they - Brown Universitycs.brown.edu/courses/cs137/2013/readings/Humphry1869.pdf · THE MYOLOGY OF THE LIMBS OF PTEkOPUS. dition in someof the loweranimals, are, in this respect, alike-are

THE MYOLOGY OF THE LIMBS OF PTEROPUS. ByPROFESSOR HUMPHRY (Pl VI. & VII.)

THE muscles of the limbs of Cheiroptera present so manyfeatures of interest that I think it worth while, although theyhave been described by others, to give an accurate descriptionof them as recently dissected by me in two specimens of Ptero-pus Edwardsii, a male and a female.

The peculiar features of interest are, first, the modificationsby which the muscles of the fore limb are adapted for flying,leading us to observe by what slight deviations from the ordi-nary mammalian type the end has been attained. The largepectorals springing from a keeled sternum, and the tensoresplice alaris constitute some approach to the peculiarities of thebird. The latter muscles, however, do not spring, as in the bird,from the shoulder girdle, but are obviously prolongations of theplatysma, and no part of them passes to the forearm; nor doesthe deeper stratum of the pectorals (pect. min. and subelavius)show any special avian characters. Moreover the resistance tothe pull of the pectoral muscle upon the shoulder is affordednot by the presence of a coracoid bone or by the union of theclavicles, but the manner in which the broad flat surfaces of thestrong clavicles are jointed with corresponding articular surfaceson the sternum. The biceps has, in one of the specimens, aslight muscular slip from the humerus; but this is not un-common in mammals or indeed in man, and it is rather a detach-ment from the brachialis anticus than a distinct origin of themuscle, as it is in the bird, from the upper end of the humerus;and unlike the same in the bird, the main (coracoid) portionof the muscle is in two parts, and these are unusually distinct.The supinator longus stops short at the upper third of the radius,acting simply as a flexor of the forearm; and the radial flexorsand extensors of the carpus are inserted as in mammals, noneof these muscles being attached, as their two representatives inbirds are, to the foremost metacarpal.

The flexor carpi ulna-is arises as usual. It does not, as in the

THE MYOLOGY OF THE LIMBS OP PTEROPUS.

bird, arise behind the inner condyle and pass beneath it, playingunder it as under a pulley. This provision in the bird forharmonizing flexion of the carpus with flexion of the elbow,like the provisions in the supinator longus (its connection withthe tensor plicae alaris above and with the metatarsus below) forharmonising extension of the carpus with extension of the elbow,is wanting in Pteropus.

In these and in the other myological differences from birdswe recognize an inferiority in the adaptation of the animal forflight which is more strongly evinced in the want of concentra-tion and simplification of the osseous and muscular structurescharacterizing the distal segment of the bird's wing, and which ismost of all shown in the dermal extension as a substitute for fea-thers, whereby the hinder limbs are involved in the work of aerialmovement and are unfitted for any other mode of locomotion'.Thus, while we admire the manner in which this order (Chei-ropterous) is specialized for flight from a class (Mammalian)which is not constructed for such movement, we are conscious ofthe great superiority of the mechanism attained in the severalmembers of that class which is throughout, or in great measure,constructed for flight, and in which the hind limbs are left freeto subserve to terrestrial locomotion, while better motor powerin air is obtained by the more perfect adaptation of the forelimbs and of the body generally. A similar illustration of thesame principle is found in the fact that the mammalian inhabi-tants of the deep show their adaptation to the watery mediumto be inferior to that of the fish, by the necessity which compelsthem to come to the surface for the purpose of breathing air.

Secondly. It is interesting to find that though the hindlimbs are unfitted for the performance of the locomotory func-tions, they, nevertheless, retain in the main the structural com-position and disposition of parts usual in those limbs. The func-tions of the hind limb in Cheiroptera are almost exclusivelythose ordinarily assigned to the fore limb; yet they are effectedby a modification of the ordinary structure of the hind limbrather than by a substitution of that of the fore limb. The

1 The reduction of the number of the phalanges in the fore limb to two ineach of the three ulnar digits has relation no doubt to their great length andattenuation, which is incompatible with the additional source of weakness thatwould have been entailed by the usual series of bones and joints.

