely through the skin and twisted around in the wrong way. I then contacted Dr. H. L. Watson, a physician and surgeon who agreed to come out and examine the bird. That evening Dr. Watson checked the joint. I had in the meantime set it as best I could and put it in splints. The joint seemed to be properly set and Dr. Watson sutured the tendons and the skin covering the joint; antiseptics were used and the damaged leg joint placed in a cast. The bird was fed from time to time. The first feeding was a little water, and the next fresh goldeneye fish. It seemed to take these reasonably well, and in a few hours seemed to look somewhat brighter. Later attempts were made with wheat soaked in water, and some milk with aureomycin given by Dr. Watson. On Sunday afternoon the plane, dispatched from Minnesota with Ross Han- son as the pilot, left Regina with the injured bird for San Antonio Zoo. Mr. Ralph Stueck of Abernathy offered to accompany the bird on this trip to feed and water it and so relieve the pilot of these duties. From Mr. Rossalius C. Hanson—February 6, 1952: Mr. Bard, or his associates, picked up the crippled Whooper from a rancher in the vicinity of Weyburn, Sask. The bird, at that time, had a crippled left wing and broken left leg. The wing injury appeared to be a shot wound, while the leg injury was believed sustained during capture by the rancher. He had an M.D. work on the bird and they amputated the wing at the wrist and set the leg in a cast. The bird was treated with penicillin and oral aureomycin. At the time I picked up the bird it had a bad rattle in its throat, indicating a respiratory condition. This may have been due partially to the administering of the oral aureomycin with a syringe and on the other hand it may have been an outcome of the bird’s lowered resistance. During the flight from Regina to San Antonio the bird was force-fed, but regurgitated most of the food during the trip. It retained some meat particles 4 THE MYOLOGY OF THE WHOOPING CRANE fed it on the second day of the trip. Soaked wheat and eggs were not retained. The bird was picked up on Nov. 2, by myself at Regina, and deposited at the San Antonio Zoo on Nov. 4. No drugs were administered during the flight. The bird was in a very weakened condition at the start and during the flight did not show any improvement. I delivered the bird to Mr. Fred Stark, Director, San Antonio Zoo, San Antonio, Texas and he administered to the bird from then on. From Dr. Clarence Cottam—December 24, 1952: Unfortunately, I do not have the exact date it died, but it was within a day or so of November 6. No measurements of significance could be made from this last speci- men which was badly emaciated and had a wing amputated at the wrist as well as a broken leg. ACKNOWLEDGMENTS Our appreciation goes to Mr. Robert P. Allen of Tavernier, Florida, for preserving the original specimen which was the major impetus in initiating this work, and to Dr. John W. Aldrich of the United States Fish and Wildlife Service for sending the specimen to the senior author. Dr. Harrison Tordoff and Dr. E. R. Hall of the Museum of Natural History, University of Kansas, were responsible for our receiving the second speci- men. Dr. Tordoff aided the senior author in skinning the bird in such a way as to make possible the preservation of a complete skin and a com- plete skeleton. He also supplied measurements and other data. Mr. Thane S. Robinson first located this bird in the field. We wish to thank the Canadian government for granting permission for us to retain the carcass of the Saskatchewan bird. Dr. Clarence Cot- tam of the United States Fish and Wildlife Service made all the arrange- ments. Acknowledgment of aid rendered is also due Mr. Orville Blosser of Griffin, Saskatchewan, Mr. Fred G. Bard, director of the Provincial Museum of Saskatchewan, Mr. Rossalius C. Hanson of Winona, Minne- sota, and Mr. Fred W. Stark, director of the San Antonio Zoo and Aquar- ium. All these individuals took part in the capture or care of this bird. Dr. Robert W. Storer of the Museum of Zoology, University of Mich- igan, kindly sent a Little Brown Crane to us. Mr. Charles A. McLaughlin of the Zoology Department, University of Illinois, inked and labeled the final drawings, and we are grateful for his skill and patience. Finally we express our gratitude to the Research Board of the Graduate College, University of Ilinois, for enabling the junior author to participate in the work in 1952 and 1953. His contribution as a research assistant was in part the result of financial support by the Board. UU METHODS Dissection proceeded very slowly and copious notes were taken on all organ systems. We attempted to gather all possible information. How- ever, having only three specimens, it was not possible to treat all systems completely; some structures were invariably displaced and ruptured in work on other features. For example, innervations in the head region could not be studied without destroying muscles, and it was not simply a matter of taking off a layer of muscle and the attendant nerves. Life- size, or in many instances twice life-size, scale drawings were made as we dissected. One major difficulty was that all specimens had been frozen before we received them. In frozen condition the tissues are very brittle and easily broken by any strain; such breakage