The Skeletal Musculatural System of the European Bison, Bison bonasus (Linnaeus 1758) 1

POLSKA AKADEMIA A C T A VOL. VI, 6. NAUK — ZAKŁAD BADANIA SSAKÓW T H E R I O L O G I C A BIAŁOW IEŻA 10.XI.1962. K rzysztof Ś W I E Ż Y Ń S K...
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POLSKA

AKADEMIA

A C T A VOL. VI, 6.

NAUK

— ZAKŁAD

BADANIA

SSAKÓW

T H E R I O L O G I C A BIAŁOW IEŻA

10.XI.1962.

K rzysztof Ś W I E Ż Y Ń S K I

The S k eletal M usculatural System of the European Bison, Bison bonasus ( L i n n a e u s 1758) 1 Umięśnienie szkieletow e żubra, B ison b on asu s ( L i n n a e u s

1758)

B ison ian a V III. [With 30 Figs, aud 1 table]

I. I n t r o d u c t i o n ....................................................................................................................................1 06 II. M aterial and m e t h o d ................................................................................................................ 167 III. D etailed s e c t i o n ..........................................................................................................................108 1.

168

A. M uscles of th e lips and c h e e k s ...................................................................................... 168 B. M andibular m u s c l e s ...................................................................................... - 171 2. N eck and t r u n k .......................................................................................................................... 1 7 3 A . M uscles o f th e shoulder g i r d l e ....................................................................................1 7 3 B. E paxial m u s c l e s ................................................................................................................ 1 78 C. Group of the sh ort m u scles of th e v erteb ra l colum n . . . - 184 D. T he la tera l and v en tra l m u scles of th e v erteb ra l colum n . . - 185 E. The m u scles of th e thoracic w a l l .......................................................................... 1 87 F. T he abd om inal m u s c l e s ............................................................................................. 1 89 3. T he thoracic l i m b ................................................................................................................ 1 9 1 A. M uscles of th e shoulder and a r m .......................................................................... 191 B. M uscles o f th e forearm and m a n u s ...........................................................................195 a. E xten sor d i v i s i o n ....................................................................................................... 1 9 5 b. F lex o r d i v i s i o n ................................................................................................................ 1 90 4. T he p elv ic l i m b ..........................................................................................................................200 A. Inner and v en tra l m u scles of th e r u m p ...................................................................200 B. T he la tera l m u scles of the h i p ...................................................................................201 C. T he m ed ial m u scles of th e th igh . 203 D. T he anterior m u scles of the t h i g h ..........................................................................205 E. The p osterior m u scles of th e t h i g h ......................................................................... 205 *) T he w ork w as carried out partly w ith the fin a n cia l assistan ce of the Z oologi­ cal C om m ittee of th e P olish A cad em y of Sciences. [16-.]

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K rzy szto f Ś w ieży ń sk i

F. T he m u scles of th e leg and foot a. D orso -la tera l group . b. P lan tar group . c. M uscles of m etatarsu s IV. D iscu ssion and resu lts . R eferences . . . . Streszczen ie . . . .

207 207 209 211 211

215 217

I. IN TRO DUCTIO N P u b lication s on th e E uropean bison are concerned ch iefly w ith th e p rob lem s of its biology, occurrence, b reeding in reserves, w h ile m ore recen t w orks deal also w ith its p hysiology and w ith in very n arrow lim its, w ith the path ology o f th is sp ecies. L iteratu re on th e anatom y of th e E uropean bison is not abundant. T he list given by W i l k u s (1957) of p u b lication s d ealin g w ith this su b ject has b een su p p lem en ted in recen t years by th e p u b lication s o f th e W arsaw C entre: Ś w i e ż y ń s k i & P i ­ l a r s k i (1956), P i l a r s k i & R o s k o s z (1957), P i ę k o ś & P i l a r s k i & R o s k o s z (1958), K r y s i a k (1960), R o s k o s z & E m p e l (1981), W ę g r z y n & S e r ­ w a t k a (1961) and th e w ork by B o c h e n e k (1955). A m ong th e fo reig n authors, know n to m e in addition to th e nam es g iv en by W i l k u s (I.e.) th e fo llo w in g d ealt w ith the anatom y of th e E uropean bison — P o l e i n e r (1932), M i l l o t (1945) and as a side lin e only A l b r e c h t (1944). The m ajority of w ork s are concerned w ith th e sk eleto n of the E uropean bison. The reason for this sta te of affairs w as probably to be found in th e d ifficu lties in volved in the co llectio n of m aterial. T he sm a ll num ber of liv in g E uropean bison, and in conn ection w ith th is th e in freq u en t op p ortu n ities of ob ta in in g cad avers m ade it n ecessary to w a it for a long tim e for th e com p letion of a su ita b le num ber of sp e ­ cim en s for research. B ones as bein g easier to store, w ere m ore su ita b le for this purpose. W orks on the “so ft p a rts“ of the body are in the m a jority of ca ses m erely con tri­ bu tions. This is th e resu lt, in addition to th e d iffic u ltie s m en tion ed above in o b ­ tain in g m aterial, of d ifficu lties in storin g and con servin g th e rem ain s of so large an anim al. One of the fe w w orks k n ow n to m e am ongst th ose d ealin g w ith the sk eleta l m u scu ­ lar system of the E. bison is th at by P o l e i n e r (1932). T he author had at his disp osal the lim bs of a fem a le E uropean bison w h ich had died in a Zoo. His ob servation s are concerned w ith the sk eleto n , m u scles and m ain n erve and vascu lar tru n k s of th e thoracic and p elvic lim b s of the E. bison in com parison w ith d om estic cattle. Ś w i e ż y ń s k i & P i l a r s k i (1956) d ea lt w ith th e cu taneous m u scu lar sy stem of th e E u ­ ropean bison. W r ó b l e w s k i (1927) refers to the m u scu lar system of th e E. bison in m ore gen eral term s. R esearch on th e sk e leta l m u scle sy stem o f th e E. bison in v o lved th e sta rt o f w ork pra ctically from its basic elem en ts. P o 1 e i n e r ’s ob servation s m ade on one sp ecim en only, and in addition referrin g on ly to the lim bs of th e E uropean bison, w ere checked and su p plem ented during in v estig a tio n s covered by th e p resen t w ork. In v estig a tio n w as also m ade of th e m orphology o f m u scu lar com ponents of the rem ain in g region s of the body of the E uropean bison. T he w ork does not in clu d e the group o f hyoid m u scles, the m u scles of th e ear, nose, ex te r n a l m u scles of th e ey eb a ll and of th e ta il. T hese groups w ill be in v estig a ted (or are in process o f in v estigation ) to g eth er w ith th e resp ectiv e system s.

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S k e le ta l m u scu la tu re of th e B. bonasus

II. M A TERIAL A ND METHOD The m aterial used for in v estig a tio n s con sisted of the cadavers of tw en ty European bison w h ich had died during th e period from 1950 to 1960. T he d etailed list b elo w show s th at th ese w ere bison of both se x es and of d ifferen t ages. T he num ber of m ales and fem ales, and th e d iffe re n t ages of th e sp ecim en s of both sex es, m ade it possible to sta te au th o rita tiv ely th at this m a teria l m ay form a basis for reaching ob jective conclusions, tak in g in to con sid eration both factors.

Table 1. 1 Połąga Poda Plazm a P latyna Purata Tatra Poziom ka P lanarie

_

_

— — — 1 1 11 18

— 7 7 5 6 8 5

1 2 1 13 10 26 16 2

9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20.

D ays

ö Z

F em ales 1153 866 900 893 943 858 674 219

No. of Pedegree B ooks

D ays

Y ears

'S o §

A ge M onths

1. 2. 3. 4. 5. 6. 7. 8.

Nam e

A ge W Ç}

Y ears

o £

1

i

No. of Pedegree Books

1

M aterial.

— — — 1 7 9 10 10 17 17 17

— 5 8 — 2 1 5 6 3 4 4

19 8 18 26 17 26 9 23 6 6 4

N am e

M ales

794 84/R 863 1086 824 572 785 597 546 228 575

F oetus Pogrom Porucznik P lastuś Pud (Burnus) P later P ołam aniec P luszcz Punkt P lu viu s II Puk Plato

T he m ajority of th e E uropean bison w h ich w e prepared w ere fir st su b jected to an atom opathological autopsy to d iscover th e cause of death. This caused partial dam age to certain region s of th e m u scu lar system (the w a ll of th e abdom en, the w all of th e thorax, the v en tra l m u scles of th e neck). For this reason only parts of certain of the cadavers w ere used for our in v estig a tio n s. T he sam e applied w hen d ifficu lties arose in con n ection w ith th e tran sp ort of th e w h o le cadaver, w h en the anim al died a long w ay from W arsaw , w h en th e specim en w as a large one, and w hen death took place during th e su m m er w h en th e processes of d issolu tion rapidly ensue. A n atom ical preparation w as carried out p artly on fresh and p artly on fix ed m a ­ terial. T he fresh m aterial w a s used w h en field w ork w as done, for as long as th e state of th e cadavers perm itted . T his applied esp ecia lly to th e first period of in v e s ti­ gations. A s the tech n iq u e for p reserv in g th e carcasses w a s im proved, d ifficu lties w ere elim inated. M aterial w as fix ed by k eep in g it in basins co n ta in in g a w ater solu tion of form alin (about 8°/o), or by in tera rteria l in jectio n s. A 5°/o form alin solu tion w as used for in ­ jection, or a m ix tu re of th e m eth y la ted e th y l alcohol u su a lly ob tain ab le in shops,

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K rzysztof S w ieży ń sk i

phenol, form alin , potato syrup and w a ter in the fo llo w in g parts: 60 : 6 : 4 : 5 : 25 2). T he m ixtu re used p erm itted of m ain ta in in g the cadaver in a sa tisfa cto ry condition for sev eral m onths even at th e w arm tim e of th e year. In addition th e m u scles (like other organs) did not harden as th ey do w h en form alin on ly is used, w h ich m akes preparation d ifficu lt. T he sp ecim en s w h ich w ere k ep t in a form alin solu tio n w ere p laced for 5— 6 days in a 3% w ater solu tion of sp irits of salts to so ften th em before preparation. T he draw ings included in th is w ork w ere m ade from photogram s. III. D ETA ILED SECTION 1.

HEAD

A. Muscles of the lips and cheeks

1. M. m. cutanei — of th e head are represented in European bison by m. cutaneus faciei, m. cutaneus frontalis, m. cutaneus nasi and m. cutaneus labiorum. The behavour of these m uscles is sim ilar to that of th e corres­ ponding m uscles in dom estic cattle, excep t as regards size, w hich is greater in the European bison ( P i l a r s k i & S w i e z y n s k i (1956). 2. M. orbicularis oris — does not d iffer in appearance from that in cattle. Its fibres run circularly along th e low er lip and pass on both sides into the upper lip. 3. M. incissivus m axillaris — is form ed by scattered m uscular fascicles passing from os incissivum and radiating into the upper lip. 4. M. incissivus m andibularis. Is slig h tly m ore strongly developed than maxillaris. The m uscular fascicles of this m uscle run from the anterior surface of the alveoral border for the incisor teeth of the m andible and reach to th e low er lip. 5. M. levator naso-labialis (Fig. 29, 30 — N1.), begins in fascia nasofrontalis at the lev el of th e ey e sockets and on th e dorsal surface of the nasal bone. In th is region it is covered by fascicles of fibres of m. cutaneus fron­ talis. The fibres of the flat ven ter run in an an terio-ven tral direction and th e ven ter divides above the upper edge of m. leva tor labii maxillaris proprius into the superficial and the deep layers. Fibres running along the surface cover m. leva tor labii m axillaris proprius, m. caninus and m. depressor labii m andibularis, n ex t radiating into m. orbicularis oris. The deep layer lies under the group of th e above m entioned m uscles, ending in m argo adentalis of the m axilla and prem axilla. The description given above agrees w ith that accepted in textbooks. 2) T he flu id w as prepared according to the p rescrip tion used in the D ep artm en t of V eterinary A n atom y of Iow a S ta te U n iv ersity , w h ere prop h yle alcohol, phenol, corn syrup and w ater are used. T he com position of th e flu id w e used w as adapted to the p ossib ilities of ob tain in g th e in g red ien ts in P oland,

S k eleta l m u scu la tu re o f th e B. bonasus

169

The problem of th e structure of th e m uscle referred to and th e appur­ tenance of its parts to corresponding m uscle units has for a long tim e form ed a subject for discussion. A s long ago as 1883 F r a n c k questioned th e appurtenance of th e deep layer of m. leva to r naso-labialis to this m uscular unit. A k a j e w s k i (1931) w h en investigatin g th ese m uscles in cattle, sheep, reindeer, pigs and dogs included the deep layer w ith m. lateralis nasi, at th e sam e tim e id en tifyin g the anterior part of m. m alaris as pars molaris m usculi levatoris naso-labialis. He based his statem ent on th e results of h is ow n investigation s and the works by H u b e r (1924), introducing som e supplem entary m aterial, w hich in his opinion w as essential to a discussion of this problem in anim als in general, and in rum inants and pigs in par­ ticular. K a m a n & H a m p l (1959) in their work on the superficial facial m us­ cles in Capreolus capreolus ( L i n n a e u s 1758) on the basis of th e origin and innervation of th e m uscle in question, distinguished three m uscular units: m. naso-labialis superficialis, m. naso-labialis lateralis profundus et 771. naso-labialis dorsali profundus. T he observations w e made, described above, on the structure and situ ­ ation of Til. leva tor naso-labialis are in su fficien t to solve the problem of th e appurtenance of its d ifferen t parts to this or the other m uscular unit of the European bison. 6. M. levator labii m axillaris proprius (Fig. 1, 29 — LI.) is attached to th e m axilla, orally of tu b e r malare. The fairly thick ven ter of this m uscle, th e fibres of w hich run dorsally, passes into a num ber of distinct fascicles, and later tendons radiating into the upper lip. Part of them are connected w ith sim ilar tendons on the opposite side. 7. M. caninus (Fig. 1, 29 — C.) runs togeth er w ith m. leva tor labii m a ­ xillaris proprius and m. depressor labii maxillaris. Near the insertion it is strongly connected w ith th e latter. The rounded ven ter of this m u scle, the fibres of w hich run len gth w ays, passe into the tendon, ending in the nasal m uscles. 8. M. depressor labii m axillaris (Fig. 1, 29 — Dl.) is connected by its ven ter w ith m. leva to r labii m axillaris proprius and w ith m. caninus. It originates, together w ith the latter, and in the oral section separates from it distinctly. It ends in tw o or three th in tendons situated ven tral to the tendons of the previous one, in the m uscles of nose and in the upper lip. 9. M. zygom aticu s (Fig. 30 — Z.) lies very su p erficially and begins in the fascia parotidico-m asseterica, on the extern al surface of th e m asseter. It form s a flat venter, to w hich the fibres of the m. cutaneus faciei radiate. In its continuation the ven ter of m. zyg o m a ticu s runs to th e corner of the

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K rzysztof Ś w ieży ń sk i

m outh, and in so doing covers m. buccinatorius and ends in m. orbicularis oris. N ear its term ination m. cutaneus labiorum reaches m. zygom aticu s, and in this place all three (m. cutaneus labiorum, m. zyg om a ticu s, m. buccinatorius) intertw ine their fibres. 10. M. depressor labii m andibularis (Fig. 1, 29 — Dm.) lies along th e low er edge of m. buccinatorius and com es into close contact w ith its super­ ficial layer. T he fibres, which at first run forward and dow nw ard and later on len gth w ays, form a flat ven ter and radiate into th e low er lip. The fibres of m. sterno-m andibularis w hich run from th e side of angulus m andibulae, lie on this m uscle. 11. M. buccinatorius (Fig. 1, 29 —- B, B ’, B ”.) form s the strom a to the cheek and ex h ib its a m ulti-strata structure. The m ost superficial, thin layer, the fibres of w hich run perpendicular­ ly and in tertw in e w ith the fibres of m. depressor labii m andibularis, covers num erous salivary glands and ends at th e lev el of th e anterior edge o f m. m asseter. The interm ediate layer has fibres running diagonally to th e long axis of the head and is thickest in th e aboral part of the m uscle, being inserted b etw een the alveolar border of th e m axilla and the aboral part of the alveolar border of th e mandible. The m ajority of th e fibres in this layer run to th e lem niscus inferior 3) and radiate into m. orbicularis oris. The d eep est and thinnest layer, only sligh tly separated from the form er one, has fibres running in several directions, w ith a preponderance of len g th w ise ones. T hey radiate both into lemniscus superior 3) and inferior. This layer is th ick est in the aboral part of the m uscle. From the interior it is covered by a m ucous m em brane w ith w hich it is closely bound. The description given above of m. buccinatorius in the European bison is sim ilar to th e relations obtaining in this respect in dom estic cattle. C h o m i a k (1947) m ade exh au stive studies of this m uscle in dom estic rum inants and W i 1 k u s (1957) referred in passing to this problem , using th e European bison as an exam ple. From these works it is clear, inter alia, that m. buccinatorius is characterised by a close m orphological and func­ tional connection w ith the long protuberances found in rum inants, situated on th e m ucous m em brane of the cheek. The results given by th e above tw o authors are confirm ed by our observations. 12. M. m en talis — partly covers m. incissivus ventralis, running from the anterior surface of the alveolar border for the incisor teeth of the m andible. It is fairly thick, strongly interlaced w ith abundant connective tissue. It radiates into the low er lip. 3) T erm s in trodu ced by M. C h o m i a k (1947).

S k e le ta l m u scu latu re of th e B. bon asu s

171

13. M. orbicularis oculi is strongly developed. T he fascicles of th e fibres of th is m uscle, attach ed to the bony border of th e orbit, follow a circular course. The w h o le of th e m uscle is interlaced by abundant con n ective tissue. 14. M. malaris (Fig. 29, 30 — Ma.) is rela tiv ely strongly expressed in the European bison and both the anterior and posterior parts can be d istin ­ guished, being clearly defined, but coalescing w ith each other. T he anter­ ior part beginning on facies facialis of th e lacrim al and zygom atic bones, form s a flat v en ter w ith radiating fibres w hich term inate at th e lev el of m. zygo m a ticu s, p enetrating into m. buccalis. The superior insertion of th e posterior part reaches further to th e back under the low er eyelid . N u­ m erous fibres of th is part run dow nw ards, p artially covering m. buccinatorius, w here th ey disappear fairly low dow n at th e lev el of th e clearly v isib le buccal branches of the facial nerve. T he anterior part is included by A k a j e w s k i (1931) w ith m. leva tor naso-labialis (see description of this m uscle). T he posterior part, also term ed m. depressor palpebrae inferioris, is m ore strongly form ed in th e European bison than in cattle. Sim ilar relations w ere noted in the goat (K o 1 d a, 1950). B. Mandibular m uscles

1. M, m asseter (Fig. 1 — M.M’.) of the European bison has tw o easily distin gu ishab le m ain parts, of w hich one is situated su p erficially and anterior in relation to the second, w hich has a posterior position and is p artly covered by the former. The superficial part begins at os m axillaris, its superior insertion reach­ ing backwards from facial tuberosity, through th e zygom atic bone and fa­ cies m asseterica of arcus zyg o m a ticu s to the end of processus tem poralis ossis zygom atici. The flat ven ter of th e m uscle is of fairly characteristic shape and has clearly defined boundaries, is constructed of fascicles of 'fibres running ob liq u ely, being decid ed ly perpendicular near th e posterior border. This layer ends in fossa m asseterica of the m andible (fain tly d efin ed in the European bison) on the border of angulus m andibulae, and p artly on the ram us m andibulae. The d eep est fibres of this part ’’tear a w a y ” during preparation and reveal a separate insertion, b eginning on t u b e r m axillae, and ending on th e craniolateral surface of ram us m a n d i­ bulae. The course follow ed by th e fibres is the sam e in it as in th e rem ain­ der and it is in fact d ifficu lt to speak of a separate layer. The deep part begins on th e zygom atic process of th e tem poral bone and ends on the extern al surface of ram u s m andibulae b elow th e condyle o f th e m andible.