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peculiarities, that is to say, of the hind limb, or most of them,are retained. The femur is jointed with one leg bone, althoughthe rotary movement of the leg upon the thigh is lost. Thetarsus is articulated with the tibia, by one bone-the astraga-lus-; the scaphoid intervenes between that bone and the distalrow of tarsals, and the os calcis stands up towards the middle ofthe flexor aspect of the limb to receive the tendo Achillis.Some approximation, however, to the osseous conformation ofthe fore limb is found in the fact that the os calcis is situatedchiefly on the fibular side of the ankle, rises to a level with theastragalus, and subtends the lower end of the fibula, remindingus of the relations of the cuneiform to the scapho-lunar and tothe ulna. Moreover the small size of the tarsus in relation tothat of the metatarsus and phalanges are reminders of theordinary relations of the carpus to the metacarpus and digits;and the foot is in the same plane with the leg. With regard tothe muscles, the gastrocnemius is derived from both the condyles,and the peroneus longus, in spite of the absence of the upperpart of the fibula, is present, and takes its usual course; andthe various muscles of the foot and leg present the usual charac-ters of the hind limb, except that the flexors of the leg run alltogether to its middle instead of passing, in two divisions, toits more lateral aspects.

Thirdly. There is, however, an assumption by the hindlimb of one character of the fore limbs, which renders theanatomy of these animals peculiarly instructive and valuable ininstituting a homological comparison between the two limbs.I mean the direction of the flexor and extensor surfaces of thelimb, which, instead of being, as it usually is, in the thigh andleg at any rate, the reverse of that in the fore limb, is, in Chei-roptera, very nearly the same with that of the fore limb. Thus,the extensor surface of the thigh and leg and the prominence ofthe knee, like the extensor surface of the arm and forearm, andthe prominence of the elbow, are directed backwards, and thehallux and tibial line, like the polled and radial line, are onthe outer side.

In another place (Essay on the Limbs of Vertebrate Animals)I have shown that the fore and hind limbs, in their primarytransitory condition in the fetus and in their persistent con-

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dition in some of the lower animals, are, in this respect, alike-are both, that is, directed in the same manner. They are bothprojected at right angles and nearly straight, from the trunkwith the dorsal or extensor surfaces superior, and the polledand hallux with the respective radial and tibial margins an-terior; and the flexion at the elbow and knee, in both, takesplace upwards, away, that is, from the abdominal aspect of theembryo. Thus far the correspondence between the two limbsis pretty exact; and if things remained in this position therewould be little difficulty in pointing out the serial homological re-lations of the several parts. But, while the flexion at the elbowand knee takes place, a rotation is going on in the two limbs andin opposite directions. The elbow with the superior or extensorsurface of the fore limb becomes turned backwards, and theknee with the superior or extensor surface of the hind limb isturned forwards. This causes the hallux, tibia, tibial condyleand side of the femur, that is, the line of the hind limb whichwas originally anterior, to be directed inwards, and the pollex,radius, radial condyle and side of the humerus, that is, the lineof fore limb which was also originally anterior, to be directedoutwards. Rather, I should say, with regard to the fore limb,this would have been the effect, but that a rotation in an oppo-site direction to that in the rest of the member takes placein the middle and distal segments (the forearm and hand).This causes the polled and lower part of the radius to bedirected inwards while the upper part of the radius and theradial line of the upper part of the limb retain the directionoutwards which I have indicated. This opposite rotation ofthe two limbs has, I need not say, proved a source of muchperplexity in their comparison and led to many errors.

In Cheiroptera the rotation of the hind limbs differs fromthat which usually takes place in those limbs, and correspondswith that which takes place in the fore limbs. The convexityof the knee and the extensor surface of the limb are directedbackwards; and the hallux, tibia, and tibial line of the femurare directed outwards: they afford us, therefore, a peculiarlyfavourable opportunity for comparing the muscles in the twolimbs, and for settling some of the disputed points. They show,for instance, unmistakeably, that the extensor muscles (triceps)

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on the arm correspond with the extensor muscles (quadriceps) onthe thigh, and that the flexors in the two limbs are in similarcorresponding relation. They give additional support to theview that the inner, or tibial, condyle of the femur correspondswith the outer, or radial, condyle of the humerus, that thetibial (usually inner, in these outer) trochanter of the femur andthe muscles passing to it are homologous with the radial (outer)tubercle of the humerus and its muscles, and consequently,that the muscles on the outer surface of the scapula are, to someextent, homologous with those on the inner surface of the ilium.Still, the rotation of the hind limb being the reverse of that whichis usual, has so far altered the parts from their ordinary disposi-tion that it is not, in every instance, easy to refer a given muscleto its correspondent in the hind limb of other animals. It isscarcely necessary to add that caution is, therefore, needed indrawing inferences with regard to serial homologies with thefore limb.