was sometimes extensive in the neck, axillary, and pubic regions. Often it was necessary to study all three specimens to obtain a clear picture of the muscles in a region. In general, the procedure was as follows: As the first bird was dis- sected, notes and drawings were made. Subsequent work on the other birds was compared with these notes and drawings. Any variations be- tween individuals or between sides of the same individual were recorded in notes and drawings. Finally, a rough manuscript was prepared and checked against all dissection notes and drawings. The final drawings represent composites in part, for we have tried in most instances to illus- trate the “usual” condition, but all the illustrations are made from speci- mens of Grus americana and to accurate scale. As discussed previously, the nomenclature of avian muscles is difficult. With one exception (M. dermoglossus), we have not introduced any new names. We have tried, by careful study and comparison, to use the most appropriate names already in literature. For names of muscles in the wing, tail, and leg we have followed Fisher (1946), who prepared comparative tables of their nomenclature. For the rest of the muscles we have included similar lists of names at the beginning of the appropriate section. The order of discussion of muscles in each section follows the sequence of listing in the table. The osteological terminology follows Howard (1929). Both authors have been active in all aspects of this anatomical study. No part can be said to be entirely the work of either one. Any part first done by one of us has been checked in detail by the other, on the same or subsequent specimens. MUSCLES OF THE SKULL AND JAWS The muscles are listed here in the sequence followed in the text. M. dermo-temporalis M. dermo-temporalis—Shufeldt (1890:5-6) part M. cranio-cervicalis—Edgeworth (1935:283) M. cucullaris, caput portion M. cucullaris, kopftheil—Fiirbringer (1888:1056), Gadow (1891:107-109) M. complexus—Gadow (1891:107-109), Shufeldt (1890:263-286), Palmgren (1949:194-195) M. complexus—Boas (1929:189-193) M. biventer cervicis M. biventer cervicis—Shufeldt (1890:270-272), Gadow (1891:107-109), Boas (1929:161-164) M. biventer—Palmgren (1949:194) M. splenius capitis and M. splenius accessorius M. rectus capitis posticus major—Shufeldt (1890:268-270) M. rectus capitis posticus—Gadow (1891:112) M. splenius capitis—Boas (1929:164-169), Palmgren (1949:194-195) M. depressor mandibulae M. biventer maxillae—Shufeldt (1890:18-19) M. digastricus s. depressor mandibulae—Gadow (1891:318-319) M. depressor mandibulae—Edgeworth (1935:109), Hofer (1950) M. rectus capitis lateralis M. rectus capitis anticus minor—Shufeldt (1890:265-266 ) M. rectus capitis anticus minor s. lateralis—Gadow (1891:120-121, pl. 18a) M. rectus capitis lateralis—Boas (1929:194), Palmgren (1949:202) M. rectus capitis superior M. rectus capitis lateralis plus M. trachelo-mastoideus—Shufeldt (1890: 289-290). Shufeldt must have been partly in error in thinking that the M. longus lateralis cervicis et capitis of Gadow was the same as these muscles of Shufeldt’s description. These muscles may correspond in part to the capitis part of Gadow’s muscle. The remainder of M. longus lateralis cervicis et capitis compares well with the M. obliquus colli slips described and figured by Shufeldt. part M. spinalis cervicis plus part M. longus lateralis cervicis et capitis— Gadow (1891:110, 116-117) M. rectus capitis superior—Boas (1929:195-196) M. rectus capitis ventralis lateral part : not described: ?part M. flexor capitis inferior—Shufeldt (1890:267-268) part M. rectus capitis anticus major s. medialis—Gadow (1891:120, pl. 18b) M. rectus capitis ventralis—Boas (1929:194-195), Palmgren (1949: 202-203) medial part part M. flexor capitis inferior—Shufeldt (1890:267-268 ) lod i M. THE MYOLOGY OF THE WHOOPING CRANE part M. rectus capitis anticus major s. medialis—Gadow (1891:120) M. rectus capitis ventralis—Boas (1929:194-195), Palmgren (1949: 202-203 ) flexor colli brevis Ppart M. rectus capitis lateralis—Shufeldt (1890:289) ? M. lateralis colli—Gadow (1891:119) M. flexor colli brevis—Boas (1929:196-197) . flexor colli profundus Ppart M. longus colli (inferior oblique part? )—Shufeldt (1890:287-288) part M. longus colli anticus—Gadow (1891:118-119) M. flexor colli profundus—Boas (1929:197) . adductor mandibulae externus superficialis part M. temporal—Shufeldt (1890:16) parts 1 and 2 M. temporalis—Gadow (1891:320-321) part M. adductor mandibulae externus—Edgeworth (1935:58-60) M. adductor mandibulae externus superficialis—Lakjer (1926:45-46), Hofer (1950:438-468, fig. 15, A. e. sf. and A. e. sf. b.) adductor mandibulae externus profundus part M. temporal—Shufeldt (1890:16) Ppart 3 M. temporalis—Gadow (1891:322) part M. adductor mandibulae externus—Edgeworth (1935:58-60) M. adductor mandibulae externus profundus—Lakjer (1926:46), Hofer (1950:464) . adductor mandibulae medius part M. temporal plus M. pterygoideus externus—Shufeldt (1890:16, 20-21) M. ethmo-maxillaris plus M. quadrato-maxillaris—Gadow (1891: 322-323) ? M. adductor mandibulae posterior plus ? M. pseudotemporalis profun- dus—Lakjer (1926:54-55, 63-65) part M. adductor mandibulae medius—Edgeworth (1935:58-59) ?deep part M. pseudotemporalis (M. quadrato-mandibularis )—Hofer (1950:478) adductor mandibulae externus