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K rzy szto f S w iezyn sk j

T he structure and course of th e fibres of th e m asseter in E. bison, described above, correspond to th e description giv en by B o r o w i e c (1950) of th e sam e relations in cattle. The on ly d ifferen ce is that w e did not note the fu sion of the deep layer of th e m asseter w ith ra. tem poralis, m ention ed by B o r o w i e c (I.e.) 2. M. p teryg o id e u s m edialis (Fig. 2, 4 — P tm , P tm ’.) the strong levator of m andible, begins on th e perpendicular part of th e palatine bone and on th e pterygoid process of th e sphenoid bone. Just after leaving this insertion it is divided by m. p te ryg o id e u s lateralis running in this d i­ rection to its insertion on th e cranium , into tw o flat b ellies, or in fact into tw o layers. T he anterior superficial layer begins as described above, its fibres running in an backward and dow nw ard direction, being inserted in fanlike form ation on to th e m edial surface of th e an gle of the m andible. The posterior deep layer begins at the back of the above layer on th e p terygoid process of the sphenoid bone and in the p terygo-p alatin e fossa, ending on th e m edial surface of th e ram us of m andible. 3. M. p teryg o id e u s lateralis (Fig. 4 — Ptl.), form s a strong venter, the fib res of w hich run in a posterior convergent direction. It begins on the border of the p terygo-p alatin e fossa and partly in th e tem poral fossa, and ends on th e m edial side of th e condyle of the m andible. T he pterygoid m u scles of th e E. bison do not in principle d iffer from th e corresponding ones in dom estic cattle. O nly m. p te r y g o id e u s lateralis does not exh ib it th e bistratiform structure, described by B o r o w i e c (1950), despite th e fact that th e places of insertion agree w ith those described by th is author in dom estic cattle. 4. M. tem po ra lis (Fig. 3, 4 —T, T’.) fills the tem poral fossa and is attached to its border. T he m ain m ass of m uscle is sim ilar in shape to an elongated oval, w ith fibres running to th e centre, in th e direction of the coronoid process of the m andible, into w hich th ey are attached. The m uscular v en ter, th e fib res of w hich run in an an terio-ven tral direction, separates from th e anterio-dorsal surface of the part of th e m uscle described above. This v en ter is attached to th e w a lls of th e tem poral fossa, partly on crista p tery g o id e a , and th en run dow nw ards and surround th e base of the coronoid process from the front and sides. The m uscular fascicles of this part reach dow nw ards alm ost to the lev el of the arcus dentalis inferior and are inserted on th e m edial and lateral surfaces of the ramus of the m andible near its anterior border. T he presence of the separated anterior portion of m. tem poralis describ­ ed above w as confirm ed by B o r o w i e c (1950) in dom estic cattle. In th e E. bison it exten d s dow nw ards, and the m andibular insertion covers

S k eleta l m u scu la tu re of th e B. bon asu s

173

not only th e m edial surface of th e ram us m andibu laris, as in th e case of cattle, b ut also th e lateral surface. 5. M. b iv e n te r m andibulae (Fig. 2 — B v.— B v ’.) originates on th e param astoid process of th e occipital bone w ith a distinct initial tendon attached under th e inferior border of th e occipito-m andibular m uscle. The thin fusiform v en ter posterior of th is m uscle runs in the direction of th e angle of m andible, w h ere it passes through th e m edial part into a short tendon situated in a posterio-m edial position in relation to th e thickened posterior border of m. m asseter. The flatten ed ven ter anterior runs from this tendon w hich ends on th e ven tral border of the body of the m andible. A long the m edial border of th e anterior b elly of the biventer, there is a tendinous strip w h ich is an exten sion of the tendon b etw een th e anterior and posterior bellies. M. interdigastricu s runs b etw een the tendinous strips of the bilateral anterior b ellies. This m uscle is better form ed in older ind ividu als (fig. 2 — d.). The m u scle described covers from th e bottom th e posterior deep part of m. m y lo h y o id e u s lyin g under it. The d ivision of m. b iv e n te r into anterior and posterior b ellies noted both in dom estic cattle (B i j v o e t, 1908; M a r t i n & S c h a u d e r , 1938) and in sm all dom estic ( T h i e l , 1954) and w ild rum inants ( T h i e l , I.e.; H a m p l & K a m a n , 1959) is also clearly defined in the E. bison. The shape and situ ation of both b ellies correspond to relations in cattle, the insertion of the anterior b elly on the m andible reaching fu rther to the front, sim ilarly to th e relations in the goat ( T h i e l , I.e.). 2.

NECK AND TRU N K

A. Muscles of the shoulder girdle

1. M. trap eziu s (Fig. 29 — Tu, Th.), form s th e m ost superficial layer of m u scles of th e w ith ers. As in other anim als, the pars cervicalis and pars thoracalis, w hich are not sharply divided from each other, can be distin gu ished in it. T rapeziu s cervicalis, attached to the dorsal border of the fu n icular part of the nuchal ligam ent, exten d s over the space b etw een the second and third cervical verteb rae and th e first thoracic vertebrae, and ends along th e spine of the scapula. A t th e lev el of the third-fourth cervical vertebrae th is m u scle in tertw in es its m uscular fascicles w ith those of m. rhom boideus w h ich lies beneath it. Its anterior border (especially in th e upper section) fu ses w ith m. cleidooccipitalis. The superior insertion of tra peziu s thoracalis reaches to the spinous processes of the term inal thoracic vertebrae (T h10— T h13, or T h 14). The fibres of th e m uscle run in an an terio-ven tral direction to th e upper 2A?

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of spina scapulae. In th is region the m u scle described coalesces w ith m. cutaneus scapulohum eralis, and the low er border w ith m. latissim us dorsi. The p o w erfu lly d eveloped (especially in m ales) w ithers and neck of th e bison are to a great ex ten t the result of th e strong d evelop m en t of the m uscles of this region. Contrary to this principle, m. tra peziu s in E. bison does not exh ib it sp ecially strong developm ent. Both the cervical and th o­ racal portions are form ed from relatively thin, flat patches of m uscle. The ex ten t of th e truncal insertion of the thoracic part is situated m ore to the back than in cattle (Th9— T h i0) and even m ore so than in rum inants w ith strongly develop ed m uscles of the w ith ers such as the Indian w ater-buffalo, in w hich it reaches T 9 (de M o u l i n , 1924). 2. M. om otransversariu s (Fig. 29 — Ot.) is a flat m uscle running from the scapular fascia at th e lev el of the collum scapulae in th e direction of the head. It is covered from th e top by m. trapezius and in th e anterior section by m. brachiocephalicus, and ends on the w ing of atlas connecting by m eans of its aponeurosis w ith m. splenius and m. longissim us atlantis. 3. M. rh om boideus (Fig. 5, 14, 15, 16, 29 — R, R ’.) is strongly developed in the E. bison. It attains great dim ensions, esp ecially in m ales, shaping (together w ith m. splenius) the outline of th is region characteristic of the bison. It d ivid es into tw o parts, differing as to situation and direction in w hich the fibres run. Pars thoracalis is inserted on the lateral surfaces of the spinal processes of the thoracic vertebrae from the 2 — 5. T he flat ven ter of this part has fibres running transversely, upwards and forwards, and ends on the m edial surface of th e scapular cartilage and the scapula. Pars cervicalis has fibres running in an arch upw ards and forwards, and is directed forw ards from the anterior part of the scapular cartilage, and as it approaches the term ination of the cervical insertion it thickens, the thickening in m ales being considerably greater than in fem ales. The part described is inserted along th e funiculus nuchalis, on w hich at th e lev el of ap proxim ately th e 2— 3 cervical vertebrae it ends su dd en ly in m ales and m ore gradually in fem ales. The cervical parts of the rhom boid m uscles of both sides connect along th e funiculus nuchalis. The structure of th e m u scle described in E. bison d iffers considerably from the relations found in cattle. These differences apply both to the degree of d evelop m en t of th e different parts and to th e ex ten t of the insertions. The cervical part which in cattle is represented by th e rela­ tiv ely flat m uscular ven ter, is strongly thickened in the bison and forms th e ridge of the m assive ’’h um p”. The vertebral insertion of pars th o ra­ cal is reaches in cattle to T h7 — Th8 ( M a r t i n & S c h a u d e r , 1938) w h ile

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in th e E. bison in the 8 specim ens prepared by us it reached T h4 or Ths, and in one case T h6. The strongly d eveloped spinal processes of the thoracic vertebrae of th e E. bison cause th e fascicles of m uscle of the pars thoracalis of the rhomboid m uscle to be relatively longer than in cattle, and th e w h ole part to be w ide. T he participation of m. rhom boideus, and strictly speaking its cervical portion, in th e structure of th e ’’h um p ” in the zebu ox w as described by B o a s & P e t e r s o n (1921) and de M o u l i n (1924). The behaviour of both parts of th e m u scle under discussion in the zebu ox differs considerably from w hat w e found in the E. bison, th e result of w hich is th e co m p letely d ifferen t silh ou ette of this anim al. This leads to th e ’’sh ift” in th e zebu ox of th e cervical part backwards to the thoracic part, w hich produces a ’’h um p ” situated at the lev el of 1st to 6th thoracic vertebrae, w h ile in th e E. bison the ’’h um p” is situated above the cervical part of th e v erteb ral colum n. W e did not find in the E. bison th e division, described in respect of th e zebu ox by de M o u 1 i n (I.e.) of th e pars cervicalis of th e rhomboid m u scle into tw o separating layers. 4. M. stern ocleidom astoideus in E. bison, as in cattle, is represented by m. brachiocephalicus and m. sternocephalicus. a) M. brachiocephalicus (Fig. 29, 30 — Br.) is situated on th e lateral su rface of th e neck. It originates on th e hum erus in the crista h um erii region. The fla t m uscular ven ter runs forwards, surrounding th e anterior part of the deep pectoral m uscle and th e cranio-lateral low er end of m. biceps brachii. Near the shoulder joint it is covered from the side and bottom by pars clavicularis m usculi pectoralis superficialis. It coalesces clo sely w ith the latter, from w hich it cannot be separated. M idw ay along th e neck the m uscular ven ter is divided by a tendinous inscription, at first fa in tly d efin ed and later m ore d istin ctly visib le, into tw o parts strongly fu sed w ith each other. It is on ly possible to separate th em in certain places. T he dorsal part described in cattle and other rum inants as m. cleidooccip ita lis term in ates in a flat aponeurosis on the occipital bone, being also in serted in the funicular part of the nuchal ligam ent near the m. rhom boideus. T he ven tral part know n as m. cleidom astoideus, coalesees at its low er border w ith m. stern o m astoid eu s and thrusting under th e parotid, term i­ n ates on the m astoid process o f the tem poral bone. From it a fe w fibres run to th e base of the cranium . T he description given above of m. brachiocephalicus in the E. bison in p rinciple corresponds to th e description of this m uscle in cattle, excep t that in th e literature at our disposal w e now here found a m ention of

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a connection b etw een m. cleidom astoideu s and m. sternom astoideus. Such a connection is, on the other hand, th e rule w ith m. brachiocephalicus in the goat, in w hich th e cranial insertion is situated only on the base of the cranium ( R e i s e r , 1903). b) M. sternocephalicus in th e E. bison, as in other rum inants, is divided into th e superficial and deep parts. Both originate on the sternum . The superficial part (Fig. 5, 7 — St.) corresponding to m. sternom andibularis in cattle runs from m a n u b riu m sterni and is situated in th is region dorsally in relation to the deep part. It runs in the direction of the head and crosses the deep part laterally at the level of the upper x/z of the neck. A t the lev el of angulus m andibu laris the m uscular v en ter passes into tw o tendons. One reaches th e anterior border of m. m asseter, closely fusing w ith it. The second is inserted on the body of the m andible. The deep part corresponding to m. stern om astoideu s (Fig. 5, 7 — Sr.) also runs from m an u briu m sterni. It coalesces in this place w ith m. sternom andibularis and is partially covered by it. The long, flat m uscular venter crosses m. sternom andibularis m edially, and laterally m. om ohyoideus, and at th e lev el of th e atlas passes into aponeurosis. It ends on th e mastoid process of the tem poral bone (join tly w ith m. cleidom astoideus) and partially also on the base of th e cranium and the m edial surface of the ram us of the m andible. The structure of and course follow ed by m. sternocephalicus in the E. bison do not in principle d iffer from the relations found in cattle. 5. M. sternoclavicularis — in th e European bison is very strongly fused w ith m. brachiocephalicus. It exten d s from m anu briu m sterni and from th e first costal cartilage, passing upw ards and sidew ays, to disappear in m. brachiocephalicus. The d egree of fusion of th e above m uscles is very considerable. A m ong the E. bison w hich w e prepared m. sternoclavicularis and m. brachiocephalicus w ere d istin ctly separated from each other only in the cadaver of one m ale called ’’P lu szcz”. 6. M. latissim us dorsi (Fig. 16, 30 — Ld.) form s a w ide m uscular layer situated caudally in relation to m. trapezius. P artially covered by m. cutaneus trunci, it originates in a w ide, flat aponeurosis closely fused with the superficial layer of the lum bo-dorsal fascia and w ith the dorso-transversal fascia. Its aponeurosis passes into a flat m uscular ven ter attached in addition in the form of tw o teeth on the low er V3 of ribs X — XI or IX — X depending on th e num ber of ribs occurring. The venter, in w hich th e fibres run convergently, in a cranio-ventral direction, partialy covers the serratus m uscles, and thickens as it approaches the posterior border of the scapula. A bove the olecranon latissim us dorsi thrusts under caput longum of the triceps brachii. On the

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m edial side of the lim b the ven ter passes into a strong tendon partially fused w ith m. teres major. It term inates on th e deltoid tuberosity of the humerus. The low er border of the m uscle occasionally intertw ines the fascicles of fibres w ith m. pectoralis profundus. T he structure of m. latissim us dorsi exh ib its relations b etw een do­ m estic cattle and sm all rum inants. The sim ilarity to th ese latter is marked in th e costal insertion of this m uscle, w hich in sheep and goats covers ribs IX— XI (M o r i k o, 1954), w h ile in cattle it reaches ribs X I— XII (M o r ik e, 1954; M a r t i n & S c h a u d e r , 1938). On th e other hand, how ever, th ese authors have em phasised the occurrence in sm all rum inants of 3— 4 digitations w hich w ere attached to the ribs, w h ile in the E. bison, sim ilarly to cattle, w e found the presence of tw o digitations. K o m a r e k (1958) describes the intertw inin g of fibres b etw een m. latissim us dorsi and m. pectoralis profundus in C ervu s elaphus ( L i n n a e u s 1758). 7. M. m. pectorales form in the E. bison tw o principle aggregations in the form of m. pectoralis superficialis and m. pectoralis profundus. a) M. pectoralis superficialis (Fig. 29, 30 — Ps.) covers th e sternum from the anterior, dividing into tw o distinct parts. The anterior and m ore superficial part, know n under th e nam e of pars clacivularis, runs from th e lateral surface of m a n u briu m sterni. The flat venter, w ith fibres running in an cranio-lateral direction, covers the low er end of m. biceps brachii, term inating on the hum erus in the region of the radial fossa and in the antebrachial fascia. The posterior part of this m uscle pars sterno-costalis originates on m a ­ n ubriu m sterni (is covered by the anterior part) and on the sides of sternebrae and reaches cau d ally to th e lev el of 5— 6 costal cartilages. The flat ven ter has fibres running in an anterio-lateral direction, and it ends in fascia antebrachii, connecting w ith m. brachicephalicus. b) M. pectoralis profundus (Fig. 29, 30 — Pp.) is strongly developed in th e E. bison and begins on the stern u m and costal cartilages (with the excep tion of the first), up to the abdom inal fascia. The stronger (compared w ith m. pectoralis superficialis) m uscular v en ­ ter, the fibres of w hich run ob liq u ely craniad, bifurcates at its term ina­ tion. The anterior part of th is attachm ent inserts on tu b e rcu lu m m aius of th e hum erus and cranio-m edially covers th e lateral branch of m. supraspinatus. The posterior part ends under the m edial branch of the above m uscle. The prescapular part of th e deep pectoral m uscles is not present in the E. bison as is the case in other rum inants. We found a few fascicles of m uscle fibre running forw ards from tu be rcu lu m m aius in the direction of

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the cervical border of th e scapula in one out of six ind ividu als dissected (’’P lu szcz”). It m ay be considered as corresponding to th e prescapular part of the deep pectoral m uscle. The presence of such fascicles has also b een described in som e dom estic cattle (M o r i k e, 1954). The pectoral m uscles behave sim ilarly in th e E. bison to those in dom es­ tic and certain w ild rum inants, such as, for instance, C ervu s elaphus, C apreolus capreolus ( R e i s e r , 1903; K o m a r e k, 1958), the gazelle (M or i k e, 1954). 8. M. serratus ve n tra lis in th e E. bison (Fig. 5, 29, 30 — Sv, S v ’.) is strongly d eveloped and originate on th e m edial surface of the scapular cartilage and the scapula. Serratus thoracis falls into regular digitations, spreading out fan w ise to the respective ribs. The first three digitations are attached to the upper V2 of th e ribs I, II, III. Starting from rib IV caudad, the digitations are longer and reach to th e costo-chondral junctions. The final digitation is attached to rib IX. In the m ajority of th e individuals dissected there w as a further sm all digitation attached above the final digitation on rib IX , w hich in three specim ens (’’Tatra”, ’’P lastu s”, and foetus) reached to rib X. Serratus cervicis ends in digitations on the transverse processes from th e 3rd to 7th cervical vertebrae. The division and ex ten t of the attachm ents of m. serratus ven tra lis in the E. bison correspond to the relations found in cattle. B. Epaxial m uscles

1. M. splenius (Fig. 5, 6, 29 — SI.) originates on the spinous processes, from the 2nd to 5th thoracic vertebrae and in fascia spinotransversaria. The m uscular fibres run ob liquely forw ards and dow nw ards, thickening considerably at the lev el of the 1st thoracic vertebra. In th e cervical part the m uscle becom es even thicker, this thickening being specially strong in adult m ales, in w hich th e division of the m uscle into three flesh y digitations can be seen. In fem ales and young specim ens th e division of th e b elly of th e m uscle is less distinct, only three tendinous layers being visible. The m ost caudal of th ese connects w ith the digitation of m. serra­ tus ventralis, and term inates on th e 3rd cervical vertebra. The second in order of the bands runs to th e w in g of atlas in com m on w ith the tendons of longissimus atlantis and longus atlantis m uscles. The m ost cranial band, in the form of a flat aponeurosis, is located together w ith the insertion of the m. longissimus capitis on the occipital bone. This aponeurosis is in com mon w ith the pars occipitalis m usculi longissimi capitis, w ith pars cleidooccipitalis m usculi brachiocephalici and w ith m. splenius.