These animals show that the direction of the rotation ineach limb has no close relation to the particular structure of thelimb; for in them, the rotation of a fore limb is associated withthe ordinary structure of a hind limb. We must, therefore,look to some other agency as the determining cause of thedirection of the rotation. At present I can discern no othercause than that mysterious agency in development, whatever itis, which determines the general conformation and the detailedstructure of a limb in accordance with the part it will have toplay in the economy of the animal.

Nor do I see that the illustrations thus furnished of themodification of one order in a class to suit, to some extent, thehabits of another class and the modification of one pair of limbsto fulfil, to s6me extent, the ordinary functions of the otherpair, aid us much in the elucidation of the great questions ofthe mode of derivation of species. In each instance the changeseems to be wrought out suddenly and completely, per saltumas it were. At least the known connecting links between theseaerial and the ordinary terrestrial mammals are too few toenable us to draw any definite inferences from them.

Some of the points of interest which suggest themselves Iwill refer to in the course of the description.

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CUTANEOUS MUSCLES

are disposed as follows. They are remarkable for their extent,their disposition, and their osseous connections.

The Platysmc portion consists of four muscular bands (1, 2,3, 4 in Pl. vi.),

(1) arising from the side of the lower jaw, where some ofthe fibres are blended with those of the muscles of the angle ofthe mouth', and from the side of the head behind the lowerjaw. It is the 'cervico-facien' of Cuvier and Laurillard2.

(2) from the hinder part of the parietal crest, from theoccipital crest, and slightly from the middle line beneath thelatter covering, in part, the space usually occupied by the cer-vical portion of the trapezium. It is the 'dorso-occipitien' ofCuvier, who regards this as the main portion, the other threebeing accessories.

(3) from the middle line of the lower part of the neck, infront of the trachea, where it is confluent with the correspond-ing muscle of the opposite side.

(4) from the fore part of the keel of the sternum, super-ficial to the pectorals major. These several bands, distinctfrom one another at their origins, converge to the free edge ofthe humero-radial expansion of skin, along which they run,forming a narrow band (5) (corresponding with the tensor policealaris in the bird8) as far as the polled into the side of the pha-langes of which it is inserted. In the middle of the edge ofthe wing-fold the muscular fibres are supplanted by a thinband of highly elastic tissue. This again, one-third from thepolled, gives way to muscular fibres which, after about an inch,terminate in common tendon. Thus there is first muscle, thenelastic tissue, then muscle, and, lastly, tendon.

The muscle thus disposed serves to give tension to the fold of thewing, bends the forearm upon the arm and the polled upon the fore-arm, also abducts the polled and so may aid in separating the digitsand expanding the wing, besides exerting some influence upon thehumerus.

1 This reminds us of the corresponding part of the muscle in Orycteropus(Vol. ii. of this Journal, p. 293) which passes from the depressor anguli oris uponthe arm and forearm.

2 Anatomie Compar6e, Planches.S This muscle in the bird is derived from the deltoid or pectoral or both.

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Another cutaneous muscle (cutaneo-pubic) arises from thefore part of the pelvis and radiates over the abdomen, thefront of the pectoral muscle, and the shoulder. A lateral band,taking a direction different from the rest of the muscle, passesinto the wing on the outer side of the thigh and leg, and runsquite to the edge of the wing. A third muscle (P1. Vi. F. C.,femoro-cutaneous) arises from the lower part of the tibial (orouter) side of the shaft of the femur and passes forwards ra-diating over the back.

A fourth (I. C., ischio-cutaneous) runs from the ischium,across the ischio-pedal fold of skin and the supplementary cal-caneal bone, upon the dorsum of the foot.

A fifth (C. C., coraco-cutaneous) arises from the coracoid incompany with the biceps muscle, leaves the latter in the upperthird of the arm, and runs through the brachio-crural flap ofthe wing to its lower edge'. Neither of these are mentionedor represented by C. and L.

Other similar bands of muscle, parallel with this last, de-scend, at nearly regular intervals, through the brachio-cruralwing-flap to its lower edge from near the arm and forearm,though they do not appear to have any direct connection withthe arm or forearm; and there is a broad thin muscular sheetextending from the back of the thorax and abdomen, near thespine, into this flap, and sending off bands which radiate invarious directions in the folds of the skin.

MUSCLES OF THE FORE LIMB.