medialis part M. temporal plus M. masseter—Shufeldt (1890:16) parts 1, 2 and ?3 M. temporalis—Gadow (1891:320-322) part M. adductor mandibulae externus—Edgeworth (1935:58-60) M. adductor mandibulae externus medialis—Lakjer (1926:46), Hofer (1950:438-468) . pseudotemporalis part M. temporal—Shufeldt (1890: 16) M. spheno-maxillaris—Gadow (1891:323) M. pseudotemporalis superficialis—Lakjer (1926:63-65) part M. adductor mandibulae medius—Edgeworth (1935:277) ? superficial part M. pseudotemporalis—Hofer (1950:468-477 ) pseudotemporalis bulbi M. pseudotemporalis bulbi pterygoideus ventralis part M. pterygoideus internus—Shufeldt (1890:20) part Mm. pterygoidei—Gadow (1891:323-325) ventral part M. adductor mandibulae internus—Edgeworth (1935:58, figs. 605c, 607) M. pterygoideus ventralis Lakjer (1926:figs. 124, 127) Lakjer (1926:65-67), Hofer (1950:481-495) MUSCLES OF THE SKULL AND JAWS 9 M. pterygoideus dorsalis part M. pterygoideus internus—Shufeldt (1890:20) part Mm. pterygoidei—Gadow (1891:323-325) dorsal part M. adductor mandibulae internus—Edgeworth (1935:58, fig. 605c) M. pterygoideus dorsalis M. protractor pterygoideus Ppart M. entotympanicus—Shufeldt (1890:19-20) part 4b M. temporalis (M. orbito-pterygoideus) 319-323, pl. 27 fig. 4) M. protractor pterygoideus—Lakjer (1926:23-25), Hofer (1950: 432-438 ) part M. spheno-pterygo-quadratus—Edgeworth (1935:57) M. protractor quadratus Ppart M. entotympanicus—Shufeldt (1890:19-20) part 4a M. temporalis (M. orbito-quadratus )—Gadow (1891:319-323, pl. 27 fig. 4) M. protractor quadratus—Lakjer (1926:23-25), Hofer (1950:432-438) part M. spheno-pterygo-quadratus—Edgeworth (1935:57) Lakjer (1926:65-67), Hofer (1950:481-495) Gadow (1891: M. dermo-temporalis Figure 1 shows M. dermo-temp. arising from the same fascial layer that forms part of the origin of M. depress. mandib. This fascia is attached to the outer face of the opisthotic process, posterior and dorsal to the external auditory meatus. From the origin the muscle, which is about 3.5 centimeters in length, courses postero-dorsad across the anterior part of M. rectus cap. lat., the most dorsal part of M. geniohy., and the side of the middle of the length of the caput part of M. cucullaris to insert on the skin near the mid-dorsal line over the articulation between the second and third cervical vertebrae. M. cucullaris, caput part It may be observed in figures | and 4 that the cephalic part of M. cu- cullaris lies dorsally on the neck just posterior to the head but passes postero-ventrad to assume a lateral and even a ventral position at the posterior end of cervical vertebra number 4. It is a thin sheet of muscle with much aponeurotic connection to the underlying muscles and, in the dorsal midline, to its counterpart of the opposite side. Origin—Although there is much fascial connection posteriorly, par- ticularly in the dorsal half of the neck, the primary origin is from the diapophysis of the fourth cervical vertebra. It is difficult to be more spe- cific regarding the origin because of the intimate relationship of fascial elements in this area. Insertion.—Anteriorly the muscle attaches to the most superficial part of the occipital crest from the midline laterad. The medial part of the insertion is immediately superficial to the insertion of M. biventer cerv., and the lateral part to the insertion of M. splenius cap. (fig. 4). Uf U U b . if ° G 1109 si Buo| ‘yd 40) “juan ‘doo snjoad ‘yd ‘yaadns yuan “BKsayd snapiokyoiueb xuAJD| 48AO DIDSD4 peyoo.y snapiokyoayon,} ‘qipuow = ‘ssaidap oAyo\o JOAyojo4a9 ‘A189 “Ub WN ss Snaployo|Ays ‘ SS _ a= =. pua ‘sod Ss = SS ‘yd "40 ‘snipaw ‘qipuow ‘ppo yd ‘yaadns ‘‘yuadns -yxa ‘qipuow ‘ppp SIAQ1Q 1]09 “xKa}} *y0| ‘dod snjoa4 ‘dns ‘dod snjoau yd yndoo ‘stpjjnono ‘dod sniuajds ‘dwajowuap dijs |D4yso4 "youd “xa ‘qipubw = “ppo dis daap ‘ysadns xa ‘qipuow “‘ppo “Bll ‘quoysod DIDSD} |O}IG’o0 MUSCLES OF THE SKULL AND JAWS elt M. biventer cervicis The long, attenuated muscle is fleshy in the anterior 4 centimeters of its length but at the posterior end of the second cervical vertebra a long, thin tendon is formed (fig. 3). The tendon goes posteriorly on the neck just lateral to the midline. Anteriorly the muscle lies between the sheets of muscle making up the caput part of M. cucullaris and M. splenius cap. In the middle of the length of the twelfth cervical vertebra the tendon becomes fleshy and forms a belly of the same proportions as the anterior belly. The posterior belly extends to the middle of the fifteenth vertebra, where a tendon is again formed. Origin —The posterior end of the tendon just mentioned arises from the anterior edge of the neural crest of the sixteenth cervical vertebra and is closely associated with other muscles attaching here. Insertion—Tendinous and fleshy insertion is made on the occipital crest just lateral to the midline and deep to the insertion of the cephalic part of M. cucullaris; it is about 0.5 centimeter wide (fig. 4). orig., pseudotemp. orig., depress. palp. orig., add. mandib. ext. medialis orig., lev. palp. orig., add. mandib. ext. superf. ae, ~ SUP. nee se deep slip : PF int, Ae i i Pet. scp bs in reel ~ orig., pseudotemp. bulbi ae Win, rect: insert., add. mandib. == ~~. orig., quad. protract. ext. superf., Willie ees superf. slip Orig., add. mandib. ext. prof., rostral slip orig., depress. mandib. insert., rectus