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T he attachm ent of th e above m uscle to th e spinous processes of the thoracic verteb rae reaches further than in the case of cattle, in w hich it covers the spinous processes of Th3 and T h 4 vertebrae. This attachm ent in addition projects further craniad than that in cattle. A ttach m en t of m. splenius on the spinous processes of th e first thoracic vertebrae occurs also in such B ovidae as the Indian w ater buffalo and zebu ox (de M o u 1 i n, 1924), but is lim ited to the first three vertebrae, w h ile in th e E. bison it reaches T h5. T his ex ten siv e reach of th e thoracic origin of th e splenius is characteris­ tic on ly of sheep and goats (M a r t i n & S c h a u d e r , 1938). 2. M. ilocostalis (Fig. 5, 6, 7, 10, 29 —- II.) in its lum bar part above the final lum bar vertebrae, is strongly coalesced w ith th e m. longissim us dorsi, below w hich it lies. Cranially from the 3rd lum bar vertebra the boundary b etw een both m u scles starts to be d istin ctly perceptible. The m ost cranial set of fasciculi of the lum bar part of the m uscle in question inserts on the fin al rib. A bove th e thoracic part of the vertebral colum n m. iliocostalis is situated la terally in relation to m. longissimus and passes to the lateral side of the ribs. It is com posed of a series of m uscle bundles w hich insert on the posterior border of the vertebral extrem ities of th e ribs. Each of the bundles crosses tw o ribs, being attached to th e third. This ap plies in the caudal part of th e m uscle (to rib IX or X), w h ile in the cranial part the m u scle bundles cross three ribs, being attached to th e fourth. Cranially from rib III m. iliocostalis coalesces w ith m. longissimus dorsi, w hich is ex trem ely thin in this place, and th ese tw o m uscles insert together on the transverse process of the 1st thoracic vertebra. In one of th e six specim ens dissected by us (’’P lu szcz”) this m ost cranial insertion w as situated on the transverse processes of th e 7th cervical vertebra and w ould correspond to pars cervicalis of th e m. iliocostalis in cattle. In the rem aining fiv e spe­ cim ens of E. bison the pars cervicalis of th e m. iliocostalis w as not found. 3. M. longissimus from th e d escriptive aspect m ay be divided into the lum bo-dorsal and cervical parts. The latter is divided into three separate m u scle units, i.e.: longissim us cervicis, longissimus atlantis and longissi­ m us capitis. a) M. longissimus lu m b o ru m et dorsi (Fig. 6, 7, 8 — Li.) is a m assive m u scle unit, exten din g along the dorsal side of the spine. In th e posterior part it blends on the m edial side w ith m. spinalis and w ith m. iliocostalis on th e ven tro-lateral side. It originates on crista iliaca, on the lateral su rfaces of th e spinous processes of the tw o first sacral vertebrae, the spinous processes of the lum bar vertebrae and on th e sam e pro­ cesses of th e tw o last thoracic vertebrae. In the lum bar section it is also attached on th e transverse processes and m am m illary processes. A t the

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lev el of the 13th thoracic vertebrae, on the dorso-m edial border of the m uscle described, th e division b etw een m. longissimus and m. spinalis becom s apparent. The b elly of the longissimus dorsi sends insertion s d ow n ­ wards to th e transverse processes of thoracic vertebrae and to th e verte­ bral ex trem ities of the ribs. A t th e lev el of the 3rd thoracic vertebra the m uscle becom es very th in and fin ally disappears, inserting m ost cranially on the transverse process of the 7th cervical or 1st thoracic vertebra. In this part m. longissimus dorsi coalesces w ith the anterior part of m. iliocostalis. The location and insertions of m. longissimus dorsi in th e European bison are sim ilar to the corresponding ones in cattle. O nly the cranial attachm ent, w hich in cattle reaches to the transverse processes of th e 7th cervical vertebra, reached to th e first thoracic vertebra in three of the four E. bison w hich w e dissected. In one (’’P lu szcz”) its behaviour w as sim ilar to that in cattle. b) M. longissimus cervicis (Fig. 7, 8 — Lr.) is a craniad exten sion of m. longissimus dorsi. It originates on the transverse processes of the first six or seven thoracic vertebrae, under the respective bundles of longissimus dorsi. It is covered in this section by its venter, and in addition is attached on fascia spino-transversalis, passing above th e cervical part of th e spine and ending on the transverse and articular processes from th e 3rd to 7th cervical vertebra. A tendinous band passes from the anterior edge of the venter h a lfw a y along its length, connecting this m uscle w ith m. longissi­ m us capitis. The m ost caudal digitation of m. splenius is connected slig h tly below the place w here it joins w ith the perim ysium of th is m uscle. M. longissimus cervicis in the E. bison behaves sim ilarly to that in cattle. c) M. longissimus capitis (Fig. 7, 8 — Lp.) originates on the articular processes of the cervical vertebrae from the 2nd to the 7th, and occasion­ ally even from the 1st thoracic vertebra (P luvius II. Plater). T hese insert­ ions are situated laterally in relation to the corresponding insertions of m. sem ispinalis capitis. The flat m uscle ven ter covers the anterior part of m. sem ispinalis capitis (in m ales only to a sligh t degree) and passes into the tendon, w hich term inates on pars lateralis of the occipital bone and on the tem poral bone, join tly w ith the cranial digitation of splenius. d) M. longissimus atlantis (Fig. 6, 7, 8 — La.) originates on the articular processes of the cervical vertebrae from the third to the sixth (w ith P lu ­ vius II also on the 7th cervical and first thoracic vertebra). The thin ven ter situated laterally in relation to m. longissim us capitis term inates on ala atlantis jo in tly w ith m. splenius,

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The m uscles longissim us capitis and longissimus atlantis are u sually described in ca ttle join tly as one m uscle. In th e E. bison w e dissected th ey w ere clearly separated, both in the anterior part and in th e v icin ity of th e vertebral insertions. The ex ten t of th e insertion on th e articular pro­ cesses of th e cervical vertebrae is greater in th e E. bison and sh ifted more caudally in relation to that described in cattle, since it exten d s in th e E. bison in m. longissim us capitis from the 3rd to the 7th cervical and even in som e cases to th e 1st thoracic vertebra, w h ile in cattle its e x ten t is li­ m ited to b etw een th e 4th and 6th cervical vertebrae ( M a r t i n & S c h a u d e r , 1938). This also applies to m. longissimus atlantis. This sh ift in a caudal direction of the vertebral insertion of m. longissi­ m us capitis occurs am ong the B ovidae in the Indian w ater b uffalo and the zebu ox (de M o u l i n , 1924), but this insertion does not reach to th e th o­ racic section, as w as th e case in tw o of the fiv e E. bison w hich w e dissected. T he rela tiv ely w id e ex ten t of th e insertion of m. longissimus capitis and m. longissimus atlantis occurs in the sheep and goat from 2nd thoracic to th e 2nd cervical vertebra — ( R e i s e r , 1903) and in C ervu s elaphus ( K a m a n & H e m p l , 1958). 4. M. spinalis et m. sem ispinalis in th e E. bison are, like those in other rum inants, strongly fused in th e lum bo-thoracic section. In th e cervical part th ey form a partly separate m uscle unit, described further as m. spi­ nalis cervicis et m. sem ispinalis capitis. a) M. spinalis et sem ispinalis dorsi (Fig. 6, 7, 8 — Sd, S d ’.) form s a group of m uscle bundles participating in th e com position of funiculus m edialis (according to the term inology introduced by B o g o r o d z k y & S t i m p e 1). Two d istin ctly separated parts can be distinguished in this m uscle: th e lateral and m edial. T he lateral part originates on crista iliaca and tuber sacrale of th e ilium and on the spinal processes of th e last three or four lum bar verteb rae and th e tw o first sacral vertebrae. In the lum bar region th e lateral part described is located m edially from m. longissimus and is blended w ith it. A t th e lev el of th e 13th thoracic vertebra m. longissimus and m. spinalis separate from each other. M. spinalis sends out several distinct m uscle d igitations w hich term inate on th e spinal processes from 10th thoracic to the 7th cervical vertebra (Foetus cf, P latyn a, Plazm a) or on the spinous processes from 10th to 1st thoracic vertebra (Pluszcz). The m edial part originates m ore cranially than that of the previous one and form s three or four tendinous m uscle bands. In the lum bar section its upper edge connects w ith the lateral part and th ey then join tly insert on th e apices of the spinous processes of the three first lum bar vertebrae and on th e lateral surfaces of th e spinous processes of the th ree final thoracic vertebrae. Cranially at th e lev el of 4th— 5th thoracic vertebrae

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the above-m entioned bands lose their tendinous character and in tertw in e m uscle bundles w ith the lateral part under w hich th ey lie. T he division betw een th e tw o parts of m. spinalis becom es indistinct in this region and the m uscle bundles of both parts term inate join tly on the spinous pro­ cesses, in the sam e w ay as that described in the lateral part. The m uscle bundles corresponding to m. sem ispinalis lu m boru m (Fig. 9 — So.) are v isib le in the lum bar section. T hey take th e form of three tendinous bands w hich pass from the m am m illo-articular processes of the two first lum bar vertebrae and the final thoracic vertebra. T hese bands blend w ith the m edial part of m. spinalis dorsi described above. The general pattern of the structure of m. spinalis et sem ispinalis dorsi in the E. bison is sim ilar to the structure of this m uscle in other rum inants. The ex ten t of th e insertions in th e caudal part is further sh ifted caudad in relation to that described in cattle, as it exten d s over the spinous pro­ cesses of th e tw o first sacral vertebrae, w h ile in cattle it is lim ited to the spinous processes of th e lum bar part of th e vertebral colum n (S t i m p e 1, 1934). This sh ift caudally of th e attachm ent of the lateral part of this m uscle is, according to S t i m p e 1 (I.e.) typ ical of sm all rum inants. In the European bison, in com parison w ith cattle, the origins of the bands corresponding to m. sem ispinalis lu m b oru m are also sh ifted sligh tly backwards. In cattle it is only in a few cases that th ey reach th e m am ­ m illo-articular processes of the first lum bar vertebra, and in the m ajority th ey are confined to th ese processes of the thoracic vertebrae (S t i m p e 1, I.e.). In all th e E. bison w hich w e dissected the m uscular bands of m. s e ­ mispinalis lu m bo ru m ran from the tw o first lum bar vertebrae and the final thoracic vertebra. b) M. spinalis cervicis (Fig. 8, 9 — Sc.) is a strong m uscle originating on the spinous processes of the 7 th cervical vertebra and the first tw o or three thoracic vertebrae (P latyna, Plazm a, foetu s c f , Pluszcz). The flat venter, th e fibres of w hich run cranio-ventrally, term inates on th e spinous processes of the 3rd to 6th cervical vertebrae. The segm ental structure is esp ecially distinct in adult m ales (Pluszcz), w h ile in fem ales and young individuals (Plazm a, P latyna, foetu s cf) the division into segm en ts is less clear. c) M. sem ispinalis capitis (Fig. 6, 7, 8 — Sm.) lies under m. splenius and is th e third m uscle in turn (in addition to m. rhom boideus and m. splenius) w hich participates in the form ation of the characteristic shape of the E. bison. In th e thoracal section it originates on the transverse processes of the thoracic vertebrae. Of the nine E. bison w hich w e dissected, in one th e origin covered Thi — Th7, in six Thi — Th8 and in tw o Thi — T h9. The d ifferen ces in th e ex ten t of the thoracic origins of m. sem ispinalis capitis w ere not connected w ith the sex of th e anim al dissected. In this

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section the m uscle described is partly covered by m. longissimus dorsi and form s a flat venter w ith fibres running cranio-dorsally. In th e cervical section the ven ter of m. sem ispinalis capitis thickens (especially in m ales) and has several origins on the articular processes of the 2nd — 7th cervical vertebrae. The m uscle bundles have fibres running in a more perpendicular direction (in relation to th e thoracic section) and form a w ide ven ter (in m ales it is thinner, and in fem ales thicker than m. splenius), covering the lam ellar part of the nuchal ligam ent from the sides. A bove the axis th e ven ter passes into an aponeurosis, term inating laterally from the insertion of the nuchal ligam ent. M. sem ispinalis capitis in th e E. bison corresponds as regards the ex ten t of its origins and insertions to the relations found in cattle. It is, how ever, m ore strongly developed and d istin ctly separated from m. spinalis et sem ispinalis dorsi, w h ile in cattle it form s an undivided exten sion of th ese m uscles on the neck region. 5. M. m u ltifidu s (Fig. 8, 9, 10 — Md, Me.) is com posed of a series of m uscle bundles running b etw een th e articular or m am m iilo-articular processes of the one, and th e spinous processes of the preceding vertebra. This form ation begins above the sacral part of the vertebral colum n and in this part, as in the lum bar part, the flesh y fibres run obliquely (cranio-dorsally), and the m uscle bands term inate on the lateral surface of the spinous process of the previous vertebra, or on one vertebra further for th e w h ole of its height. In the thoracic part the m u scle bundles of the m uscle described run in a more perpendicular direction, do not com pletely cover the lateral surfaces of the spinous processes and do not cover m ore than three vertebrae. In the cervical part the d ivision into m uscle bundles is less distinct. T hey insert on the articular processes of the cervical vertebrae and term inate at the base of th e spinous processes, form ing a uniform flat m uscle ven ter covering from th e sides th e anterior part of m. spinalis cervicis, and are th em selves covered by m. longissimus capitis. The m ost cranial insertion of this m uscle is attached on the caudal border of the spinous process of th e axis. Our observations of m. m u ltifid u s in th e E. bison agree w ith the description of this m uscle by K r ii g e r (1927) in cattle and goats, excep t for th e lum bar part of this m uscle. In th is part, according to the above author the shortest bundles of fibres u su ally cross one vertebra, w h ile in th e E. bison w e found bundles running from vertebra to vertebra. It is only the question of the occurrence in th e E. bison of m. rotatores w hich is not clear, th ese m uscles according to K r u g e r (I.e.) being the m ost d istin ctly separated from m, m u ltifid u s in rum inants. This separation w as

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not found in the E. bison w hich w e dissected. This problem , and also th e one dealt w ith by K r ii g e r (I.e.), using dom estic rum inants as exam ples, as to the form ation of th e articular surfaces of verteb rae and the m utual dependence of th ese tw o factors, require in m y opinion fu rth er and m ore detailed exam ination. C. Group of the short m uscles of the vertebral column

1) M. obliquus capitis cranialis (Fig. 9, 10 — Or.) in th e E. bison is w eaker in relation to m. obliquus capitis caudalis. It runs from th e cranial border of the w ing of the atlas and from the fossa atlantis and inserts on th e occipital bone, along sutura parieto-occipitalis, to the base of the param astoid process. 2) M. obliquus capitis caudalis (Fig. 9, 10 — Ou.) in th e E. bison is a strongly developed m uscle, originating on th e lateral surface of th e spinous process of th e axis. The thick, triangular ven ter (the base of w hich is directed tow ards the atlas) inserts on th e caudal border o f the w ings of atlas and cranially of this border. 3) M. rectus capitis dorsalis m ajo r (Fig. 9, 10 — Rj.). Its origin is located on the edge of the spinous process of th e axis. T he ven ter of this m uscle ex h ib its tendencies to division into a superficial and a deep part. The latter on account of its ex ten t corresponds to m. rectu s capitis dorsalis in term ed iu s described by certain authors (K o 1 d a, 1950). In th e E. bison w e dissected th is division w as not su fficien tly distinct to authorise d ifferen tiation into tw o independent m uscle units. Both parts term inate on the cranium under the tendon of th e sem ispinalis capitis w ith w hich it som etim es blends. 4. M. rectus capitis dorsalis m inor — passes from tu b e rcu lu m dorsale atlantis. The m uscle venter, situated under the b ellies of th e previously described m uscle, is stronger and better developed, but shorter than it. and term inates on the skull below its insertion. 5. M. atlantooccipitalis (Fig. 7, 8, 9, 10 — Ao.) described in the sheep and goat ( K r u g e r , 1927) and, am ong w ild rum inants, in Capreolus capreolus and C ervu s elaphus ( R e i s e r , 1903), also occurs in the E. bison. It is located in the exten sion of the insertion of th e m. longus atlantis. It originates on the cranial tendons of the m. longus atlantis and m. longissim us atlantis on the atlas. The flat ven ter of m. atlantooccipitalis runs craniad ending in a flat aponeurosis on th e tem poral bone. 6. M. m. intertran sversarii — in the E. bison, as in other m am m als, can be distinguished only in the cervical and lum bar parts, the m uscle bands of th e lum'bar part being com pletely fused w ith m, longissimus dorsi and m. iliocostalis.

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M. m. intertran sversarii in the cervical section are form ed from m uscle bands running b etw een the transverse processes and articular processes of th is section. T hree groups can be distinguished am ong them , d ifferin g both as to the direction in w hich th ey run and as to their insertions. T he first group (Fig. 9 — It.) form bundles running from the transverse processes of the one vertebra to th e articular processes of the next. The second group (Fig. 9 — It’) is form ed by th e bundles running b etw een the transverse processes of tw o neighbouring vertebrae. The third group (Fig. 9 — It”), th e m ost strongly developed, is form ed by bundles running from th e transverse processes of one vertebra to the ven tral edge of th e ven tral branches of the transverse processes of the n ex t vertebrae. In .th is group th e m ost strongly d eveloped are the bundles running b etw een th e third, fourth and fifth cervical vertebrae. D. The lateral and ventral m uscles of the vertebral column

1. Musculi scaleni in the E. bison, as in cattle, are represented by tw o m uscles, the m. scalenus supracostalis and m. scalenus prim ae costae. M. scalenus supracostalis (Fig. 5, 6, 7, 8 — Ss.) is a flat m uscle w hich is located on the lateral side of th e thoracic w all, under the digitations of th e serratus tharocis. It originates on th e posterior border of the third rib in its low er part and by m eans of a short aponeurosis on the anterior border of the fourth rib in th e upper part. The flat venter, the fibres of w h ich run in th e form of an arch, passes into a tendon w hich is attached on th e transverse process of th e 5th cervical vertebra. M. scalenus prim a e costae d ivid es in turn into tw o separate parts. Pars ve n tra lis (Fig. 5, 6, 7, 8 — Spv.) begins on the sternal half of the first rib. The thin ven ter of th e m uscle ends in an indistinct tendon on the lateral branch of th e transverse process of the 6th cervical vertebra, and then together w ith m. scalenus supracostalis (beneath it) passes on to th e transverse process of th e 5th cervical vertebra. Pars ventralis (Fig. 5, 6, 7, 8 — Spv) begins on the sternal half of the first rib in the form of a w ide, flat ven ter and ends in distinct digitations under the lateral branches of th e transverse processes of the 5th to 7th cervical vertebrae. D espite the joint pattern of structure, th e musculi scaleni in the E. bison exhibit, in com parison w ith th e corresponding m uscles in cattle, d ifferen ­ ces in th e ex ten t of the insertions. M. scalenus supracostalis inserts on the transverse process of the 5th, and not 6th to 3rd cervical vertebrae, as is the case w ith cattle. Insertions of th is kind of th e m uscle discussed in E. bison are m ore lik e the relations found in the goat, in w hich m. scalenus supracostalis inserts on th e 5th— 4th cervical vertebrae ( R e i s e r , 1903).

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The dorsal part of the scalenus prim ae costae reaches in th e E. bison to the transverse processes of th e 5th and 6th cervical vertebrae, w h ile in cattle it reaches to 7th cervical vertebra. The ventral part of this m uscle does not d iffer as regards insertions from that found in cattle. 2. M. divided ventral both as

longus colli (Fig. 11) in the E. bison, as in other m am m als, is into the thoracic part and cervical part. Both are located on the surface of corresponding parts of th e spinal colum n and differ to location and to direction in w hich th e fibres run.

The thoracic part (Fig. 11 — b) is form ed from m uscle bundles located in pairs (on th e left and right sides) of w hich the m ost caudal pair runs from the ventral crest and lateral surfaces of the body of the 6th thoracic vertebra. The fusiform ven ter is form ed from the n e x t pairs of m uscle bundles running sim ilarly from the fourth and second thoracic vertebrae, and passes cranial inserting on th e w ay into th e lateral surfaces of the bodies of the thoracic vertebrae, and fin ally inserts w ith m igh ty tendons on the ven tral branches of the transverse processes of the 6th cervical vertebra. The cervical part (Fgi. 11 — a) has tw o layers. The m uscle bundles originate along th e low er border of the ventral branches of the transverse processes of the 6th cervical vertebra form the superficial layer, which ends on the crista corporis of the 4th cervical vertebra and the deep layer ending in the sam e place on the 5th cervical vertebra. The bundles originating on th e 5th, 4th and 3rd cervical vertebrae behave sim ilarly, that is the superficial layer covers one vertebra, and the deep one ends on the previous vertebra. The m uscle bundles originating on the transverse processes of the axis form distinct b ellies (left and right), w hich are attached on the ventral tubercle and on the posterior border of the ventral arch of atlas. The description of m. longus colli given above in the E. bison agrees w ith the description of this m uscle in cattle. 3. M. longus capitis (Fig. 6, 9 — Lk.) in th e E. bison, and in particular in the m ales, is strongly developed and is located on the lateral side of the neck. The m uscle in question passes from the ventral branches of the transverse processes of the cervical vertebrae, from the 3rd to the 6th (below th e m. longus atlantis, w hich is also attached in this place). The long venter, w hich in its anterior end connects w ith m. sternom astoides et cleid om a sto ideu s, term inates together w ith them on the tem poral bone and on th e base of th e skull. 4. M. longus atlantis (Fig. 5, 7, 8, 9 — Lt.) passes from the ventral branches of th e transverse processes of the 2nd to the 6th cervical verte­ brae (in the case of ’’T atra” to the 5th) under the corresponding insertions

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of m. longus capitis. Stronger in relation to th e latter, th e m uscle venter ex h ib its tendencies to bifurcation and term inates on the edge of the wings of atlas. T he behaviour of m. longus capitis and m. longus atlantis in the E, bison is sim ilar to that in cattle, excep t that th ey are m ore strongly separated from each other, w h ile in cattle this separation is not so distinct. As a result certain authors have described the tw o m uscles as one m uscle unit. 5. M. rectus capitis lateralis (Fig. 7, 8, 9 — Rl.) form s a strong short ven ter, originating in th e fossa atlantis (in the depression located caudo-v en tra lly to foram en alare and term inates on the m edial surface and caudal border of th e param astoid process of th e occipital bone. 6. M. rectu s capitis ve n tra lis originates on the caudal border of the ven tral arch of the atlas (laterally from th e ventral tubercle). The large, thick m uscle vfenter term inates on th e bony prom inences of the basilar part of the occipital bone. 7. M. stern ohyoideus (Fig. 2 —J.) originates on m an u briu m sterni jointly w ith m. stern oth yreoideu s, partly above and below the insertion of m. sternom astoideus. Crossing the latter, im m ediately after leaving the insertion it passes to th e ventral side of the trachea, ending on the body of th e hyoid bone. 8. M. stern o th yro id e u s originates togeth er w ith the form er one, in itially coalescing w ith it. In its continuation it runs laterally in relation to m. stern ohyoideus, term inating on the thyroid cartilage of the larynx. M. m. stern oh yo ideu s and stern o th yre o id eu s in their insertions and course do not exh ib it d ifferen ces from the sam e m uscles in cattle. 9. M. om ohyoideu s (Fig. 2, 5, 6, 8 — Om.) in the E. bison, as in other rum inants, passes from fascia colli profunda at th e lev el of the transverse processes of th e third and fourth cervical vertebrae. The flat and relativ­ ely w ide m uscle ven ter, th e fibres of w hich run cranio-ventrally, ends on the hyoid bone. On its w a y the ven ter connects w ith m. sternom astoideus E. The m uscles of the thoracic w all

1. M. serratus dorsalis in the E. bison is divided into m. serratus dorsa­ lis cranialis and m. serra tu s dorsalis caudalis. a) M. serratu s dorsalis cranialis (Fig. 6 — Si.) originates on fascia spinotransversalis at the lev el o f the fourth or fifth thoracic vertebra, form ing a fla t ven ter w ith fibres running caudo-ventrally. The venter, by m eans of fa in tly p erceptible d igitations, inserts on the anterior borders from the 6th to 8th ribs.

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b) M. serratus dorsalis caudalis (Fig. 29 — Se.), in th e foetu s and th e 6-m onths-old ’’P la ty n a ” w hich w e dissected, form ed a uniform flat m u scle venter, passing from fascia lum bo-dorsalis at th e lev el of th e fin al thoracic vertebra and first lum bar vertebra and term inating on th e caudal border of the tw o last ’’P la ty n a ”) or th e three last (foetus) ribs. In th e rem aining m ales (Pluszcz, Punkt) th is m uscle form s three separate digitations inserted on the caudal borders of the three final ribs. Each of the d igitat­ ions divides into a superficial part covering m. intercostalis extern u s, and a deep part, penetrating under it and term inating low er in relation to th e superficial part, join tly w ith m. intercostalis internus, w ith w hich it fuses. M. serratus dorsalis in m am m als exh ib its great sp ecific and individual variation. This variation is m anifested by d ifferen ces in the ex ten t of th e costal insertions of both parts of this m uscle, or even the retrogression of one of them . M. serratus dorsalis cranialis s. pars inspiratoria is form ed in cattle from three to six digitations, having costal insertions from th e 4th, 5th or 6th to 8th or 9th rib (Ma r t i n & S c h a u d e r , 1938). In sheep the costal insertion is situated on the anterior border of the 4th, 5th or 6th rib (R e is e r, 1903), in goats from the 4th to 6th, or 7th rib ( R e i s e r , I.e.). In the C ervu s elaphus th e exten t of the costal insertions of th e serratus inspiratorius covers ribs 5, 6, 7 or 8 (R e i s e r, I.e.). In th e Indian w ater buffalo and the zebu ox this part is strongly reduced or does not occur at all (de M o u l i n , 1924). M. serratus dorsalis caudalis s. exp ira to riu s , is form ed in cattle from 4 to 5 m uscle digitations, term inating on the caudal borders of the 10th or 11th to th e 13th ribs (M a r t i n & S c h a u d e r , 1938), in sheep and goats from th e 9th, or 10th to 12th, or 13th ( R e i s e r , 1903), in C ervu s elaphus from the 10th to 13th rib ( R e i s e r , I.e.). In the Indian w ater buffalo pars expiratoria m usculi serrati dorsalis inserts by m eans of d igit­ ations from the 6th to 13th rib (de M o u l i n , 1924). From th e description given above it is clear that m. serratus dorsalis is characterised by individual variation in the E. bison also. The ex ten t of the costal insertions is d ifferen t from that in the other rum inants consider­ ed for purposes of com parison. 2. M. transversu s costarum (Fig. 5, 7 — Tc.) in the E. bison is located on the lateral surface of the thorax in th e form of a w ide and relatively thick venter, w hich reaches from the sternal extrem ity and costal cartilage of the first rib to the 4th costal cartilage. The ex ten t of this m uscle in the E. bison is shorter than in cattle, in w hich it reaches to the 6— 7th costal cartilage ( M a r t i n & S c h a u d e r , 1938), and corresponds to the relations found in sm all w ild and dom estic rum inants, in w hich it reaches the 4th costal cartilage ( R e i s e r , 1903).

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3. M. m. levato res costarum (Fig. 8, 9 — Lc.) consist of a group of m u scle bundles running from the costal tubercles and transverse processes o f th e thoracic vertebrae to the anterior border of the n ext rib. T hey occur in th e E. bison in all th e intercostal spaces, being less strongly d eveloped b etw een th e 1st and 9th rib, and caudally separating more d istin ctly from the intercostal m uscles. 4. M. m. intercostales ex tern i et interni behave sim ilarly to th e corres­ ponding m u scles in cattle. M. m. intercostales e x tern i are form ed of strongly tendinous m uscle bundles w ith fibres running cau do-ventrally. In the final th ree or four intercostal spaces the direction in w hich the fibres run becom es more horizontal in relation to the fibres in the anterior intercostal spaces and th ey in tertw in e strongly w ith the internal ones. M. m. intercostales in te r ­ ni have fib res running in th e opposite direction to that of th e extern al ones. 5. M. retractor costae (Fig. 29 — Rc.) runs from the transverse process of the first lum bar vertebra and ends on the posterior border of th e final rib, coalescing by m eans of th e final tendon w ith th e final digitation of the serratus dorsalis caudalis. The upper edge of m. obliquus abdom inis internus thrusts in b etw een th ese m uscles. 6. M. tra nsversu s thoracis (Fig. 13 — Tt.) in th e E. bison form s a sym ­ m etrical accum ulation of m uscle bundles located on th e intern al surface of the sternum and on the costal cartilages. T hese bundles run from the m iddle lin e of the dorsal surface of the sternum beginning from the 2nd sternebra. C audally th ey term inate in turn on the costal cartilages from the 2nd to th e 8th and on the sternal extrem ities of th e corresponding ribs. The tw o fiñal portions attached to the 7th and 8th cartilages are clearly separated. The w h ole corresponds to the structure of m. tra n s v e r ­ sus thoracis in cattle. The m uscle bundles of th e opposite sid es do not connect w ith each other in th e m edial line, and their location and insertion correspond to the ”p aired -h oof” typ e of this m uscle described by T u r k i e w i t s c h (1928). F. The abdominal m uscles

1. M. obliquus abdom inis extern u s (Fig. 30 — Oe.) is included in the group of m uscles form ing th e abdom inal w alls, and its structure in the European bison does not in principle d iffer from th e relations encountered in other rum inants. It originates on the extern al surface of the final 7— 8th ribs in the form of a series of digitations thrusting b etw een the d igitations of the thoracic part of the serratus ventralis, and m ore caud­ a lly , under m. latissim us dorsi. The fla t m uscle form ed by th e above-

K rzysztof Ś w ieży ń sk i

-m entioned digitations, th e fibres of w hich run obliquely, cau do-ventrally, form s a uniform w hole. The upper edge coalesced w ith fascia lum bo-do rsalis reaches caudally to tu b e r coxae. The cranio-ventral part thrusts b etw een m. pectoralis profundus and ra. rectus abdominis. The w h ole passes into a strong aponeurosis coalesced w ith tunica flava abdominis. This aponeurosis w idens considerably caudally and connects w ith the aponeurosis of m. obliquus abdom inis internus surrounding rectu s abd o­ minis from below . In the m edial line the low er edge of this aponeurosis connects w ith th e one like it from the opposite side. The caudal edge is attached along the sh aft of ilium and partly on the pubic bone. 2. M. obliquus abdom inis internu s (Fig. 29 — Oi.) is located under m. obliquus abdom inis ex te r n u s , being included in th e com position of the abdom inal w all. It originates on tu b e r coxae and fascia lumbodorsalis. The flat m uscle, th e fibres of w h ich run fan w ise in a cranio-ventral direction, can be divided into tw o parts for purposes of description. The dorsal part, w ith fibres running more parallel, is attached on the caudal edge of the fin al rib, to th e costo-chondral junction. The upper edge of th is part attached on th e tranverse processes of th e lum bar vertebrae, thrusts b etw een m. serratus dorsalis and m usculus retractor costae and term inates on tu b e r coxae and on lig am en tu m inguinale. The ven tral part, thinner than the previous one, has fibres running m ore ob liq u ely (near th e caudal edge th ey run alm ost perpendicularly). It passes in the form of an arch (at th e lev el of a im aginary line drawn from the costro-chondral junction of the fin al rib to the fold of th e flank) into a aponeurosis w hich connects in linea alba w ith the one like it from th e opposite side. This aponeurosis is strongly coalesced w ith the aponeu­ rosis of m. obliquus abdom inis extern u s and covers m. rectus abdominis from th e exterior. The general pattern of th e structure of m. obliquus abdom inis internus in the E. bison is sim ilar to the structure of this m uscle in cattle. The division of this m uscle into tw o parts, described by M a x i m e n k o (1928) in cattle and other rum inants, takes place in the E. bison also. This d ivis­ ion is m anifested in th e d ifferen ce in the direction in w hich th e fibres run and in th e d ifferen t thickness of the m uscle layer, w h ile th e transverse band of fibrous tissue (inscriptio tendinea) separating th ese parts, w hich is present in cattle, is here absent. 3. M. rectu s abdom inis (Fig. 12, 29 — Ra.) in the E. bison is a flat, w ide m uscle, originating by m eans of a distinct tendon on th e caudal tendon of the m. transversu s costarum and on the costal cartilages beginning from th e 5th caudad, and a d istin ctly separated m edial part on th e ventral surface of the sternum (at the lev el of the junction of th e sternum w ith the costal arch). Both parts connect caudally, m aintaining distinct bound­

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aries on ly in th e places w h ere nerve branches and v essels pass through them . The flat v en ter is divided by fiv e tendinous inscriptions. C audally the m uscle term inates on the pubic bone and on th e prepubic tendon. The location and num ber of tendinous inscriptions in m. rec tu s a b d o m inis in the E. bison correspond to th e structure of this m uscle in cattle. The sternal attachm ent, w hich divid es up and is partly tendinous (on the costal cartilages) and partly m uscular (on the ven tral surface of th e stern ­ um) occurs in sheep and in C ervu s elaphus, but in th ese rum inants the num ber of tendinous inscriptions is seven ( R e i s e r , 1905). 4. M. transversu s abdom inis (Fig. 12 — Ts.) form s th e d eep est m uscular layer of th e abdom inal w alls. It passes in a flat aponeurosis from the transverse processes of th e lum bar vertebrae and from the caudal edge of the fin al osseous rib, and further on attaches on the m edial surface of the costo-chondrial junctions of all th e asternal ribs as far as the fin al sternal rib. The m uscle ven ter passes in an arch (along a h ypothetical line conn­ ecting the xiphoid cartilage w ith the region of the stifle joint) into an aponeurosis covering the dorsal surface of m. rectus abdominis. In the interm edial line it connects w ith th e corresponding one from th e opposite side. The structure of m. transversu s abdom inis in the E. bison is sim ilar to th e structure of th is m uscle in cattle, as far as its insertions ar conc-r erned. It differs, how ever, by reason of th e m ore strongly d evelop ed pars aponeurotica. 3.

T H E T H O R A C IC L IM B

A. M uscles of the shoulder and arm

1. M. deltoideus (Fig. 14, 15, 29, 30 — D, D ’.) in th e E. bison is a flat, broad m uscle w hich inserts along th e spina scapulae dow n to th e acrom ion, and coalesces w ith m. infraspinatus lyin g beneath it, throughout alm ost the entire surface. In th e cranial part it is thinner, and thicker caudally. B eginning from the caudal edge of m. infraspinatus. M. delto id eu s is surrounded by a joint fascial sheath w ith m. teres minor. It m ixes num ero­ us fibres w ith m. teres m inor and also w ith th e lateral head of triceps brachii, w h ich it covers from the sides. It term inates on th e ven tral end of crista hum eri and on th e deltoid tu berosity of hum erus. I n th e caudal part of this m uscle th e part w hich passes by m eans of an aponeurosis on to th e lateral head of tricep s brachii separates from it. The division, occurring in all rum inants, of this m uscle into pars acrom ialis et pars scapularis, w as perceptible in the E. bison w hich w e dissected, but w as not so distinct as that in cattle. 2. M. supraspinatus (Fig. 5, 14, 15, 16, 30 — S.) originates in th e supra­ spinous fossa and on th e anterior an gle and anterior edge of th e scapula.

K rzysztof ¡Sw iezynski

The m u scle venter, protruding cranially slig h tly beyond th e anterior edge of th e scapula, attains its greatest dim ensions at the lev el of tu b e r scapu­ lae. A bove the shoulder joint it divid es into tw o branches of w hich the lateral one, as a rule stronger, is attached w id ely on tu b e rcu lu m maius craniale, and the m edial one on tu be rcu lu m minus craniale. 3. M. infraspinatus (Fig. 5, 15 — I.) occupies th e infraspinous fossa and the greater part of it blends w ith m. deltoideus, w hich covers it. It is attached on the caudal surface of th e spine of scapula in infraspinous fossa and on th e caudal an gle and in its ’’fr ee” part (from m. deltoideus) it form s a thick, fla t ven ter term inated by a strong tendon on facies m usculi infraspinam and by several tendinous fibres on tu bercu lu m m aius caudale. P o l e i n e r ’s statem en t (1932) asserting that contrary to cattle, the tendon of insertion of th is m u scle in the E. Bison w as less tendinous, w as not confirm ed by our observations. 4. M. te res m inor (Fig. 15 — Ti.) is a m uscle alm ost en tirely fused w ith m. infraspinatus and is d ifferen tiated from it on ly by its exten t. T he fibres of its v en ter form a w h ole w hich, lying caudally in relation to th e m ain m ass of m. infraspin atu s, does not term inate on tu b e rcu lu m m aius caudale, but runs slig h tly further to facies teres (although only fain tly perceptible in th e E. bison) and to th e upper section of crista humeri. 5. M. subscapularis (Fig. 16 — Su, S u ’, S u ”.) located on th e m edial side of th e scapula, perm its of d istinguishing three parts clearly separating from each other. The cranial one, in the upper section, coalesces w ith m. su praspin atus and is the th ick est of the three. The m iddle one occupies fossa subscapularis and is strongly tendinous. Its fibres run at an acute an gle in relation to th e cranial part and thrust under it to pass in turn into a joint aponeurosis. The caudal part, lying behind and partly under the m iddle one, coalesces in th e upper section w ith m. teres major. A ll three connect at the lev el of the shoulder joint in a com m on tendon (prev­ iously arranging th em selves one on top of the other in the order given) w hich thrusts under the distal head of coraco-brachialis and inserts on tu b e rcu lu m m inus caudale and below in its vicinity. D uring th e dissection of the thoracic lim bs of tw o fem ales (Poziom ka and Purata), w e encountered a fairly d istin ctly separating fourth part of this m uscle, w ich m ust be considered as corresponding to that part des­ cribed by L e i c h n e r (cit. acc. to M a r t i n & S c h a u d e r , 1938) in m. subscapularis in the goat. 6. M. teres m a jor (Fig. 14, 15, 16 — Ta.) originates on the caudal angle of th e scapula and in th e upper section of the caudal edge, coalescing near its origin w ith m. subscapularis. The flat ven ter thrusts under m. latissim us dorsi, w ith w hich it also in tertw in es fibres and term inates m edially to it on tuberositas te r e s of the hum erus.

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7. M. coracobrachialis (Fig. 16, 18 — Cb.) is a flat m uscle, th e venter of w hich is divided by a branch of N. m usculo-cutaneus into tw o parts: proxim al and distal. Both originate in a distinct tendon on scapular tuber­ osity. The proxim al part is located more su perficially and term inates above teres tuberosity. The distal part in the in itial section is located deeper than the previous one and term inates below teres tuberosity, reaching crista epicon dyli medialis. This division into the d ifferen t parts is visible in older anim als (Poziom ka, Plazm a, Plastus, Pluszcz). It cannot be seen at all in th e calves of E. bison (Poda), apart from the place w here the n erve crosses and from a certain d ifferen ce in th e direction in which the fibres run. 8. M. tensor fasciae antebrachii (Fig. 16, 30 — Tf.) originates on the caudal edge of the scapula and on th e aponeurosis of m. latissim us dorsi. It passes on to the m edial side of caput longum of triceps in th e form of a flat venter, term inating on olecranon and in fascia antebrachii. 9. M. triceps brachii occupies th e angle b etw een the scapula and hum erus, form ing a group of m uscles in w hich four distinct heads can be distinguished. Caput longum (Fig. 14, 15, 16, 29, 30 — Tr.) originates on th e caudal angle of the scapula and below its caudal edge, up to the lev el of the neck of this bone. The triangular m uscle ven ter, w ith fibres running caudo-ven traly, term inates w ith ou t the visib le participation of th e tendinous com ponent on the olecranon, covering from th e m edial side, from the top, and laterally, the tendon of the lateral head. Caput laterale (Fig. 14, 15, 29 — Tr’) originates on the lateral side of the hum eral bone at th e lev el of crista anconea and on the caudal border of tendon of m. teres minor. It is covered in its anterior part by m. deltoideus and in four out of the six E. bison dissected by us fu ses w ith it. In itially thick, th e fla t ven ter n ext term inates on the lateral surface and caudal border of tu b er olecrani. Caput m ediate (Fig. 16 — Tr”) originates above and below teres tube­ rosity. It is covered in its anterior part by m. coracobrachialis, and in the posterior by caput longum. A bove the m edial epicondyle of th e hum erus this head passes into a tendon, term inating on the m edial surface of the processus olecrani. Caput accessorium (Fig. 18 — Tr’”) originates at the lev el of the low er V 3 of th e hum erus on its caudo-m edial side. The flat, thin m uscle venter, w ith fibres running len gth w ays, term inates on the lateral side of processus olecrani under the lateral head of the m uscle described. R. P o 1 e i n e r (1932) in his oth erw ise very detailed description of m. triceps brachii in the Et bison m akes ab solu tely no m ention of th e caput