cap. lat. pt. insert., add. mandib. medius, lat. pt. insert., add. mandib. ext. prof., rostral slip pt. insert., add. mandib. medius Origs., add. mandib. medius + add, mandib. ext. prof. i insert., add. mandib, neg oapemac ext. prof., caudal slip Fic. 2. Diagram of the muscular attachments on the skull and lower jaw. M. splenius capitis The thick, triangular belly lies on the dorso-lateral part of the vertebral column over the atlas and axis (figs. 1, 3). It is, for the most part, covered by the caput part of M. cucullaris and M. biventer cerv. The antero- ventral portion of the muscle lies just beneath the insertion of M. rectus cap. lat., while the postero-lateral edge covers parts of Mm. splenius access. and rectus cap. sup. Origin—There is fleshy and tendinous origin from the side of the neural crest of the axis and from fascia overlying the atlas and extend- ing to the foramen magnum. On the axis the origin is dorsal to the middle 2 THE MYOLOGY OF THE WHOOPING CRANE of the belly of M. splenius access. and to the Mm. interspinales arising from the antero-ventral part of the crest (fig. 3). Insertion —A wide line of fleshy insertion (fig. 4) is present in the dorsal part of the occipital region and extends ventro-laterally on to the base of the opisthotic process. At its ventral end the insertion is medial to the insertion of M. rectus cap. lat. and the dorso-lateral part of the origin of M. depress. mandib. and is almost continuous with the heavy, medial origin of this same muscle. M. splenius accessorius Although the muscle does not connect directly with the skull, it is in- cluded here because it is apparently a part of the splenius complex (figs. 3, 5). It is a flat band which is covered anteriorly by M. splenius cap. and which overlies the anterior end of the M. spinalis cerv. complex; it runs from the third cervical vertebra to the atlas. The dorsal edges of the second and third fasciculi of M. rectus cap. sup. lie under the ventral border of the anterior half of M. splenius access. Origin.—The lateral surface of the neural crest of the third cervical vertebra provides fleshy and tendinous origin. Insertion.—The insertion is primarily tendinous on the posterior sur- face of the most ventral and lateral extent of the atlas (fig. 5), and con- tiguous with the ventral edge of the interspinales muscles. M. depressor mandibulae In figure 1 the muscle may be seen to lie across the suspension of the lower jaw; the medial part of the muscle is best illustrated in figure 7. Posteriorly and distally the lateral and medial parts are inseparably fused, but the inserting tendon of M. rectus cap. lat. forms the separation in the dorsal part. Deep to the medial part is the ligamentum mandibulae. The antero-ventral edge is in contact with M. pteryg. vent., and M. stylohy. arises just anterior to the anterior edge of the belly of M. de- press. mandib. Above the origin of M. stylohy., M. depress. mandib. is separated from the external auditory meatus only by the membrane lining this canal. It would seem that any swelling of the belly, as when the muscle contracts or the posterior end of the mandible arcs dorsad, would tend to close the opening of the ear. The dorsal fascial part of the lateral portion is in part covered by M. dermo-temp. Origin—The extent of the origin from the postero-lateral area of the occipital region and the posterior face of the opisthotic process may be observed in figures 2 and 4. On the lateral surface of the process the origin is tendinous and fleshy, but it goes dorsad as a tendinous apo- neurosis to the level of the deep slip of M. add. mandib. ext. superf. From the most ventral and medial tip of the opisthotic process, a strong tendon forms a major part of the origin of the deeper, medial portion of the muscle; in fact, many of the tendinous fibers pass through the fleshy belly and thus go from origin to insertion and serve as a ligament. MUSCLES OF THE SKULL AND JAWS 13 Insertion—The superficial and most ventral part of the insertion is tendinous on the postero-ventral corner of the lateral surface of the mandible. The primary insertion is fleshy on the entire area of the pos- terior end of the mandible, bordered anteriorly by the posterior end of M. pteryg. vent. (fig. 1). M. rectus capitis lateralis The asymmetry visible in the Mm. recti cap. vent. is not so evident in the present muscles. On the left side the belly of M. rectus cap. lat. is intimately connected with the deep lateral edge of the belly of M. rectus cap. vent., probably as a secondary effect of the asymmetry of the latter pseudotemp. add. mandib. medius, add. mandib. ext. superf., deep slip biventer cerv. splenius cap. splenius access. Srd. cerv. vert. spinalis cerv. lat. pt. rectus cap. rectus cap. lat. vent., med. pt. add. mandib. ext. prof., rostral slip rectus cap. sup. flex. colli brevis longus colli Fic. 3. Lateral view of a second layer of muscles of the head and the anterior vertebral region. muscle. On the right these two recti muscles are separate and distinct. In lateral view the single slip of M. rectus cap. lat. emerges from the neck musculature just dorsal to the lateral part of M. rectus cap. vent. (figs. 1, 3, 7) and ventral to the various slips of M. rectus cap. sup. Origin—There is fleshy and fascial origin from the ventro-lateral cor- ners of the hypopophyses of