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accessorium . He does not even refer to its absence w h en discussing d ifferen ces compared w ith cattle, w hich gives rise to the suspicion th at this m uscle unit w as overlooked. In all the specim ens dissected by us th e caput accessorium of m. triceps brachii is relatively strongly form ed. 10. M. anconeus (Fig. 17 — A.) forms a flat m u scle ven ter, located laterally in relation to th e accessory head of triceps brachii, and d irectly contiguous w ith the hum erus. It originates on the lateral condyloid crest and passes on the m edial surface of the lateral epicondyle w h ere it coalesces w ith the tendon of th e lateral head of the triceps. This m uscle is relatively strongly developed in th e E. bison, and corresponds to the relations encountered in sm all dom estic rum inants, but does not possess an independent term inal tendon, as is th e case in Capreolus capreolus and C ervu s elaphus ( R e i s e r , 1903; K o m a r e k, 1958). 11. M. biceps brachii (Fig. 14, 15, 16, 18, 20 — Bb.) originates in a m assive tendon on tu b e r scapulae. It passes into the intertubercular groove b etw een th e branches of m. supraspinatus and exten d s into a fusiform ven ter, running on th e cranio-m edial side of the hum erus. It term inates w ith one tendon on the collateral m edial ligam ent of the elb ow joint, on tu b e rcu lu m ligam entosu m m ediate of radius and on th e radial tuberosity, and w ith a second tendon on the m edial border of the radius. On the m edial surface of m. biceps in old anim als an elongated groove is visible w hich indicates a tendency, as it w ere, to a division of this m uscle into tw o secondary bellies. In th e anim als w e dissected w e did not find the presence of lacertus fibrosus in the form present in cattle. O nly in old individuals (Pluszcz, Poziom ka) does this m u scle exh ib it coalescence w ith the fascia antebrachii w hich is thickened on the m edial side. 12. M. brachialis (Fig. 14, 15, 18 — Be.) originates on the neck of the hum erus in th e caudal h alf of its circum ference, continuing to the beginning of th e anconeal crest. The strong m uscle ven ter is located in th e m usculospiral groove of th e hum erus, in itially covered by the lateral head of th e triceps brachii and by m. deltoideus. On reaching th e lateral side it passes b etw een m. brachiocephalicus and m. ex ten sor carpi radialis, and n ex t runs on to th e m edial side to term inate on th e m edial border of the radius in th e vicin ity of tu be rcu lu m lig am en to su m m ediate. In the ven tral end m. brachialis is covered by a tendon of insertion of m. biceps brachii. A strong tendinous band passes from the insertion to term inate on th e ulna near sp a tiu m interosseum antebrachii. The insertion, described above, of m. brachialis in th e E. bison exhibits interm ediate relations b etw een those encountered in cattle, w here the insertion is located on tuberositas radii, and in sm all dom estic rum inants

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in w hich th is insertion is located on processus coronoideus ulnae ( R e i s e r , 1903). M. brachialis term inates in Capreolus capreolus and C ervus elaph us in a sim ilar w ay to that in th ese latter rum inants ( R e i s e r , I.e., and K o m a r e k, 1958). B. M uscles of the forearm and manus

a. E xtensor division 1. M. e x ten so r carpi radialis (Fig. 14, 15, 16, 17, 18, 20, 29, 30 — Er) originates on th e lateral supracondyloid ridge, on the lateral epicondyle and in the radial fossa. Near the origin it coalesces w ith th e ven ter of m. exten so r digiti tertii proprius. S lig h tly low er, th e ven ter of th is m uscle undergoes d ivision into tw o bellies: the superficial and th e deep, w hich above the carpus pass into tw o d istin ctly separating tendons. The superficial one, running m edially, term inates on tuberosity Me. III. The deep one — lateral — term inates more laterally. In adult individuals a tendinous band passes through the deep venter, beginning above the fossa radialis and joining the lateral tendon. The tendons of insertion are crossed from the front by the tendon of m. abdu ctor pollicis longus. R. P o 1 e i n e r (1932) in his description of this m uscle in th e E. bison refers to th e division of the m u scle ven ter, describing in detail th e course follow ed by both tendons. R e i s e r (1903) also w rites of the division of m. ex ten sor carpi radialis in rum inants and referring to M ii 11 e r & L e i s e r i n g as authority for this, considers it as u sually occurring in sheep, and in certain cases also in cattle. M a r t i n & S c h a u d e r consider th e division of m. ex ten so r carpi radialis as a characteristic of th e m uscle sy stem of sm all rum inants. A ll th e above authors consider this accessory head of m. ex ten so r carpi radialis as corresponding to m. ex ten sor pollicis longus of th e five-fin gered limb. 2. M. e x ten so r carpi ulnaris (Fig. 14, 15, 17, 19, 29, 30 — Eu.) originates on the epicon dylu s extensorius and on crista epicon dyli lateralis, and in one of the E. bison w hich w e dissected (Purata) also on th e m argin of the fossa radialis of the hum erus. The large ven ter runs towards the fingers and above th e carpus passes into a tendon ending in one branch on the accessory bone of carpus, and in the other on Me. IV. According to P o 1 e i n e r (1932) th e tendon of insertion of this m uscle sends out fibres to the rudim entary Me. V, w hich w e noted on ly in one case (Poziom ka). 3. M. a bdu cto r pollicis longus (Fig. 17, 29, 30 — Ap.) originates on the dorso-lateral surface of the forearm bones in a flat, tendinous triangular venter. A bove the carpus it passes into a tendon crossing from th e front

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the tendon of insertion of m. ex ten so r carpi radialis, and term inates on the m edial side of the proxim al extrem ity of Me. III. M a r t i n & S c h a u d e r (1938) and K o 1 d a (1950) in describing th is m uscle in cattle do not distinguish th e ulnar part of origin. The presence of this part of the insertion is characteristic of sheep, goats and C ervu s elaphus ( R e i s e r , 1903). 4. M. ex ten so r digitalis com m unis (Fig. 14, 15, 17, 29, 30 — Ec.) begins w ith a superficial head on epicon dylu s lateralis and in fossa radialis of the hum erus, and w ith a deep head on the forearm bones, in th e neighbourhood of the proxim al interosseous space. H alfw ay along the radius both heads join, th en pass into a tendon, th e location of w hich w as described togeth er w ith th e tendon of m. ex ten so r digiti tertii proprius. A bove the m etacarpo-phalangeal joint th is tendon bifurcates and runs to the exten sor processes of the third phalanges. 5. M. ex ten so r digiti tertii proprius (Fig. 14, 15, 17, 18, 29, 30 — Et.) originates on the epicon dylu s lateralis of the hum erus (on the cranio-lateral side) above th e origin of th e ex ten so r digitalis communis. In th e upper section th e b ellies of th ese m uscles exch an ge fibres. A t the level of th e low er xl\ of the radius, the slender ven ter passes into a tendon, w hich runs more dorsally than the tendon of th e e x ten so r digitalis communis. In the section b etw een th e carpus and tu b erosity of Me. Ill both tendons are surrounded by a com m on syn ovial sheath. On leaving th e sheath the tendon of the m. exten so r digiti tertii proprius runs more m edially, inserting on Ph. II and Ph. I ll of th e third digit. This tendon connects in the final part of its course w ith th e tendon of m. interosseus medius. 6. M. ex ten so r digiti quarti proprius (Fig. 14, 15, 17, 19, 29, 30 — Eq.) originates on the epicon dylu s exten sorius of the hum erus, near tuberculum ligam entosu m laterale of th e radius and on th e lateral border of the ulna, to w hich it is attached by m eans of a strong fascia surrounding the m uscle venter. B y m eans of this fascia it is in addition attached to the dorsal ligam ent of th e carpus, to w hich strong tendinous bands pass out from it. A bove the carpus the fusiform m u scle v en ter passes into a tendon running tow ards the fingers, w id en in g along Ph. I ow ing to reinforcem ent by the tendon of m. interosseus medius. It d ivid es into tw o branches, attached on Ph. II and Ph. I ll of th e fourth finger. b) F lexor division. 1. M. flex o r carpi radialis (Fig. 16, 18, 19, 20 — Fr.) is a flat m uscle originating on the m edial epicondyle of th e hum erus and on the m edial ligam ent of th e elbow joint. The m uscle v en ter at th e lev el of the low er 1/d of the radius in young individuals, and th e upper V 3 in adults, passes

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into a tendon running along the m edial edge of this bone. This tendon above the carpus receives a syn ovial sheath, strongly coalesced w ith the fascia antebrachii, w hich is thick in this place, and passing on th e m edial side of canalis carpalis, inserts on the second carpal bone ,and partly on Mc. III. P o 1 e i n e r (1932) in his description of the final tendon of th is m uscle did not take into account the insertion on th e carpus. 2. M. fle x o r carpi ulnaris (Fig. 16, 18, 19 — Fu.) located m ost caudally in the flex o r group, originates alm ost w ith ou t a tendon on th e m edial con d yle of th e hum erus and on the fla t aponeurosis, thrusting under the m edial head of the triceps brachii on the olecranon. The group of m uscle fibres running from this aponeurosis form s a distinct ulnar head, although closely coalesced w ith the m ain venter, the w h itely-gleam in g tendon of insertion of w hich runs on th e caudal edge of the m ain ven ter. It is sp ecially d istin ctly visible in you ng anim als. The tendon of insertion of the flex o r carpi ulnaris w hich begins at th e lev el of the carpus, coalesces w ith fascia antebrachii and inserts on th e accessory carpal bone, partly also on Mc. IV. 3. M. fle x o r digitalis superficialis (Fig. 17, 18, 19, 20 — Fs, F s’.) originates on th e m edial epicondyle of th e hum erus and d ivid es into tw o heads blended w ith each other in th e upper part. The superficial head passes into a tendon at the lev el of th e carpus, w h ile the deep head becom es tendinous sligh tly higher up. A t the lev el of the carpus both tendons, w hich are separated from each other by liga m en tu m carpi volare superficiale, are located su p erficially in relation to the tendon of th e deep digital flexor. T hese tendons coalesce h alfw ay along the m etacarpus to divide at th e lev el of the m etacarpo-phalangeal articulation into lateral and m edial branches. These branches, togeth er w ith the tendons of the superficial head of the interosseus m edius, surround the tendons of the fle x o r digitalis profundus, form ing a sheath round them . Both final branches of th e tendon of the m uscle described divide once again and each of them ends laterally on th e Ph. II. A band runs from the anterior edge of the ven ter of the deep head of the superficial digital flexor at th e lev el of the low er Vs of th e radius w hich connects w ith the tendon of the deep digital flexor. This is m. interflexorius proxim alis (Fig. 20 — a.) w hich in all the individuals w hich w e dissected contains little m uscular tissue. The situation is d ifferen t w ith m. in terflex oriu s distalis (Fig. 20 — b.) (it connects the hum eral head of the deep d igital flexor w ith th e deep head of the superficial digital flexor) w hich regardless of the age of th e anim al dissected, exh ib ited the presence of m uscle fibres,

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The occurrence of in terflexor m u scles in th e m uscle system of th e thoracic limb, has been described both in cattle and in sm all dom estic rum inants (P i t z o r n e, 1905). K o m a r e k (1958) also described it in C ervu s elaphus. P o 1 e i n e r (1932) notes the existen ce of m. interflex orius in European bison w ith o u t stating p recisely w h eth er he found the presence of both, or of one. Our observations show that both m usculi in terflexo res occur in the E. bison. 4. M. flex o r digitalis profundus (Fig. 17, 18, 19 — Fp.) is a strong flex o r m uscle consisting of three independent heads, as follow s: the hum eral radial, and ulnar heads. C ap u t hum erale (Fig. 19, 20 — Fph.) origin ates on th e lateral epicon dyle and on its crest, and is covered in this place by m. triceps brachii. The m assive ven ter of this head, threaded w ith num erous tendinous bands, exh ib its a tendency to division into tw o or three bellies. T hey pass above th e carpus into a fla t tendon, concave m edially. In th is hollow , on the boundary of the transition of the ven ter into tendon, m. interflexorius distalis is located, in young anim als com p letely m uscular, in older specim ens slig h tly tendinous, the tendon of w hich connects w ith th e tendon of insertion of the deep head of the superficial flexor (see description of th is m uscle). C ap u t radiale (Fig. 20 — Fpr.) is located th e d eep est in the im m ediate v icin ity of ossa antebrachii. It inserts on th e m edial side of the caudal surface of the radius and partly near th e interosseous space. The narrow flat ven ter passes into a tendon, w hich connects in the region of canahs carpalis w ith th e tendon of the hum eral head. The d ifferen ce, em phasised b y P o l e i n e r (1932) in the origin of th is head in relation to that in cattle does not appear ju stified to us. In all the ind ividu als dissected by us (ex cep t P latyna) th e origin did not exten d beyond the low er h alf of the radius. C ap u t ulnare (Fig. 14, 15, 16, 17, 19, 20 — Fpu.) originates on the lateral su rfaces and caudal border of th e olecranon, form ing a triangular ven ter thrust b etw een caput hum erale and m. exten so r carpi ulnaris. A t th e level of th e proxim al interosseous space the b elly of the ulnar head passes into a strong, fla t tendon, lyin g cau d o-laterally on the hum eral head of this m u scle and coalescing w ith its tendon of insertion above th e carpus. A ll the heads described have a com m on tendon located m ost deeply w ith in canalis carpalis and contained at this lev el in a syn ovial sheath reaching to th e lev el of the carpo-m etacarpal joint. At the level of the m etacarpus the tendon of the deep digital flexor is located under the tend on of the superficial digital flexor and, not reaching th e m etacarpo-

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-phalan geal joints, bifurcates into tw o branches. A t the lev el of the m etacarpo-phalangeal and proxim al interphalangeal joints th ese branches are surrounded by sheaths form ed by the tendons of the superficial digital flex o r and th e branches of the tendons of insertion of the superficial part of th e interosseus m edius. T hey th en term inate on the third phalanges. 5. M. pron ator teres w as not found by us in young specim ens. The scan ty group of m uscle fibres w h ich can be d istinguished in fascia an tebrachii in this region can only be considered as evid en ce that it is present in adult anim als. P o 1 e i n e r (1932) does not m ention this m uscle in the E. bison at all, neither does he include its absence in the chapter devoted to th e d ifferen ces b etw een th e m uscular system s of the E. bison and cattle. R e i s e r (1903) found that this m u scle w as absent in C ervu s elaphus and Capreolus capreolus. 6. M. interosseus m ed iu s (Fig. 17, 18, 19, 25, 26 — Im.) located d irectly on th e volar surface of the m etacarpus, is strongly tendinous in th e E. bi­ son and seen from the exterior gives the im pression of a tendon, but after m aking len g th w a y s cuts it is possible to find som e bundles of red m u scle fibres in its interior. It consists of tw o parts, beginning as a uniform m u scu lo-tend inou s form ation on th e lig am e n tu m carpi volare p r o fu n d u m and on the volar surface of the proxim al extrem ity of the m etacarpus. S lig h tly b elow th e origin it d ivid es into superficial and deep parts. The deep part divides h alfw ay along the m etacarpus into three branches, th e lateral and m edial of w hich run to th e extern al proxim al sesam oid bones, and th e interm edial branch d ivid es up once again into three. The w eaker, collateral branches run to th e internal proxim al sesam oid bones. The interm edial branch thrusts under th e intersesam oid ligam ent, en ters the interd igital space and passes out on th e dorsal side. Here it joins by m eans of tw o thin tw igs w ith th e tendons of th e ex ten sores proprii of the third and fourth digits. The superficial part, slig h tly h igh er than the one previously described, also divides into three branches. T he interm edial branch divides in the low er 1U of th e m etacarpus into tw o and join tly w ith the branches of th te tendon of insertion of th e su perficial digital flexor covers the tend on of th e deep d igital flexor. T he collateral branches are very w eak ly form ed and run in th e direction of th e accessory digits (II and V). The above description agrees w ith th e description of this m u scle in cattle ( S t e r b a , 1958),

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4.

T H E P E L V IC L IM B

A. Inner and ventral m uscles of the rump

1. M. iliopsoas ex h ib its in European bison, as in other species, a distinct division into tw o units: a) M. psoas m a jo r (Fig. 10, 24 — Pj.) originates on the vertebral ex trem ity of th e 13th rib, coalesces w ith m. transversus abdom inis ju st behind this rib, and inserts on the bodies and ventral surface of th e transverse processes of the lum bar vertebrae. The venter, at first broad and flat, becom es round at the lev el of th e sacrum and is surrounded from th e sides and above by m. iliacus. The strong, long tendon of this m uscle term inates on troch anter m inor of the fem ur. b) Two heads w ith d ifferen t origins can be distinguished in m. iliacus. C aput laterale (Fig. 23, 24, 29 — la.) is stronger and originates on th e tu b e r coxae, m argo sem ilunaris and on facies pelvin a of th e w in gs of th e ilium and sacrum . The concave b elly surrounds m. psoas m a jo r from th e side and from above, interchanging fibres w ith it. Caput m ediate (Fig. 24 — la.) is w eaker, and originates on th e p elvic surface of th e w in g of the sacrum and on the shaft of th e ilium . A flat ven ter interchanging w ith m. psoas m inor surrounds m. psoas major, blending w ith it, and then w ith caput laterale. Both haeds term inate in a short aponeurosis on trochanter minor. The description given above of m. iliopsoas does not in principle d iffer from the description of this m uscle in cattle, of course taking into account the differen ce in size. 2. M. psoas m inor (Fig. 24 — Pm.) is th e m ost m edially located of this group. It originates on th e caudal part of the lateral surface of the body of th e 13th thoracic and on the lateral surfaces of the bodies of all the lum bar vertebrae. The fusiform b elly at the lev el of prom ontoriu m ossis sacri begins to becom e tendinous, passing into a tendon located laterally from the lateral branch of origin of the sartorius. It term inates on tuberculum psoadicum w hich is very fain tly m arked in th e E. bison. 3. M. quadratus lu m b oru m is located m ost d orso-laterally of the w hole group. Its structure is segm ental in character, exh ib itin g individual var­ iation in arrangem ent. It originates, in all th e individuals w hich w e dis­ sected, in m uscle bundles w hich run from the caudal part of the body of 11th thoracic vertebra. Sim ilar bundles also run from the bodies of the 12th and 13th thoracic vertebrae. In som e individuals th ese bundles all term inate on the transverse process of th e first lum bar vertebra, and in others on ly the fin al one term inates in this w ay, w h ile the tw o previous ones term inate on the transverse process of the last thoracic vertebra and

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on the verteb ral extrem ity of the last rib. In th e lum bar section th e bundles of th is m u scle run from th e transverse processes of one vertebra to th e transverse processes of another vertebra, crossing som etim es over one, and som etim es even over tw o vertebrae. The final portion of th e bundles of th is m u scle term inates on the inner surface of th e w in g of the ilium . Som e few fibres insert on the ven tral surface of th e w ings of sacrum also but th is w e found in one individual only. 4. M. o btu rato r internus is a flat, fanshaped m uscle, originating on pars sacrotuberale of th e sacro-sciatic ligam ent, on tu b e r ischii, arcus ischiadicus and along th e dorsal side of s y m p h y s is pelvis. The flat b elly w ith fibres running in convergent radiation is located in th e region of foram en obturatu m , passing through this foram en and form ing a m assive tendon attached in fossa intertrochanterica of the fem ur. 5. M. o bturator extern u s originates on the outer surface of th e body of ischium and along sy m p h y s is pubis and round the border of fora m en ob­ tu r a tu m , w ith th e excep tion of the cranio-lateral part of this border. The flat ven ter surrounds the tendon of insertion of m. obturator internus and inserts by m eans of short tendon in the trochanteric fossa. 6. M. qu a dra tu s fem oris (Fig. 22, 23 — Qf.) is located caudally from the hip joint and covered from the back by m. sem im em branosus. It originates near the origin of the tendinous band described together w ith m. biceps fem oris (see description of this m uscle) and passes into a fusiform ven ter w ith fibres running caudo-ventrally. It inserts on the linea trochanterica caudalis. 7. M. g em ellu s (Fig. 22, 23 — Ge.) originates on the acetabular branch of ischium along the lesser sciatic notch and on tu b e r ischii. The flat single ven ter term in ates in a short tendon in th e trochanteric fossa. B. The lateral m uscles of the hip

1. M. biceps fem oris (Fig. 21, 27, 29, 30 — Gb.) is, in the E. bison as in other rum inants, fused w ith the caudal part of the m. g luteus superficialis, form ing a m u scle unit know n under th e nam e of m. gluteobiceps. The vertebral origin of this m uscle exten d s along th e spinal processes from the 2nd sacral to th e 2nd coccygeal vertebra. The sciatic head is attached on the lateral tu b ercules of the tu b e r ischii and on the sacro-sciatic ligam ent in its caudal part. The ven ter is d istin ctly divided into tw o parts, differin g as to location and the d irection in w hich the fibres run, w hich is particularly appa­ rent in their low er ends. A bove the stifle joint both parts pass into an aponeurosis m aking contact w ith the very strong fascia cruris. The anter­ ior part ends on the p atellar ligam ents and on the tuberosity of tibia from

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th e lateral side. The posterior part ends w ith one branch below th is tu b er­ osity, and the other (after joining w ith th e tendinous bands of the s e m i­ m em b ran osu s and sem itendinosus) on the tu b e r calcis. On th e boundary b etw een the anterior and posterior parts on th e m ed ­ ial side of the m uscle ven ter, a very strong tendinous band runs origin at­ ing on th e ven tral surface of ischium . It runs from the prom inence located cau d o-laterally in relation to foram en obturatum . This band, bordering w ith th e caudal edge of m. addu ctor fem oris (see description of this m uscle) as it approaches th e stifle joint, becom es increasingly strong and is so directed cranially that as a result it reaches to th e anterior edge of the m u scle and term inates on liga m e n tu m rec tu m patellae. The description of m. biceps fem oris in P o 1 e i n e r’s work (1932) d iffers slig h tly from our observations of th is m uscle. The d ifferen ces are p articularly evid en t in th e description of the low er part. The tendons of insertion of both parts are, according to this author, connected and term inate join tly. According to our observations this division applies both to th e m uscular parts and to th e tendons of insertion, certain d ifficu lties b ein g encountered in separating th ese aponeuroses in older specim ens. The joining of th e tendon of m. biceps w ith m. gastrocnem ius described by P o 1 e i n e r (I.e.) has, according to us, the character of a tendinous band, strongly coalesced w ith fascia cruris, w hich runs under the tendon of m. triceps surae to tu b e r calcanei. Sim ilar relations w ere found in cattle (M a r t i n & S c h a u d e r , 1938), goats, sheep ( R e i s e r , 1903), and Cervu s elaphus ( R e i s e r , I.e., S t e r b a & H e g e r o v a , 1958). T he structure and course follow ed by th e tendinous band situated on th e inner surface of th e m uscle corresponds to th e behaviour of this band in sh eep (M a r t i n & S c h a u d e r , 1938). 2. M. tensor fasciae latae (Fig. 21, 30 — Tl.) connects closely in its origin w ith th e aponeurosis of m. gluteus superficialis (w ith th e cranial part of w h ich it coalesces) and w ith th e deep layer of th e glu teal fascia covering m. glu te u s m ediu s from th e m edial side. The osseous origin of this m uscle is located on the tu b e r coxae and lateral border of the ilium . The venter, w hich is triangular in cross-section, passes on the lateral side into fascia lata, and on the m edial side of the thigh into fascia fem oralis. T he division, visib le in cattle, of this m u scle into an anterior and pos­ terior part (corresponding to th e cranial part of m. g lu teu s superficialis), w as not found by us in th e E. bison w hich w e dissected. The com plete fusion of th ese m uscle units corresponds to the relations in sm all dom estic rum inants ( R e i s e r , 1903). 3. M. g lu teu s m ediu s (Fig. 21, 22, 29, 30 — Gm.) is covered in its caudal part by m. gluteobiceps, and in the cranial part by the rela tiv ely thick

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g lu tea l fascia, w hich fu ses w ith its perim ysium . The origin is located on tu b e r coxae, crista ilica and on facies glutea of ilium , above th e insertion of m. glu teu s accessorius. In addition it also attaches on pars sacrospinalis of th e sacro-sciatic ligam ent. The ven ter of this m uscle coalesces closely w ith m. piriform is form ing its dorso-caudal m argin. The clearly visib le tendinous inscription separates th e tw o m uscles and term inates on th e ap ex of trochanter major. The d ifferen ce is also visib le in the d ifferen ce in th e direction in w hich the fibres of both b ellies run. T he b elly corresponding to th e true m. gluteus m ediu s term inates on th e lateral su rface of troch an ter major. 4. M. piriform is (Fig. 22 — Pi.) originates on the facies glutea of ilium , and surrounds tro ch a n ter m ajor w ith a fla t venter, inserting on th e caudal border of troch anter m a jo r and below . 5. M. g lu teu s accessorius (Fig. 23 — Ga.) is an elongated m u scle origin ­ ating on tu b e r coxae and facies glutea of th e iliu m b etw een the insertions of m. gluteus m ediu s and m. gluteus profundus. The venter, w ith fib res running cau d o-ven trally, term inates in a strong tendon, crossing tro ch ­ a n ter m a jo r laterally on th e caudal border of this trochanter. T his m uscle is considered by som e scien tists ( M a r t i n & S c h a u d e r , 1938; S t e r b a & H e g e r o v a , 1958) as part of m. gluteus m ed iu s in rum inants. Both the innervation of th ese m uscles and their close b lending found in certain sp ecies argue in favour of this view . As both m u scles separated d istin ctly from each other in the specim ens w e dissected, th e u se of the term m. glu te u s accessorius m ay be considered as ju stified . 6. M. gluteus p rofu n dus (Fig. 23 — Gp.) originates on the outer surface of spina ischiadica, on pars sacrospinalis of the sacrosciatic ligam en t and on th e sh aft and low er part of th e glu teal surface of the ilium . S in gle m u scle bundles are attached in addition on the lateral border of th e ilium . T he flat v en ter consists of fibres radiating outw ards and term in ates in a strong, fla t tendon along the border of troch anter major. W e did not find th e insertion of m. gluteus profundus on th e anterior surface of the p roxim al extrem ity of the fem ur as is th e case in cattle ( M a r t i n & S c h a u d e r , 1938), C ervu s elaphus and Capreolus capreolus ( R e i s e r , 1903). C. The medial m uscles of the thigh

1. M. sartorius (Fig. 24 — Sa.) is a flat, th in m uscle, one branch of w h ich runs from the m edial surface of m. psoas m ajor, and the other from crista iliopectinea, this la tter branch crossing th e tendon of insertion of m. psoas m in o r from th e m edial side. Both branches join in a narrow ven ter, runn­

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ing tow ards th e foot and coalescing in its caudal border w ith the cranial border of m. gracillis. The aponeurosis of this m uscle ends on tu berositas tibiae and below it. 2. M. gracillis (Fig. 24 — G.)is th e m ost superficial m uscle of th e m edial side of th e thigh area. The broad, flat m uscle originates in a flat tendon on th e tendon of insertion of the m. rectus obdom inis and along the ven tral border of sy m p h y s is pelvis. The upper part of th e ven ter of this m u scle covers m. addu ctor fem oris and m. sem im em branosu s, partly coalescing w ith it. It also fu ses along the m edial line w ith th e ven ter of this sam e m u scle from th e opposite side. The anterior border tou ch es the caudal border of m. sartorius and is cennected w ith it by m eans of the m edial fem oral fascia. The cross-section of the ven ter increases gradually*from th e caudal border craniad, interchanging fibres near th e th ick est part w ith m. p ectin eu s lying beneath it. A t th e lev el of the low er lU of th e fem ur, th e ven ter of m. gracillis pass­ es into an aponeurosis, closely connecting w ith fascia cruris and term inat­ ing: a) b y m eans of th e aponeurosis of m. sartorius (w ith w hich aponeu­ rosis it closely blends) on the m edial epicondyle of th e fem ur, b) on the m edial con d yle of th e tibia and below it, c) blending w ith th e fascia cruris described previously, w hich is very thick, on th e tibial tu berosity and b elo w it, d) by m eans of a tendinous band running from th e caudal, thin border of this m u scle on to tu b e r calcis. This band soon a fter leaving con­ n ects w ith a sim ilar one running from m. sem im em branosu s. P o l e i n e r (1932) in describing th e aponeurosis of insertion of m. gracillis, lim its h im self to m entioning its com m on insertion w ith m. sartorius, only adding that th e aponeurosis passes into fascia cruris. As during dissection w e succeeded in accurately tracing th e insertions by m eans of this fascia, it has been taken into consideration in describing the m u scle. In su pplem enting P o l e i n e r ’ s description em phasis is also laid on th e presence of th e tendinous band w hich is attached on tu b e r calcis and w h ich w as clearly visib le in all th e E. bison w e dissected. 3. M. pectineus (Fig. 24 — P.), originates on the sym p h ysial branch of th e pubis and partly on the cranial part of origin of m. gracillis, w hich it stro n g ly blends. T he relatively short, round ven ter in th e upper V3 of the fem u r passes into an aponeurosis covering th e tendon of m. adductor fe m o ris from the cranio-m edial side and term inating along the caudolateral border of th e fem ur, as far as the m argin of fossa flexoria from th e lateral side. M. addu ctor fem oris w as described join tly w ith m. sem im em branosus (see description of this m uscle).

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D. The anterior m uscles of the thigh

1. M.quadriceps fem oris — the strong exten sor of the knee joint, consists of four heads as it does in cattle. a) M. vastu s lateralis (Fig. 21, 22, 23, 24, 26, 29 — VL.) is a strongly d eveloped m uscle lyin g cranio-laterally in relation to th e fem ur. It takes origin in a strong, flat tendon on th e caudal and dorsal edge of troch a n ter m a jo r (the m ost su p erficially located of all those attached here). The venter, w ith fibres running cranio-ventrally, form s a h ollow on th e m edial surface for m. rectu s fem oris and m. va stu s interm edius. A bove th e stifle joint the v en ter passes into an aponeurosis attached on lig a m e n tu m re c tu m patellae and on the patella. b) M. va stu s in te rm e d iu s exh ibits a certain tend en cy to d ivision in connection w ith th e d ifferen t origin. The fibres originating b elow t r o ­ ch an ter m ajor form a narrow flat ven ter w ith fibres running tow ards th e foot, located la terally in relation to the bone. A t the level of the low er 1/* of th e fem ur th e ven ter passes on the dorsal side of th e bone and joins w ith the second part of the m uscle, w hich takes origin on the rough crest ex ten d in g cranially b etw een troch anter m a jor and caput femoris. Both parts pass into a com m on tendon, term inating under the tendon of m. va stu s lateralis. c) M. rectus fem o ris (Fig. 23, 24, 26 — Rt.) is a strongly d eveloped round m uscle originating on th e sh aft of th e ilium . It runs on th e dorsal side of th e fem ur and term in ates on the patella. d) M. vastu s m edialis (Fig. 24 — Vm.) is a flat m uscle taking origin on the cranio-m edial surface of the proxim al extrem ity of the fem ur. It term i­ nates, tegother w ith the rem aining parts of m. quadriceps on patella and lig am e n tu m r e c tu m patellae. The description g iven above of m. quadriceps fem oris in the E. bison agrees in principle w ith that given by P o 1 e i n e r (I.e.). He w as unable to take th e insertion of origin of m. rectu s fem oris into consideration, as he had only the lim b w ith ou t th e p elvis at his disposal. The description as a w h ole corresponds to th e relations found in cattle. E. The posterior m uscles of the thigh

1. M. sem iten din osu s (Fig. 21, 22, 29, 30 — Sn.) originates by m eans of an inperceptible tendon on the body of th e ischium and on tu b e r ischii. The triangular m uscle ven ter thrusts b etw een m. sem im em bran o su s and m. biceps femoris. This m uscle passes tow ards the foot on to the m edial side of the fem ur and at th e lev el of th e low er V 4 of this bone form s an aponeurosis connecting w ith fascia cruris and term inating on tubero sitas

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tibiae and on crista tibie from the m edial side. From the caudal border of this m uscle a tendon runs w hich is inserted on the tu b e r calcis — teania calcanea4). P o 1 e i n e r (1932) in describing th e insertion of this m uscle indicates the presence of th e thin tendinous bands running from its caudal border, w hich h a lfw a y along th e tibia connect w ith the tendon of m. flex o r digi­ talis pedis superficialis. T hese fibres w ere described by us as teania calcanea, esp ecially as in th e m ajority of th e cases th ey reached to the tu b e r calcis independently. The presence of th e teania calcanea in m. semitendinosus in rum inants w as established in sheep ( M a r t i n & S c h a u d e r, 1938) and am ong w ild rum inants in C ervu s elaphus ( R e i s e r , 1903). S t e r b a & H e g e r o v a (1958), in describing this m uscle in C ervu s elaphus, state that ’’The thin tendinous branches connect w ith the tendon of A ch illes”, w hich d iffers sligh tly, it is true, from R e i s e r’ s opinion (I.e.) but m ay be confirm ation of the existen ce of teania calcanea in th is mam mal. 2. M. sem im em branosu s (Fig. 21, 22, 23 — Sb.) in the European bison w hich w e dissected blends com pletely, or occasionally on ly partly, w ith m. add u ctor fe m o r is . T hey form a large, flat m uscle located under m. gracillis, partly blending w ith it. The caudal part corresponding to the true m. sem im em bran o su s takes origin on tu b e r ischii, on th e body of th e ischium and on th e sym p h y s is pelvis. Thick in its cross-section, the triangular upper part of the ven ter of this m uscle thrusts b etw een m. sem iten din osu s and m. gracillis, n ext becom ing flat and at the lev el of the low er V 3 of the fem ur passing into an aponeurosis, w hich is attached on the m edial condyles of the fem ur and tibia and on liga m en tu m collaterale m ediale of the stifle joint. The tendinous band passing from the low er V 3 of the caudal border of this m uscle term inates on tu b e r calcis. 3. M. add u ctor fem oris — separates from m. sem im em bra n osu s only in tw o of th e six individuals w hich w e dissected and on ly in the lower section. It takes origin on s y m p h y s is pelvis and partly on th e acetabular branches of the pubis and ischium . The relatively thick ven ter term inates in a broad aponeurosis on the low er 2/z of th e labium m ediale of the fem ur. The tendinous band of biceps fem oris (see description of m. biceps fe ­ moris) contacts th e cranial border of this m uscle. P o 1 e i n e r did not find, in the lim b of th e E. bison w hich he dissected, the adhesion b etw een m. addu ctor fem oris and m. sem im em b ra n o su s, In describing in great detail th e first of them , he gives in conclusion 4) T erm introduced by R. P o p l e w s k i (1939).

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a description not d ifferin g in principle from th e description giv en above. In m. sem im em branosu s the above author does not distin gu ish teania calcanea, w hich w e m ost undoubtedly did find. The presence of the teania calcanea of m. sem im em b rano su s has been established in sm all dom estic rum inants (G r a u, cit. acc. to M a r t i n , & S c h a u d e r , 1938). The adhesion b etw een the above m uscle occurs in cattle, sheep and goats. F. The m uscles of the leg and foot

a. D orso-lateral group 1. M. tibialis anterior (Fig. 22, 23, 24, 25, 26, 27 — Tb.) takes origin on the lateral surface of tuberositas tibiae and crista tibiae and on condylus lateralis tibiae. The flat venter, partly covered from the side by m. fibularis tertius, passes into a thin tendon located at the level of th e hock joint under th e tendon of m. fibularis te rtiu s and term inating on the m edial side of the proxim al extrem ity of m etatarsus. P o l e i n e r (1932) also noted the lack of division of the origin of m. tibialis anterior into tw o heads. In cattle tw o heads occur, w h ich in itially m aintain their autonom y, and blend h alfw ay along the venter. The caudal head corresponds to m. exten so r hallucis longus. The ind ivisib le m uscle ven ter of m. tibialis anterior is typ ical of sm all dom estic rum inants. (R e is e r, 1903) and of C ervu s elaphus ( S t é r b a & H e g e r o v á , 1958). 2. M. fibularis te rtiu s (Fig. 22, 23, 25, 27, 29, 30 — Ft.) takes origin togeth er w ith ex ten so r digitalis pedis longus lyin g beneath it, w ith w hich, particularly in th e upper section, it m ixes num erous fibres. The large v en ter passes above the distal extrem ity of the tibia into a tendon covering th e tendon of m. tibialis anterior. This tendon is inserted by m eans of a series of fibres on the calcaneus, II and III tarsal bone and on th e m edial side of th e m etatarsus. The description g iven above of the course taken by m. fibularis tertius and its insertions in the E. bison corresponds to the one given by P o 1 e in e r and is in accordance w ith th e relations found in cattle. 3. M. fibularis longus (Fig. 26, 27, 29, 30 — Fi.). The flat, triangular ven ter of this m uscle takes origin on the lateral fem ore-tibial ligam ent of th e stifle joint and on th e lateral condyle of the tibia, partly covering m. fibularis tertius. A t the lev el of the upper l/s of th e tibia th e ven ter greatly narrows, passing into a thin tendon running su p erficially in th e d irection of th e hock joint b etw een m. fibularis tertius and m. ex ten sor digiti quarti proprius. A bove the tarsus th e above-m entioned tendon passes m ore to th e lateral side togeth er w ith the tendon of m. exten sor d igiti quarti proprius, crossing it from th e front and entering into the

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canal bordered by os centotarsale and m etatarsus. In this canal it runs along th e m edial side and term inates on th e lateral surface of os tarsale prim um . The origin, described by P o 1 e i n e r. on tu bercu lu m lig am en tosu m laterale of the fem ur, instead of the insertion given by us on th e lateral fem oro-tibial ligam en t of the stifle joint, w ould seem to form a m atter for discussion. In v iew of th e abundant m aterial w e had at our disposal, our obser­ vations should be considered as being more objective. Regardless of w h eth er the origin on th e fem ur takes place d irectly on tu b e rcu lu m liga m en tosu m of this bone (P o 1 e i n e r) or by m eans of th e lateral ligam ent (our observations) the presence of this insertion renders the behaviour of th is m u scle sim ilar to that described in sheep ( R e i s e r , 1903). In cattle th is m uscle takes origin on th e tibial bone. 4. M. e x ten sor digitalis pedis longus (Fig. 25, 26, 27 — El.) takes origin on the lateral condyle togeth er w ith m. fibularis tertius. The v en ter of th is m uscle d ivid es at th e lev el of th e upper lU of the tibia into tw o heads, the deeper of w hich is called m. exten so r digiti terti proprius and as such w ill be described later. The superficial head h alfw ay along th e tibia passes into a tendon running tow ards th e digits, and at th e lev el of the upper l U of th e m etatarsus blends w ith th e ven ter and n ex t w ith the tendon of m. ex ten sor digitalis pedis brev is (Fig. 25, 26, 27 — Eb.) w hich runs at the lev el of the tibial tarsal bone from the dorsal surface of the tarsus. The tendon of m. ex ten so r digitalis pedis longus bifurcates im m ediately above th e m etatarso-phalangeal joint into branches running to th e third p halanges of the third and fourth digits. The description given above of this m uscle in E. bison agrees w ith the description g iv en by P o 1 e i n e r and corresponds to the relations described in cattle. 5. M. ex ten so r digiti tertii proprius (Fig. 25, 26, 27 — Et’.) takes origin join tly w ith m. exten so r digitalis pedis longus, the initial sections of the b ellies of both m uscle closely blending w ith each other. Certain authors describe them as tw o heads of one m uscle. The ven ter of m. ex ten sor digiti t^rti proprius is located below the ven ter of m. exten so r digitalis pedis longus and above th e tarsus passes into a tendon, w hich in this region runs b etw een the tendon of insertion of m. fibularis tertiu s and m. ex ten so r digitalis pedis longus and above th e m etatarsus m edially in relation to the latter. It term inates on the second phalange of the third digit, receiving above th e m etatarso-phalangeal joint reinforcem ent in the form of the branch of the deep part of m. interosseus m ediu s in a sim ilar w ay to that in th e corresponding thoracic limb.

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6. M. exten sor digiti quarti proprius (Fig. 26, 27, 29, 30 — E q\), takes origin on the lateral fem oro-tibial ligam ent, on the lateral condyle and proc. fibularis of tibia and on the lateral border of this bone, along 3A of its length. The fusiform ven ter above the hock joint passes into a tendon crossed from th e side by the tendon of m. fibularis longus. A bove the m etatarsus it runs along its dorso-lateral side, joining, in th e m ajority of cases, w ith the v en ter of m. exten sor digitalis pedis brevis and term inating on the dorso-lateral side of the second phalanx of the fourth digit. A bove th e m etatarso-phalangeal joint it is reinforced by branches of the deep part of m. interosseus m edius, like the corresponding exten sor in the thoracic limb. P o 1 e i n e r (1932) does not distinguish the origin of this m uscle on the fibular process of th e tibia. This insertion w as found by us in all the E. bisons w e dissected. The presence of this insertion w as noted in sheep, w h ile in cattle it w as absent ( M a r t i n & S c h a u d e r , 1938). b. Plantar group 1. M. triceps surae falls into tw o m uscular units: a) m. gastrocnem ius in th e E. bison is strongly developed and consists of tw o heads. C aput m ediate (Fig. 23, 24, 25, 28 — G s’.) takes origin near th e distal end of labium m ediale of th e fem ur and on th e m edial condyle of th is bone above fossa ligamentosa. In addition a strong tendinous band running from the m edial condyle of th e tibia also radiates into the ven ter of this head. C aput laterale (Fig. 22, 23, 26, 28 — Gs.) takes origin on th e lateral border of the shaft of the fem ur at the lev el of fossa plantaris, then passes into a strong venter. Both the above heads blend together below the passage b etw een them of the tibial nerve, and envelop the superficial digital flex o r from the sid es and back .At the lev el of the distal x/z of the tibia the b ellies of both heads pass into three tendons (Fig. 28 — a, b, c,) of w hich th e m iddle is th e strongest and com m on to both heads. In addition the m edial head has its ow n tendon, in itia lly located on the m edial side, then the caudal side and fin a lly passing on to th e lateral side of the com m on tendon. The lateral head has a sim ilar tendon, considerably stronger and d istin ctly separating. A bove tu b e r calcis all three tendons join and insert on this tuber below the tendon of the superficial digital flexor. In very young individuals the tendon of the m edial head cannot be clearly separated from the m iddle tendon. In older specim ens (in our m aterial specim ens over 6 m onths) this tendon is clearly separated.