cervical vertebrae 3 and 4. Insertion—A heavy tendon attaches to the posterior face of the base of the opisthotic process, deep and posterior to the lateral origin of M. depress. mandib. (figs. 2, 3, 4) and superficial in part to the most lateral edge of M. splenius cap. M. rectus capitis superior The muscle complex to be considered consists of five different fasciculi which arise from the first five cervical vertebrae and extend to the ventro-lateral region of the head (figs. 1, 3, 5, 6, 7). They are arranged 14 THE MYOLOGY OF THE WHOOPING CRANE serially so that the most dorsal goes anteriorly from the atlas and the most ventral and posterior slip comes from the fifth vertebra. From this description it is apparent that the slips increase in length from anterior to posterior in the series. In lateral and dorsal view the bundles are var- iously covered by M. splenius cap., the caput part of M. cucullaris, and M. rectus cap. lat. On the ventral side they are deep to the latter muscle and to the lateral part of M. rectus cap. vent. Origin —Fasciculus number 1, the most anterior and dorsal, arises from the most lateral extent of the anterior surface of the postzygopophyseal area of the atlas. Number 2 comes from the same part of the postzygo- pophyseal process of the axis, but its origin goes forward on a mid-lateral line on the body of this vertebra. The same part of vertebra 3 gives rise to slip 3 which is twice as large as slips 2 and 4; the origin is completely tendinous and continues anteriorly across the joint between vertebrae 2 and 3. There is fleshy origin from the ventro-lateral corner of the centrum of vertebra 2 and beneath the origin of the second part of the muscle complex. Part number 4 has fleshy and tendinous origin from the lateral tip of the transverse process of vertebra 4; the origin is closely intercon- nected with the fascia giving rise to the Mm. intertransversarii. Bundle 5 arises fleshily from the anterior edges of the posterior zygopophyseal proc- esses of vertebrae 3 and 4 and from a mid-lateral line on the centrum of vertebra 4; there is also tendinous origin from the anterior edge of the transverse process of vertebra 5. The deep surface of this fifth part of M. rectus cap. sup. gives off a small slip antero-mediad. Insertion —The four most anterior slips insert tendinously and in com- mon on the lateral three-fourths of the length of the posterior edge of the basitemporal plate (figs. 4, 8). The fifth and by far the largest slip inserts tendinously on the most antero-lateral corner of the axis and the postero- lateral aspect of the atlas. There is heavy fascia covering the articulation between the first two cervical vertebrae and attaching anteriorly to the base of the skull; the fifth slip uses this as part of its insertion. Hence the functional insertion of the slip is at least in part on the middle of the pos- terior edge of the basitemporal plate. M. rectus capitis ventralis The muscles of the two sides are not symmetrically arranged; on each side of the ventral aspect of the neck the muscle consists of a medial and a lateral part. On the left side the posterior end of the lateral portion extends fleshily to the caudal end of the fifth cervical vertebra; on the right side it goes only to the fourth vertebra. This asymmetry is also visible in the medial parts of the muscles; the left goes to the fourth vertebra and the right to the third. The variation between sides is appar- ently correlated with the position of the trachea. Superficial to the caudal end of the right M. rectus cap. vent. the trachea moves from its position on the right side of the neck to a ventral location (fig. 9). MUSCLES OF THE SKULL AND JAWS 15 In figure 7 it may be observed that the lateral part is superficial to M. rectus cap. lat. and is separated from its opposite member by the paired veins; this separation is only superficial, for along the posterior two-thirds of their dorsal edges the muscles meet in the midline. Where the two medial parts are side by side they are inseparable. Although in their posterior parts the lateral and medial slips are distinct, they are more or less fused opposite and anterior to the region of M. depress. mandib. Pesed, 3,7). Origin —The left lateral part comes from the hypopophyses of cervical vertebrae 4 and 5 (sometimes 6 as well) and from the midventral line of the centra between these hypopophyses. The anterior end of this line is continuous with the posterior end of the line of origin of the medial part, which latter origin continues on this line to the base of the skull. The right lateral part takes its principal origin from the hypopophysis of vertebra 4 but may extend on to number 5. The medial part on this side is similar in its origin to that on the left except that it goes posteriorly only as far as the hypopophysis of the third vertebra. Insertion —The lateral parts of this pair of muscles insert tendinously on the posterior surface of the occipital process (fig. 8), and the medial parts attach fleshily to the entire area of the basitemporal plate (figs. 7, 8). There may be tendinous insertion at the apex of the basitemporal plate. M. flexor colli brevis Topographically the muscle could be considered as slips of the M. rectus cap. sup. complex (figs. 1, 3, 5, 6). However, its anterior end does not reach to the head. Consequently, functionally at least, it is a separate muscle. It lies posterior and ventral to the most posterior slip of the supe- rior rectus group and is just dorsal to the cephalic end of M. longus colli. In its passage antero-ventrally it traverses the most ventral and lateral parts of the Mm. intertransversarii present here. Origin—The origin is in three parts. The most anterior comes from the ventro-lateral corner of the entire length of the axis, beneath the an- terior end of the posterior fasciculus of M. rectus cap. sup., and from the anterior end of the third cervical vertebra. Origin 2 is on the lateral end of the transverse process of the fifth vertebra, virtually in common with, but deep to, the origin of the posterior parts of M. rectus cap. sup. The last origin is tendinous from the end of the transverse process of the sixth vertebra and is superficial to the ventral edge of the Mm. intertrans- versarii (fig. 6). Insertion —The most anterior insertion is fleshy on the dorso-lateral edge of the anterior end of the most posterior slip of M. rectus cap. sup., which in turn attaches primarily to the atlas. The other slips of this flexor insert, one above the other, on the most posterior area of the transverse process of the third cervical vertebra. At its insertion the more posterior 16 THE MYOLOGY OF THE WHOOPING CRANE of these parts is fused with the anterior part of M. longus colli (fig. 6). M. flexor colli profundus This group of fasciculi (fig. 6) has sometimes been considered a part of M. longus colli. It does in fact replace M. longus colli on vertebrae 2 and 3 and coexists with this muscle on vertebrae 4 and 5. There are six distinct bundles of fibers constituting M. flex. colli prof., which is the deepest of the ventral muscles in the anterior part of the neck. insert., biventer cerv. insert., cucullaris, caput pt. occipital ridge insert., splenius cap. insert., rectus cap. lat. > orig., depress. mandib. insert., 4 slips quadrate rectus cap. sup. Fic. 4. Diagram of the muscular attachments on the posterior end of the skull. Origin.—The anterior part (slip 1) arises fleshily from the lateral part of the antero-ventral surface of the transverse process of vertebra 3. Slip 2 comes from a more medial and posterior part of the same process (fig. 6). Number 3 takes fleshy origin from the antero-ventral edge of the transverse process of the fourth vertebra and from the ventro-lateral cor- ner of the centrum of the third vertebra. The fourth slip, counting from the anterior end of the complex, originates as a fleshy and tendinous belly from the transverse process and anterior part of the centrum of vertebra 5 and from the side of the posterior end of the centrum of the fourth vertebra. Bundles 5 and 6 come from the ventral surface of the antero- lateral corner of the sixth vertebra, and to a lesser degree from the same surface of the postero-lateral corner of vertebra 5 (fig. 6). Insertion.—Slips 1 and 2 insert fleshily on the ventral part of the ante- rior end of the axis (fig. 6); the attachment of number | is dorso-anterior to that of number 2. Number 3 goes tendinously to the posterior end of the hypopophysis of the axis. Four inserts tendinously on the postero-lat- eral corner of the hypopophysis of the third cervical vertebra. Number 5 attaches to the same place on the fourth vertebra, in common with the second most anterior slip of M. longus colli. Slip 6 inserts on the same area of vertebra 5. M. adductor mandibulae externus superficialis The nomenclature of the muscles of the adductor and pterygoideus groups is in a state of vast confusion, despite the major studies by Lakjer MUSCLES OF THE SKULL AND JAWS IL (1926), Edgeworth (1935), Fiedler (1951), and Hofer (1951). Because of interspecific variation and lack of agreement between previous work- ers it is virtually impossible to assign names with confidence. We agree with Beecher (1951:415) that “the jaw musculature appears confusing when fully labelled... .” We have not, with our few specimens, con- ducted any independent investigations of homologies, and we only use the existing names (pp. 7-9). The most superficial of the adductors is M. add. mandib. ext. superf., which has two parts—a superficial and a deep layer. The outer layer does not always cover the dorsal part of the deep layer (fig. 1), lies just be- neath and somewhat posterior to the eye, and is parallel and anterior to M. add. mandib. ext. prof. At its antero-ventral extent it is immediately anterior to the outer part of M. add. mandib. medius. The deep pinnate fasciculus of M. add. mandib. ext. superf. (figs. 1, 3) in some specimens is the most superficial muscle of the upper part of the temporal fossa; Mm. pseudotemp., add. mandib. ext. medialis, and add. mandib. ext. prof. are deep to it. In its distal part the muscle is covered by the superficial part of M. add. mandib. ext. superf. and by the lateral part of M. add. mandib. medius. Origin—The superficial layer originates from the fascia overlying the middle of the length of the deep layer and from a tendon which passes beneath the postero-ventral part of the deep layer to attach to the pos- terior rim of the temporal fossa. The deep part comes fleshily from an extensive area of the dorso-posterior region of the temporal fossa (fig. 2). Insertion —The superficial fasciculus inserts tendinously