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P o 1 e i n e r, in describing the tendons of insertion of both heads of the m. gastrocnem ius in the E. bison, found that th ey are joined. This ’’ju n ction ” on account of th e direction in w hich the fibres run, the close blending w ith th e b ellies of both heads and also the possibility of dissecting it from the tw o rem aining tendons of insertion, has been described b y us as a separate, m iddle tendon of insertion of th e gastrocnem ius com m on to both its heads. b) m. soleus (Fig. 22, 23, 26 — Sf.) is a w eakly form ed m uscle running from th e lateral condyle of tibia and blending w ith th e tendon of origin of m. exten so r digiti quarti proprius. The very thin, flat ven ter joins the lateral head of m. gastrocnem ius by its tendon. We did not find the tendinous band, described by P o 1 e i n e r, w hich according to th is author w as supposed to correspond to ten do solei, in any of the specim ens w e dissected. 2. M. flex o r d igitoru m pedis superficialis (Fig. 23, 25, 26, 28 — Fsc.) takes origin in a short but thick tendon of origin in fossa flexoria of the fem ur. A strong tendinous ven ter blends on its caudal border w ith the heads of m. gastrocnem ius. In th e distal V 3 of the tibia this ven ter passes into a tendon located in itia lly under, then laterally, and fin ally above th e tendons of m. triceps surae. This tendon from above and from the sides en velop s tu b e r calcis, inserts on it, and then runs tow ards th e digits, being located su p erficially on the plantar surface of th e foot. A bove th e m etatarso-phalangeal joint this tendon divides into tw o branches, both of w hich connect w ith the corresponding branches of the superficial part of m. interosseus m edius. H alfw ay along the first phalanx each of the branches of the tendon of the superficial digital flex o r again divid es into tw o, th u s adm itting into this bifurcation the corresponding branch of the tendon of the deep digital flexor and inserting on the proxim al extrem ity of the second phalanx on plantar side. The description given above agrees w ith the description of this m uscle in the E. bison given by P o 1 e i n e r, and does not in principle d iffer from th e description of this m uscle in cattle. 3. M. flex o r digitalis pedis profundus (Fig. 25, 26, 29, 30 — Fpc.) is a group of flex o r m uscles term inated by a com mon tendon. T hese are: a) M. flex o r hallucis longus (Fig. 23, 25, 26 — Fh.) takes origin on the caudo-lateral side of the lateral condyle of the tibia and on the fascial septum lying b etw een it and m. ex ten sor digiti quarti proprius. The m uscle ven ter above the tarsus passes into the tendon running from th e m uscle groove above su sten tacu lu m tali. B elow th e hock joint this tendon joins w ith the tendon of m. tibialis posterior. b) M. tibialis posterior (Fig. 25 — Tp.) takes origin on the lateral condyle of the tibia near the popliteal notch and on the m uscular lines of this bone,

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The flat, th in ven ter blends in th e proxim al section w ith the ven ter of the p revious m u scle and h a lfw a y along the tibia passes into a tendon, to w hich th e tendon of the m. flex o r digitalis pedis longus is joined b elow the tarsus. c) M. flex o r digitalis pedis longus (Fig. 25 — FI.) takes origin on the lateral con d yle and fibular process of the tibia. The m u scle' ven ter h a lfw a y along th e tibia passes into a fla t tendon, running on th e caudo-m ed ial side o f th e tibia in the groove on m. flex o r hallucis longus, and later in a sim ilar grove on the tibial bone. This tendon continues on the m edial side of th e hock joint and joins w ith th e tendon of th e tw o rem aining heads below th e fibular tarsal bone. The tendon of insertion com m on to th e three above-m entionad m u scles runs tow ards th e digits along m. interosseus under the tendon of the su perficial d igital flexor. A bove the m etatarso-phalangeal joint this tendon is surrounded by a sheath form ed by branches of th e superficial part of m. interosseus m edius. It divides into tw o branches term inating on th e third p halanges of th e third and fourth digits. The description of this m uscle in the E. bison agrees w ith th at of the sam e m uscle in cattle. 4. M. po pliteu s (Fig. 22, 23, 25 — Po.) takes origin on the lateral condyle of the fem ur in a strong tendon, thrusting under the lateral fem oro-tibial ligam en t of th e stifle joint. The m uscle ven ter, in itially rounded, covers capsula articularis from th e back and th en becom es flat, inserting fan w ise in the m edial border of th e tibia. T his m u scle b eh aves sim ilarly in E. bison to that in cattle. c. M uscles of m etatarsus 1. M. interosseus m ediu s is located on the plantar surface of the m etatarsus and consists of tw o parts: th e deep and superficial, th e further division and course of w hich correspond to the relations present in the thoracic limb. 2. M. e x ten so r digitalis pedis brevis (Fig. 25, 26, 27 — Eb.) originates in th e European bison on the dorsal ligam ents of the hock joint. T he flat m u scle venter, situated b etw een the tendons of m. exten sor digiti qu arti proprius and m. ex ten sor digitalis pedis longus, joins the latter at th e lev el of the proxim al 1/z of th e m etatarsus. IV. D IS C U S S IO N A N D R ESU LTS

The results given of our observations of the sk eletal m uscular system of th e European bison indicate the existen ce of specific featu res in the form ation of m any m uscle units com pared w ith the corresponding m u scles

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in dom estic cattle. This refers here both to features of structure and to th e topography of individual m uscles of d ifferen t regions of th e body of the E. bison. A m ong th e m uscles of the head of the E. bison the follow in g are w orthy of note: the degree of d evelop m en t of the cutaneous m uscle system and th e d ifferen ces in d etails of structure of such m uscles as m. tem poralis and m. biven ter. The stronger d evelopm ent of the cutaneous m uscle system in the E. bison is connected w ith the considerable thickness of the skin. This connection w as em phased in greater detail by S w i e z y n s k i & P i 1 a r s k i (1956). D ifferen ces in th e structure of th e m andibular m uscles result in the greater ex ten t of the insertions of m. tem poralis and m. b iv e n te r m and ibulae. The degree of form ation of m. tem poralis in rum inants is, accord­ ing to B o r o w i e c (1950) dependent on the participation of th e incisor teeth in the process of taking in food. The function of th e ven ter anterior of the m. b iv e n te r is ch iefly to depress the m andible. Consideration of these observations leads to th e conclusion that the incisors play a greater part in the m echanics of taking in food in the E. bison than in dom estic cattle. This is confirm ed by observations of th e biology of th is species. Hard shoots and th e bark of trees form a large part of the food of th e E. bison, and to bite th ese off requires strong pressure of the incisors and w ider opening of the m outh. The appearance of a live E. bison is in itself su fficien t to rouse interest in the structure of th e m u scles of the w ith ers (regio dorso-scapularis) in that th ey participate in the form ation of the characteristic ’’h um p” of the European bison. This ’’h um p” is the result on the one hand of the u nusually high spinous processes of the thoracic vertebrae, and on the other of the strongly d eveloped m uscles of this region w h ich contribute to its form ation. The m uscles involved in this form ation are prim arily pars cervicalis m usculi rhom boidei, m. splenius, m. sem ispinalis capitis and m. spinalis cervicis. The exp lanation th at th e greatly developed m. splenius and m. sem ispi­ nalis capitis are due to th e effect of the considerable w eigh t of the head of this anim al does not seem convincing, despite th e fact that the opinions of certain authors are unanim ous in this respect and su ggest the inter­ dependence b etw een th e w eigh t of the head and th e degree of d evelop­ m ent of th e spinous processes of the m uscles of the w ithers region. This v iew is refuted both by th e results of K r y s i a k ’s investigations (1951) and those of R o s k o s z & E m p e l (1960). The occurrence of m assive m uscle units intended so lely for a fu n ction so little dynam ic as supporting th e head, w ith the sim u ltaneous existen ce for this purpose of an ’’econom ­ ica l” construction in th e form of ligam en tu m nuchae w ould be unjustified

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’’p rod igality” in N ature. T hese m uscles, in addition to th e function of liftin g the head, particularly im portant to th e m ales during fig h ts (hence probably, their sp ecially strong developm ent in th e represen tatives of this sex) act on one side on ly and bend the spine in their direction. T he livin g conditions of the E. bison, com bined w ith th e great mass of their bodies, m ake it essential for this m am m al to be very agile, w hich is connected, in te r alia, w ith the ’’ela sticity ” in the lateral m ovem en ts of its spine, esp ecially in the cervical region. This requirem ent is m et by th e strong d evelop m en t of the m u scles referred to above. C onfirm ation of these suppositions is provided by th e structure of certain m uscles of th e epaxial group. M. iliocostalis and m. longissimus dorsi, lim ited in th e m ajority of E. bison to the lum bo-thoracic section of the spine, do not reach the cervical vertebrae, as is the case in dom estic cattle. M. sem ispinalis capitis and m. spinalis cervicis are strongly developed in the E. bison and clearly separated from m. sem ispinalis et spinalis dorsi. Such ’’au ton om y” in the ex ten t to w hich the d ifferen t sections of th e spine are capable of m ove­ m ent is of great assistance to the E. bison under its livin g conditions. The im portance of m. rh om boideus in the m echanics of w alking in these anim als has not been sp ecially em phasised, despite th e fact that it fu lly d eserves attention. T he cervical part of the rhom boideus, during th e free phase of th e thoracic limb, contracts the cartilage and anterior an gle of th e scapula, thus draw ing the shoulder joint backwards. P o p l e w s k i (1937) in his in vestigation s of the m echanics of w alking, considers the shoulder joint as the ch ief centre of m ovem ent in this phase, and the group form ed by biceps brachialis and m. brachiocephalicus as th e gen er­ ators of this m ovem ent. He considers that the cause of the great d evelop ­ m ent of m. brachiocephalicus in quadrupeds is th e im portance of this v ery function. A ll other m ovem ents take place in the rem aining joints and, being the result of contractions of other m uscles, possess in this sphere, according to this author, a secondary, com pensatory character in relation to the action taking place in th e region of the stylopodium . The question as to w h eth er in the case of E. bison th e a ctivity of th e cervical part of rhom boideus is of such a secondary character requires, in m y opinion, further investigation. W r ó b l e w s k i , in describing th e w alk of the European bison w rites: ”... it gives the im pression that the anim al supports th e w h ole w eigh t of its body on the hind legs, testing the ground w ith the front ones, ready at any tim e to draw back. The hum p, w hich assists this original w ay of w alk in g” 5). Both W r o b l e w s k i ’s observations cited above and the structure of this m uscle, described in the d etailed part of this work, indicate its im portant participation in th e m e­ 5) E xtract cited above tran slated from P olish,

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chanics of w alking. The function of th is m u scle in th e support phase of the lim b w ould seem to be of even great significance and worth investigating. The rem aining m uscles of the thoracic lim b in th e E. bison do not exh ibit special d ifferen ces in com parison w ith the corresponding m uscles in dom estic cattle. The sole excep tions to this are m. ex ten sor carpi radialis, and m. abdu ctor pollicis longus, w hich in the E. bison u n exp ected ly inserts in the sam e w ay as in sm all rum inants. In th e E. bison the m. p ro­ nator teres is absent, as it is in Capreolus capreolus and C ervu s elaphus. The m uscular system of the w alls of th e thorax is sim ilar to the m uscular system of th is region in dom estic cattle — apart from the insertions of m. transversu s costarum, w hich also behave sim ilarly in the E. bison, to those in sm all rum inants. A m ong the m uscles of the abdom inal w alls of the E. bison, the strong d evelop m en t of the aponeurosis of m. transversu s abdom inis and the w ide ex ten t of m. rectu s abdom inis are notew orthy. Such structure of the above m uscles, togeth er w ith the very pow erful fascia system , result in the abdom en of th e E. bison being suspended very tau tly, and even in h igh ly pregnant fem ales th e abdom en does not hang dow n or becom e flaccid. A m ong m uscles of the p elvic lim b w hich d iffer from the corresponding m u scles in dom estic cattle are m. gluteobiceps, m. ten sor fasciae latae, m. gracilis, m. sem im em branosu s, m. a d d u ctor fem oris and also m. tibia­ lis anterior, m. fibularis longus, m. exten so r digiti qu arti proprius, m. ga­ strocnemius. The first fiv e m uscles form a group of units closely connected w ith the motor properties of the p elvic limb. The functional im portance of th is group is esp ecially distinct in the support phase of the lim b (w ith the excep tion of m. tensor fasciae latae), w h en the ch ief centre of m ovem ent is situated in th e stifle joint, and the effect of contraction of m. gluteobiceps is m anifested by the pushing forw ards on its p elvico-vertebral attachm ents. The effect of this is to ’’push” th e p elvis forwards, and w ith it the w h ole trunk. M. sem im em bra n osu s and sem itendinosus in this phase of m ovem en t co-operate w ith m. gluteo-biceps and in cattle are bi-articular m u scles (hip and stifle joints). In the E. bison, ow ing to the presence of teania calcanea of these m uscles, th ey becom e tri-articular and are sim u ltaneously exten sors of the foot on account of the insertion of th ese bands on tu b e r calcis. P o p l e w s k i , in the work cited above, lays special em phasis on the im portance of th e m ovem ent of the foot in th e function of the motor m echanism of th e p elvic lim b. He em phasises the part played in this m ovem ent of teania calcanea of the biceps fe m o ­ ris, and of the w hat he term s ”tuberal fle x o r e s ”. In th e light of th ese reasonings m, sem im em bra n osu s and m. sem iten din osu s in the E. bison

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also play an au xiliary part in th e m ovem en ts of the foot and in this w ay increase the motor capacity of th e p elvic limbs. A sim ilar functional im portance m ust be attributed to the teania calcanea of m. gracillis. M. addu ctor femoris, ow ing to its blending w ith m. sem im em branosus, co-operates w ith it in th e functions described above. M. tensor fasciae latae plays an im portant part in th e sphere of th e free lim b, w h en the centre of m ovem ent sh ifts to the hip joint, and the distal extrem ity of the fem ur sh ifts forwards. The blending of both parts of m. tensor fasciae latae w hich occurs in the E. bison m ay affect its functional im portance, not in th e sense of th e direction in w h ich it acts, but in its dynam ic capacity. The functional consequences of th e preservation in th e E. bison of the m. m. tibialis anterior, fibularis longus and ex ten so r digiti quarti proprius are d ifficu lt to defin e w ith ou t m aking biom echanical in vestigation of these m uscles. As stated in th e introduction th is w as not th e aim of our work. These m uscles (sim ilarly to m any others m entioned above) exh ib it sim i­ larity to th e relations occurring in th is respect in sm all rum inants, both dom estic and wild. REFERENCES 1. A k a j e w s k y, A., 1931: Zur M orphologie des M. le v a to r nasolabialis und des M. m ala ris bei ein igen H au stieren . A nat. A nz., 73, 1/3: 1— 22. Jena. 2. A 1 b r e c h t, R., 1944: Zur A n a to m ie des B ovid en h erzen s. D iss., Leipzig. 3. B i j v o e t, W. F., 1908: Zur v e r g le ich en d e M orfologie der Musculus digastricus m a n d ib u la e bei den S ä u g etieren . Z tschr. Morph. A ntrop., 11. B erlin (acc. to Thiel). 4. B o a s & P e t e r s e n , 1921: D u k k elen hos Z ebuochsen. P e n K gl. V eterinair og L andbohöjskole A arsk rift. (acc. to M a r t i n & S c h a u d e r , 1938). 5. B o c h e n e k, S., 1955: Szczątk i żubra B ison bonasus (L.) z Podhala. A cta theriol., 1: 15—25. W arszaw a. 6. B o r o w i e c, S., 1950: W spózależność m ięśn io w o -zęb o w a . F olia m orphol., 1(9), 1: 88— 120. W arszaw a. 7. C h o m i a k, M., 1947: M ięsień p o liczk o w y (m u scu lu s buccinatorius) u p rzeżu ­ w aczy (krow a, ow ca, koza). Ann. U n iv. M. C u rie-S k łod ow sk a, C2, 9: 214— 233. L ublin. 8. F r a n c k, L., 1838: H andbuch der A n a to m ie der H au stiere. B erlin, (acc. to A k ajew sk y, 1931). 9. H a m p l , A. & K a m a n , J., 1959: H lu b ok ä sv a lo v in a ni. facialis u srnce a je le na. Cs. M orfol., 7, 2: 144— 156. Brno. 10. H u b e r , E., 1924: U ber d ie B ed eu tu n g der ex p e r im e n te lle n M ethode in der F a cialisforschun g, n eb st B etra ch tu n g en über d ie p h y lo g en etisch e E n tw ick lu n g der F acialism u scu latu r in der V erteb ra ten -R eih e. A nat. A nz., 58, 24: 177— 205. Jena. 11. K a m a n, J. & H a m p 1, A., 1958: S v a ly k rku jelen a (C e rv u s elaphus L.). S b ornik VSZL, B, 6 (27), 2: 134— 146. Brno. 12. K a m a n , J. & H a m p l , A., 1959: P o v rch o v ä sv a lo v in a ni. facialis u srnce (Copreo lus capreolus L.). Sb orn ik VSZL, B, 7 (38), 1— 3: 93— 118, Brno,

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13. K o 1 d a, J., 1950: S ro v n â v a ci an atom ie zvirat dom âcich. 3— 4: 1— 426, S tu d . Org. C. S. V et.-M ed., Brno. 14. K o m â r e k , V., 1958: N ék teré poznâm ky ka sv a lo v in e hrudni k on ćetin y jelen a. Sbornik VSZL, B, 6(27), 2: 147— 159. Brno. 15. K r ü g e r , W., 1927: M u scu lu s m u ltifid u s und Musculi rotato re s der H a u ssä u g e­ tiere und ih re B ezieh u n g en zur D reh fä h ig k eit der W irbelsäule. A nat. A nz., 63, 21/24: 305— 327. Jena. 16. K r y s i a k, K., 1951: W ięzadło k a rk o w e (L ig a m e n tu m nu chae) żubra (B ison bonasus L.). F olia m orphol., 2(10): 271— 283. W arszaw a. 17. K r y s i a k, K., 1960: T he E uropean B ison (Bison bonasus). S ta te Counc. C onserv. N at. P oland, 10: 1— 38. K raków . 18. M a r t i n, P. & S c h a u d e r, W., 1938: L ehrbuch der A n atom ie der H au stieren . 3: 1— 560. Sch ick h ard -E b n er, Stuttgard. 19. M a x i m i e n k o, A., 1928: Zur M orphologie des M. o bliq uu s a b d o m in is intern us ein iger S äu getiere. A nat. A nz., 64, 18/19: 358— 371. Jena. 20. M i 11 o t, J., 1945: Les bisons eu rop éen s des collectio n s du M uséum d ’H istoire N atu rrelle. M am m alia, 9, 1: 1— 19. Paris. 21. de M o u 1 i n, F., 1924: D ie o b erfläch lich en R ü ck en - und H alsm u sk eln des in d i­ schen B ü ffels und des Z ebus. A nat. A nz., 58, 19/21: 460— 471. Jena. 22. M ö r i k e, K., 1954: V e r g leich a n d e-fu n k tio n elle M orphologie der R u m p f-O b er­ arm m u skulatu r der S äu g etiere und des M enschen. G egenbaurs m orph. Jb., 94, 1— 2: 165— 237. L eipzig. 23. N i c k e l , R., S c h u m m e r , A. & S e i f e r 1 e, E., 1954: L ehrbuch der A n atom ie der H au stiere, B ew egu n gsap p arat., 1: 1— 502. P aul P arey. B erlin — H am burg. 24. P i ę k o Ś, M., P i l a r s k i , W. & R o s k o s z, T., 1958: O b serw acje nad dłu gością jelita u żubra (Bison bonasus L.). F olia m orphol., 9(17), 1: 69— 79. W arszaw a. 25. P i l a r s k i , W. & R o s k o s z, T., 1957: Z jaw isk o u k rzyżow ien ia (sacralisatio) o statn iego kręgu lęd źw io w eg o u sam ic żubra (Bison bonasus L.). F olia m orphol., 8(16), 2: 110— 119. W arszaw a. 26. P i t z o r n o, M., 1905: M uscu lu s in terfle x o riu s. A tti. Soc. Tosc. sc. nat. Pisa. Proc. verb., 14. Pisa. 27. P o 1 e i n e r, R., 1932: D er an atom isch e A ufbau der E x trem itä ten beim Euro­ p äisch en W isent (Bison bonasus L.) in verg leich zum H ausrind. D iss. W ien. 28. P o p 1 e w s k i, R., 1937: B adania nad m ech an ik ą chodu. W iad. w et., 204: 249— 278. W arszaw a. 29. P o p 1 e w s k i, R., 1939: A n atom ia ssak ów . 3: 1— 142. Kom. W yd. Pol. A kad. W ar­ szaw a. 30. R e i s e r , E., 1903: V erg leich en d e U n tersu ch u n gen über S k elettm u sk u la tu r von H irsch, Reh, S ch a f und Z iege, 1— 42. D iss. B ern-W ien. 31. R o s k o s z, T. & E m p e l , W., 1961: T he size of th e head and the h eig h t of sp in ou s processes in th e region of th e w ith ers of the E uropean bison, Bison bo­ nasu s (L i n n a e u s 1758). A cta th eriol. 5, 6: 63—71. B iałow ieża. 32. S t ë r b a, C., 1958: V olârni a p la n ta m i krâtk é svaly m etap od ia skotu. C eskoslov. M orfol., 6, 3: 252— 256. Brno. 33. Ś t e r b a , E. & H e g e r o v a , E., 1958: M yologie p an evn i k on ćetin y jelen a (Cerv u s elap hu s L.). Sbornik VSZL., B, 6(27), 2: 162— 169. Brno. 34. S t i m p e 1, J., 1934: D er M orphologie des m edialen M u sk elstran ges der S ta m m ­ zone b ei den H au stieren . M orph. Jb. 74, 3: 337— 363. Leipzig. 35. S w i e ż y ń s k i , K. & P i l a r s k i , W., 1956: U m ięśn ien ie sk órn e żubra (Bison b on asu s L.). F olia m orphol., 7(15), 2: 133— 138. W arszawa.