and fleshily on the dorso-lateral edge of the mandible for a distance of 1 centimeter, starting just anterior to the anterior edge of the dorsal part of the inser- tion of the external part of M. add. mandib. medius. (figs. 2, 7). This at- tachment continues forward, on the dorsal rim of the mandible, to the region of the junction of the maxillary and quadratojugal bones where it ends on the connective tissue in the corner of the mouth. Laterally the insertion is dorsal to the anterior part of the insertion of the external portion of M. add. mandib. medius. Immediately ventrad to the postorbital process of the skull, the bi- pinnate belly of the deep slip forms a calcified tendon (fig. 3) which inserts on the coronoid process of the mandible between the two layers of M. add. mandib. medius (fig. 5). The insertion is posterior to the in- sertion of the superficial slip. M. adductor mandibulae externus profundus The muscle has two parts—a rostral and a caudal (figs. 1, 3, 5). Other than the posterior part of M. add. mandib. medius (?M. add. mandib. post. of Lakjer, 1926) this is the most posterior of the external adductor muscles. It is posterior and in antero-ventral part deep to M. add. mandib. medius. The dorsal end of the rostral part is covered by the deep slip of 18 THE MYOLOGY OF THE WHOOPING CRANE M. add. mandib. ext. superf. and in turn completely covers the caudal part. The rostral slip is bipinnate (figs. 1, 3) and the caudal slip simple (fig. 5). Origin.—Figure 2 shows the fleshy origin of the rostral slip from the postero-ventral area of the temporal fossa and from the dorso-lateral sur- face of the orbital process of the quadrate. Beneath the fleshy origin there are several strong bands of calcified tendon which extend from origin to insertion; some of these come from the antero-dorsal part of the supra- meatic process. The anterior edge of the orbital process of the quadrate provides fleshy origin for the caudal slip (fig. 2); the area is limited to the ventral half of this edge, for the posterior slip of M. add. mandib. medius arises from the dorsal half and is continuous and inseparable from the dorsal end of the caudal part of this profundus muscle. Insertion—The rostral slip inserts tendinously on the dorso-lateral corner of the mandible, beneath the middle of the orbital fossa, deep and dorsal to the insertion of the main, external body of M. add. mandib. medius, and anterior to the insertion of the caudal fasciculus. The pos- terior end of the line of attachment is 2 centimeters from the posterior end of the mandible. See figure 2 for an illustration of both insertions. M. adductor mandibulae medius As we understand the conformation of the muscle, it consists of a medial and a lateral part, which together make this the largest of the jaw adductors. The lateral part may be seen external to the mandible (figs. 1, 3, 7), and the medial slip is beneath the eye as the deepest of all adductors in this area (figs. 5, 7, 8). The lateral part is, in its dorsal half, deep to the superficial fasciculus of M. add. mandib. ext. superf. Its disto-anterior edge is in contact, on the outer surface of the mandible, with this superficial fasciculus; its posterior border is over the M. add. mandib. ext. prof. In some specimens the postero-ventral corner is in contact with the superficial part of M. pteryg. vent. The lateral and medial parts are separated on the dorsal ridge of the mandibular ramus by the insertions of both fasciculi of M. add. mandib. ext. superf. and by that of the rostral bundle of M. add. mandib. ext. prof. (figs. 2, 5). Origin.—A calcified tendon from the postero-ventral part of the squa- mosal bone, but primarily from the post-temporal process of this bone, constitutes the origin of the lateral part (fig. 3). The tendon is deep to the ventral edge of the deep part of M. add. mandib. ext. superf. and immediately dorsal and superficial to the upper edge of the rostral slip of M. add. mandib. ext. superf. The medial part arises fleshily from the dorsal part of the lateral face and from the anterior surface of the orbital process of the quadrate (fig. 2); the medial area of this process provides some fleshy origin. The dorsal part of the fleshy origin is overlaid by MUSCLES OF THE SKULL AND JAWS 19 heavy fascia. Since M. pseudotemp. inserts on the dorso-medial edge of the medial part, it should probably be considered a part of the origin. Insertion.—The lateral part inserts directly on the external surface of the mandible by means of muscular fibers; the region of insertion in- cludes all that part of the mandible covered by the muscle in figures 1 and 3. (See also fig. 2). The medial part of M. add. mandib. medius at- base of neural crest splenius colli no. 6 intertransversorii splenius colli no. 5 on vert. no. 6 — splenius access. tendon, pseudotemp. longus colli flex. colli brevis 8 add. mandib. ext. medialis interspinales pseudotemp. bulbi pseudotemp. lacrimal gland 5th cranial nerve Fic. 5. Lateral view of the deep muscles of the head and a third layer of muscles in the anterior vertebral region. ea o ee) 23 ess Co So = je 39 Sa wr = ao) , ® = = @ . . a as a za 5° 5 : a no) oo s ys o = ~- oa i =] Za £ 20 THE MYOLOGY OF THE WHOOPING CRANE taches fleshily to the inner side of the mandibular ramus in that depres- sion beginning posteriorly at the internal articular process and going an- teriorly to the region of the surangular. At its posterior end the insertion extends dorso-laterad on the mandible just anterior to the articular facets; this latter extension may correspond to the insertion of M. add. mandib. post. of Lakjer. In lateral aspect the wide, inserting tendon of M. pseudo- temp. crosses the medial part and attaches to the medial side of the mandible. M. adductor mandibulae externus medialis As may be seen in figure 5, this is the deepest of the muscles of the adductor group in the temporal fossa; it is superficial only to M. pseudo- temp. and to the origin of M. pseudotemp. bulbi (fig. 2). Origin.—It arises tendinously from a narrow line in the temporal fossa, between the dorso-posterior part of the origin of M. pseudotemp. and the antero-dorsal section of the origin of the deep slip of M. add. mandib. ext. superf. (fig. 2). Insertion.—At the postero-ventral corner of the orbital fossa the thin band of muscle becomes tendinous and inserts as a weak aponeurosis or dense fascia on the connective tissue covering the dorso-medial border of M. add. mandib. medius (fig. 5). M. pseudotemporalis Although this is the deepest of the muscles in the temporal fossa, the anterior corner of the dorsal end of its thick belly is visible superficially (figs. 1, 3). In cross-section the belly is triangular; at the ventral end of the post-orbital process the belly is crossed superficially by the belly of M. pseudotemp. bulbi. Edgeworth (1935:277) included M. pseudotemp. as part of his M. add. mandib. medius. However, in the Whooping Crane it is a very distinct muscle as Lakjer (1926:63-65) has indicated. We were not able to find the two parts described by Lakjer. Origin.—The origin is well depicted in figure 2; usually it is fleshy but in one instance aponeurotic origin was also found. Insertion——At the postero-ventral corner of the eye, the heavy belly gives way to a strong, flat, calcified tendon which goes ventrad super- ficial to the medial part of M. add. mandib. medius (fig. 5). The tendon inserts on the middle of the width of the mandible, about 3 centimeters from the posterior end of the bone. Thus the insertion is covered medially by the insertion of the medial layer of M. add. mandib. medius. M. pseudotemporalis bulbi The thin band of M. pseudotemp. bulbi lies ventral to the post-orbital process, deep to M. add. mandib. ext. medialis and superficial to the middle of the belly of M. pseudotemp. (fig. 5). Origin.—The origin is fleshy and tendinous from the middle of the ventral part of the temporal fossa, posterior to the origin of M. pseudo- MUSCLES OF THE SKULL AND JAWS 21 temp., and deep to the anterior edge of the origin of the rostral slip of M. add. mandib. ext. prof. (fig. 2). Insertion —There is fleshy insertion on the fascia beneath the postero- ventral corner of the lacrimal gland (fig. 5). A second insertion, tendi- nous, comes from the muscle immediately ventral to the lacrimal gland and attaches to the connective tissue at the postero-ventral corner of the eye. rectus cap. sup., post. slip flex. colli brevis vert. no. 3 flex. colli prof. rectus Cap. sup., WAN ) cut end post. slip~€\\/! \) \ \ \ iZ vert. no. 4 flex. colli brevis intertransversarii Vp ; vert. no.6 Fic. 6. Ventral view of the deep muscles of the anterior vertebral region. M. pterygoideus ventralis Edgeworth (1935) synonymized all of Lakjer’s pterygoideus muscles with the muscle he called M. adductor mandibulae internus. Embryo- logically the entire muscle mass may be a single unit but, as Lakjer has shown, it breaks into a number of distinct, well-defined muscles which are variously developed in the different groups of vertebrates and which show major differences between some related species. It is of course easiest to combine all these into one unit, but this makes descriptive treatment of the variations an arduous task and leads to a glossing-over of the quite manifest diversities and separate entities of this complex of muscles. We have, therefore, retained as a matter of convenience the terms—M. pterygoideus ventralis and M. pterygoideus dorsalis. And since these pterygoideus muscles are composed of several fasciculi in the Whooping Crane, we designate these subdivisions as lateral and medial parts. The pterygoid muscles are in three layers in the Whooping Crane—a 29 THE MYOLOGY OF THE WHOOPING CRANE ventral, which is itself in two layers, and a dorsal stratum. The superficial ventral layer is undivided, but the deep, ventral part exists as lateral and medial slips. The dorsal layer is similarly separated into lateral and medial fasciculi; this sheet is here designated as M. pteryg. dors. Both ventral sheets are called M. pteryg. vent. In figure 7 both layers of M. pteryg. vent. are shown, as is the division of the deep layer into lateral and medial parts. The superficial layer is the outermost muscle of the ventral surface, crossed only by the small belly of M. stylohy. and a branch of the facialis vein. Its posterior end is marked by contact with M. depress. mandib. (fig. 1). The postero-medial os pal. [)} \ mandib. : quadratojugal J fe add. mandib. ext. superf., superf. pt. add. mandib. medius, med. pt. add. mandib. medius , lat. pt. branch, facialis vein ar SS ts. IWS = SS >< Li pteryg. dors. <4 stylohyoideus pteryg. vent., superf.