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36. T h i e l , W., 1954: D er tie fe F acialm u sk u latu r und die H y o id verb in d u n g des M uscu lu s b iv e n t e r der S ä u g etiere. G egenbaurs m orph. Jb., 94, 3— 4: 391— 451. L eip ­ zig. 37. T u r k i e w i t s c h, K., 1928: Zur M orphologie des M. tr a n s v e r s u s tr u n ci der S äu g etiere. A nat. A nz., 66, 16/18: 281—292. Jena. 38. W ę g r z y n, M. & S e r w a t k a, S., 1961: L ig a m e n tu m sacro tu bera le la tu m bei B iso n bonasus ( L i n n a e u s 1758) und Bos ta u r u s ( L i n n a e u s 1758). A cta th e riol. 5, 7: 73—97. B iało w ieża . 39. W i l k u s , E., 1957: P o ró w n a w czo -a n a tcm iczn e badania nad gło w ą żubra, B ison b on a su s (L.). I. C a v u m oris. A cta theriol. 1, 7: 184— 307. W arszaw a. 40. W r ó b l e w s k i , K., 1927: Żubr puszczy b ia ło w iesk iej. W yd. P olsk ie: 1— 232. P ozn ań. 41. Ż a b i ń s k i, J., 1947— 1951: K sięgi R odow odow e Ż ubrów . W yd. M iędzyn. Tow. Ochr. Żubra: 1— 155. W arszaw a. W arsaw A gricu ltu ral U n iv ersity , D ep a rtm en t of A n im al A n atom y, W arsaw , G rochow ska 272. STRESZCZENIE P iśm ien n ictw o z zakresu an atom ii żubra n ie jest bogate i d otyczy przede w sz y st­ kim szk ieletu . T rudności w zdobyciu i p rzech ow yw an iu m ateriału w p ostaci tak rzad k iego a jed n ocześn ie tak dużego ob iek tu badań sp o w od ow ały, że n ieliczn e p u ­ b lik a cje dotyczące an atom ii części m ięk k ich tego gatu n k u m ają w w ięk szo ści ch a­ rak ter przyczynków . Jako m ateriał do badań p o słu ży ły zw ło k i 20 żubrów różnej płci i w iek u , pad łych w latach 1950— 1960, będące w d ysp ozycji O środka B adań nad A n atom ią Żubra przy Z ak ład zie A natom ii Z w ierząt W ydziału W eteryn aryjn ego S.G.G.W . w W arszaw ie. P rep aracje anatom iczną p row adzono częściow o na m a teria le św ieży m , g łó w n ie zaś na u trw alon ym drogą nastrzy k iw a ń d otętn iczych p łyn am i k o n serw u ją cy m i lub przez p rzech o w y w a n ie zw łok w basen ach z w od n ym roztw orem form alin y. Z am ieszczon e w pracy rycin y w yk on an o na p od staw ie fotogram ów .8) P o ró w n a n ie w y n ik ó w badań nad u m ięśn ien iem szk ieleto w y m żubra z dan ym i z tego zakresu u bydła d om ow ego oraz in n ych p rzeżu w aczy dom ow ych i dzikich, w yk azało obecność szeregu różnic. D otyczą one zarów no zew n ętrzn ych cech m o rfo ­ logiczn ych jak i topografii jed n o stek m ięśn io w y ch różnych okolic ciała żubra. W śród m ięśn i g łow y żubra na u w agę zasłu gu ją m ięśn ie sk órn e i szczegóły b u d o­ w y tak ich m ięśn i jak rn. t e m p o r a lis i m. b iv e n t e r m an dib u lae. S p ecjn a lie w yraźn e różnice m ięd zy m ięśn ia m i sz k ieleto w y m i żubra i b yd ła d om o­ w eg o zaznaczają się w o k olicy g rzb ieto w o -ło p a tk o w ej, regio dorsoscapularis. D o ty ­ czy to m. rh o m b oid eu s, m. splenius, m. sem ispin a lis capitis, m. spinalis cervicis. W gru p ie m ięśni n ad osiow ych żubra ze w zględ u na to p ografię p rzyczep ów pod ­ k reślen ia w ym agają tak ie m ięśn ie jak: m. iliocostalis, m. longissim us dorsi, m. s p i­ nalis e t sem isp in a lis dorsi. Z m ięśn i kończyn, które b u d ow ą sw ą od b iegają od sto su n k ó w w y stęp u ją cy ch u b yd ła d om ow ego, w y m ie n ić należy: m. e x te n s o r carpi radialis, m. a b d u c to r pollicis longus, m . g luteob icep s, m. te n s o r fasciae latae, m. gracillis, m. s e m im e m b ra n o s u s, 8) R ysu n k i w yk on ał lek. w et. M ieczy sła w W ę g r z y n .

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m. a d d u c to r fem o ris, m. tib ialis an terio r, m. fibu la ris longus, m. ga strocn em iu s, m. e x te n s o r d igiti ą u a rti p ro p r iu s k oń czyn y m ied n iczn ej. Oprócz w y m ien io n y ch w yżej m ięśn i m niej w y ra źn e różn ice w porów n an iu z u m ięśn ien iem szk ieleto w y m byd'a w y k a zu je szereg in n ych jed n o stek m ięśn io ­ w ych żubra. Z aob serw ow an e cech y u m ięśn ien ia szk ieleto w eg o żubra dają p ew n e p od staw y do tłom aczen ia ich sp ecyficzn y m i w aru n k am i b yto w a n ia tego zw ierzęcia. W yniki pra­ cy stw arzają ponadto m orfologiczn e p od staw y do dalszych badań tego zagad n ien ia pod kątem analizy biom ech an iczn ej.

FIGURES

P la te X X X . Fig. 1. M uscles of head of E uropean bison (The su p erficia l m u scles are rem oved). Fig. 2. M uscles of la ry n g ea l region of European bison. M. m ylo h yo id eu s : a — anterior part, a’ — posterior part.

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P la te X X X .

P la te X X X I. F ig. 3. M. t e m p o r a l is of E uropean bison. F ig. 4. M andibular m u scles of E uropean bison. D eep d issection . B ody and ram us of th e le ft h a lf of m an d ib le are p a rtia lly rem oved.

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P late X X X I.

P la te X X X II. Fig. 5. M uscles of neck and th orax of E uropean bison, cf. R ight side. T he su p erficia l m u scles and thoracic lim b are rem oved. F ig. 6. T opography of M. splen ius of E uropean bison, d1. T he preparation is v iew ed from th e righ t sid e and so m ew h a t dorsally.

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P la te X X X II.

P la te X X X III. F ig. 7. M uscles of th e neck, back and loin s of E uropean bison, cT. R igh t side. M. splen ius is rem oved.

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P la te X XX IIT.

P la te X X X IV . Fig. 8. M uscles of th e neck, back and lo in s of E uropean bison, cf. R ight side. M. iliocostalis is rem oved . M. m. sem isp in alis capitis, long issim u s capitis and lon gissim us a tlan tis are partially rem oved. Fig. 9. M uscles of th e neck, back, and loins of E uropean bison, cf. R ight side. L ateral part of rn. spin a lis e t sem isp in a lis dorsi and m. lo n g issim u s cervicis are e n tirely rem oved.

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P la te X X X IV .

P la te X X X V . Fig. 10. M. m u ltifid u s of E uropean bison. R ight v iew . Fig. 11. M. longus colli of E uropean bison. V en tral v iew , a — cerv ica l part, b — thoracic part.

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P la te X X X V .

P la te X X X V I. Fig. 12. M. rectu s a b d o m in is of E uropean bison. V en tral v ie w . Ra —- la tera l portion; R a’ — m ed ial portion; 1 — p la cem en t of th e xip h oid cartilage; 2 — p lacem en t of linea alba', 3 — p lacem en t of th e atach m en t on the p elvis; 4,5 — fou rth and fifth costa l ca rtilages; 6 — p lacem en t o f the costal arch. Fig. 13. M. tr a n s v e r s u s thoracis of E uropean bison. D orsal v iew . stern i; b — d ia p h r a g m a (cut).

1 — m a n u b r iu m

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Ra'

P la te X X X V II. Fig. 14. M u scles o f sh ou ld er and arm of E uropean bison. L a tera l v iew . Fig. 15. M u scles o f sh ou ld er and arm o f E uropean bison. L ateral v ie w . M.m. d e l t o id eu s and bic ep s brach ii are p a r tia lly rem oved.

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P la te X X X V II.

P la te X X X V III. Fig. 16. M uscles of sh ou ld er and arm of E uropean bison. M edial v ie w . M. tensor fasciae a n te b r a c h ii is p artiaiy rem oved.

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P la te X X X V III.

P la te X X X IX . Fig. 17. M uscles of rig h t thoracic lim b of E uropean bison from elb o w d ow n w ard s. L ateral v ie w . Fig. 18. M uscles of rig h t thoracic lim b of E uropean bison from elb ow dow nw ards. M edial v ie w . Fig. 19. M uscles of righ t th oracic lim b of E uropean bison from e lb o w dow nw ards. V olar v iew .

a

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P la te X X X IX .

P la te XL. F ig. 20. M uscles of an teb rach iu m o f E uropean bison. D eep d issection , m ed ia l v iew . M. f l e x o r ca rp i u ln aris is co m p letely and ca p u t su p erficialis m i. fle x o ris su perficialis is p a rtia lly rem oved , a — m . in te r fle x o r iu s p ro x im a lis; b — m. in te rfle x o riu s distalis. Fig. 21. M u scles of g lu tea l and fem o ra l region of E uropean bison. L eft side.

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P la te X L

P la te X LI. Fig. 22. M uscles of g lu tea l and fem o ra l region of E uropean bison. L eft side. M. g lu te o b ic e p s is rem oved. F ig. 23. M uscles of g lu tea l and fem ora l region of E uropean bison. D eep d essection . 1 — tu b e r sacrale; 2 — tu b e r coxae; 3 — tu b e r ischii, 4 — N. tibialis.

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P ía te XLI

P la te X L II. F ig. 24. M uscles of p elv is and thigh of E uropean bison. L eft sid e, m edial view .

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P la te X L II.

I

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P la te X L III.

Fig. 25. M uscles of le g ' and fo o t of E uropean bison. M edial v ie w . 1 — patella; 2 — fe m u r; 3 — tib ia ; 4 — calcaneus; 5 — m e ta ta r su s. Fig. 26. M uscles of righ t leg and foot of E u ro p e a n . bison. L a tera l v iew .

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P la te X L III.

P la te X LIV . Fig. 27. M uscles o f righ t leg and fo o t of E uropean bison. D orsal v iew . F ig. 28. M. g a stro c n e m iu s of E uropean bison. P la n ta r v iew , b — m id d le tendon; c — la tera l tendon.

a— m ed ial tendon;

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P la t e X L ÍV .

P la te X LV . F ig. 29. M u scu latu re o f E uropean bison, cT. T he su p erficia l m u scle

are rem oved.

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P la te XLV.

P la te X LV I. Fig. 30. M u scu latu re of E uropean bison, cf.

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Píate XLVT.

K EY TO THE LETTERIN G A — to, anconeus A p — to. abductor poZiieis lo n g u s A o — m. a tla n lo c cip ita lis B , B ’, B ” — to. bu ccin ato riu s B b — m, biceps bracZiii B e — m , bracliialis B r — to. brachiocephalicus B v — m . b i v e n l e r (v e n t e r a n te r io r) B v ’ — m . b ioen ter (ven ter posterior) C —■m . caninus Cb — to, coracobracZiialis Cf — to, cutaneu s /row talis Cl — to. cleid om asfoid eu s Co — to. cutan eus lab ioru m Cu •— to. cutaneus faciei D — m. deltoid eus D ’ — m. d e lto id e u s (p. acrom ialis) D1 — m . d e p resso r tab ii m a x illa r is D m — to, d ep res so r la b ii rnandibularis Eb — m. ex t. digit, p e d is b re v is Ec — m . e x t . digit, co m m u n is E l — m . e x t . digit, p e d is longus Eq — to. ext. d igiti IV p ro p r iu s E q ’ — to. ex t. d ig i ti IV p ro p r iu s (pelvic lim b) Er — m . e x t . ca rp i radialis E t — to. ext. d ig iti III p ro p r iu s E t’ — m . e x t . d ig iti 111 p ro p r iu s (pelvic lim b) Eu — m. e x t. c a rp i u ln aris Fh — to, flex , hallucis longus F i — m .f a b u la r i s longus FI — to. flex , digit, p e d is longus Fp — to. flex, digit, p ro fu n d u s F ph — to, flex , digit, p r o fu n d u s caput h u m era le Fpr — to. flex , digit, p r o fu n d u s c a pu t radiale F pu — to. flex , digit, p r o fu n d u s c a pu t u ln are Fpc — to. flex', digit, p e d is p r o fu n d u s Fr — to. flex, ca rp i radialis Fs — m. fle x , digit, su perficia lis F s ’ — to. flex , digit, su p erficia lis (caput p ro fu n d u m ) F sc — to. flex , digit, p e d is superficial. F t — to. f ib u la ris te r tiu s F u — to. fle x o r c a rpi u ln a ris G — to, gracillis Ga — to. g lu teu s accessorius Gb — w . glu teo bic eps G b’ — to. g lu teo b icep s (antei'ior part) G b” — to. g lu teo b ic ep s (m iddle part)

G b’” — to. g lu teob icep s (posterior part) Ge — to. g e m e llu s Gm — to. g lu te u s m ed iu s Gp — to. g lu te u s pro fu n d u s Gs — m. g a stro cn em iu s (caput la terale) G s’ — to. g a stro c n em iu s (caput m e d i a le ) I — m. in fra sp in a tu s la — m . iliacus (caput la te ra le ) la ’ — to. iliacus (caput m ed ia le) Ic — w . in terco sta lis e x te r n u s II — to. iliocostalis Im — to. in tero ss eu s m e d iu s In — to. in terd ig a str icu s Is — to. in te r spinalis It, I t ’, It” — to. in t e rtra n sv e rs a riu s J — m . stern o h y o id e u s La — m. lo ngissim us atla n tis Lc — to. leo a fo r costae Ld — to. la tis sim u s d o rsi L g — to. longus colli Li — m . lon gissim us d o rsi Lie — m . longus capitis LI — to. le v a to r labii m a x illa ris p ro p r iu s Lp — to. lon gissim u s cap itis Lr — to. lo n gissim u s c ervic is L t — to. longus a tla n tis M, M’ — to, m a s se ie r Ma — m. m a la ris Me — to. m u l ti fi d u s c ervicis Md — to. m u l ti f i d u s d o rsi N1 — m . l e v a t o r nasolabialis Oe — m. ob liqu u s a b do m in is e x te r n u s Oi — to. o bliq u u s a b do m in is in tern u s Om — to. o m o h y o id e u s Or — to. ob liq u u s ca p itis cranialis Ot — to. o m o tra n sv e r sa riu s Ou — to. ob liqu u s ca pitis caudalis P — to. p e c tin e u s P i — to. p ir ifo r m is Pj — p soas m a jo r P m — m, p soas m inor P o — to. p o p lite u s P p — to. p ecto ra lis p ro fu n d u s P s — to. p e cto r a lis su perficia lis P tl — to. p te r y g o id e u s lateralis P tm — to. p te r y g o id e u s m ed ia lis (su p erficial layer) P tm ’ — to. p te r y g o id e u s m ed ia lis (deep layer)

Q f — m. q u a d r a tu s fe m o ris Q1 — w . q u a d ra tu s lu m b oru m Q uf — m. q u a d rice p s fem o ris R — to. r h o m b o id e u s (pars ih orttad iaf R ’ — to. rh o m b o id e u s (p. cervim W i) Ra — to. re c tu s ab d o m in is R c — to. re tr a c to r costae R j — w. re c tu s ca p itis dorsalis tun ¡in R1 — to. r e c tu s cap itis latere lis : R t — w . r e c tu s fem o ris S — m. su p ra sp in a tu s Sa — to. sartoriu s Sb — to. s e m im e m b r a n o s u s Se — to. sp in a lis cervicis Sd, S d ’ — to. spinalis e l s(’.mií¡p¡níihh d orsi (pars la te ra lis ef iñádialíñi Se — to. ser ra tu s dorsalis (pars ca.udalis) S f — m. soleus S i — to. se rra tu s dorsalis (pars cranialis) SI, s r , SI”, SI”’ — to, sp ienins Sm — to. s em isp in a lis c a p ia s S n — to. se m ile n d in o su s So — to. sem isp in a lis lamlm«»?»* Sp — to, sca len u s prim n c amUt? (p. dorsalis) S p v — to. sca len u s prim er (p. v e n tr a lis) Sr — to. sternoTOasioidcK/f Ss — to. scalenus supruroshdib S t — to. stern om an d ib alu ris S u ,S u ’, S u ” — m. snbsropHiüí S v, S v ’ — to. serra l us i >c a ! ¡ni (p. th o raca lis el cerU/icidíMÍ • T, T ’ — to. te m p o r a l is Ta — to. teres m a jo r T b — m . tibia lis anterior ' Tc — to. t r a n sv e rs a s emu »»o*« T f — to. ten so r fascial? Th — to. tr a p e ziu s (jr. i h o i i j r v i y i ;■ Ti — to. te r e s m in o r ' : T1 — to. te n s o r fascia»* laid*. • ■■ . '• Tp — to. tib ia lis paid «ribí ' .' y.- ' Tr, T r’, Tr”, T r’” — ith i f № № Ts — to. transversuil T t — to. tr a n s v e r a s .• / Tu — to. tr a p e z iu s |Í Í S I Í Í iÍ i ! |í VI — w . v a s tu s later id U V m — to. v a s tu s mviitnH« Z — to. zy g o m a tic a s

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