Pectoral in three portions:(1) Sternal portion (Pect.), of great size and thickness,

arises from the. side of the keel of the sternum and the sternalends of the sternal ribs, slightly from the lateral portion of the

1 In Echidna setosa and in the Jerboa a thick band of muscular fibresarises from the anterior tubercle of the humerus, close to insertion of thepectoral muscle, and runs into the pannicle on the side of the abdomen. Partof the pannicle is also connected, in the Echidna, with the edge of the ulna; andin the Jerboa it is, as in Pteropus, attached to the face and side and back ofthe head, and by a distinct slip to the sternum. In it, as well as in Pteropus,the latissimus dorsi is nearly covered by this part of the pannicle, and is, as itwere, stunted by it, being very small. In the Mole and the Rat, also, the pan-nicle is connected with the sternum, the humerus and the ulna. It is rarely,however, attached to the coracoid.

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manubrium sterni but not at all from the clavicle, to the ridgeon the outer side of the bicipital groove of the humerus.

(2) Clavicular portion, from the deeper part of the innerthird of the clavicle, lies beneath the preceding, and beneaththe deltoid, and might perhaps be regarded as belonging toeither. It is inserted into the upper part of the same ridge inclose relation to the fibres of the pectoral and of the deltoid.It is the 'petit pectoral' of C. and L., but may, probably, bemore correctly regarded as a part of the pect. major. It doesnot arise, in either of my specimens, as represented by Cuvier,from the rib, but is connected only with the clavicle. It hasnot the ordinary characters of the clavicular part of the pectoralismajor; for this usually lies more superficially, and crosses infront of the rest of the muscle to a lower part of the pectoralcrest of the humerus.

(3) Abdominal portion arises from the pubes in companywith the rectus abdominis, forming the outer part of the com-pound muscle, separates from it as it ascends to the ribs, passesbeneath the sternal portion of the pectoral, between it and thebiceps, to the upper and inner margin of the same ridge justbeneath the preceding.

It is called ' portion ventrale' by C. and L. It' might be saidthat the rectus divides as it ascends and thus forms its outer de-tachment. In many animals (Fawn, Pig, Calf) a portion of thepectoralis arises from the sheath of the rectus or the linea alba.In Batrachians it is, as in this instance, blended with the muscleitself. The relation of this portion (which may probably be re-garded as therepresentative of the Pectoralis minor), and of thesubelavius in Pteropus as well as of the scaleni, in some animals, tothe rectus suggest these may all belong to one series or one muscularplane commencing at the pubes, terminating in the cervical vertebraeand diverging to the shoulder, sometimes continuous, but more com-monly interrupted by one or more ribs'.

1 The Pyramidalis is large, has a broad origin from the front of the pubes,and covers a considerable portion of the lower and fore part of the abdomen.The fibres ascend with an inclination inwards, and are chiefly inserted into thelinea alba. Some however run on to the sternum and adjacent cartilages of theribs.

In the female Pteropus, the Pyramidalis arises from the pubes on the sideof the pubic gap, just in front of the rectus. The fibres pass obliquely inwardsto the line alba. The lowest are almost transverse, parallel with and quiteclose to the symphysial ligament, so as completely to cover all the part abovethe ligament in the interval between the recti. The upper fibres are moreoblique.

Rectus is attached above to the ribs wide of the sternum, as high as the first rib,

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Meckel's (Vergleich. Anat. iII. 454) description of the relation ofthe pectoral with the rectus in Vespertilio corresponds with theabove. This anatomist believes that in cases where the pectoralisminor appears to be absent it is really blended with the scalenus(s. 490) and says the same is also occasionally the case with thesubelavius (s. 416).

Subelavius (Subel.) is large. It passes from the under sur-face and anterior edge of the broad sternal portion of the firstrib (the rectus abdominis is inserted into the posterior edge),along the lower surface of the clavicle, to be inserted beneathits outer part. It is inserted exclusively into the clavicle.

Serratus magnus anticus is in two portions, as Meckel de-scribes. The upper, thick and muscular, arises from the first ribbehind the subclavius and beneath the tendinous insertion ofthe scalenus and also slightly from the second rib, and is in-serted into the upper part of the hinder margin of the scapula,internal to the insertion of the levator scapulae, but not extend-ing so far down the scapula as it does. It is quite distinctfrom the levator scapule in its whole extent. The lower por-tion is broader, arises from the eight ribs below the first andis inserted into the inferior margin of the scapula between theteres major and the subscapularis.

It is nearly the same in the two-toed Anteater, but the lowerportion is in it attached more to the angle and hinder margin of thescapula than in Pteropus and does not advance so far forward betweenthe teres and subscapularis. This muscle undergoes subdivision inBirds, Reptiles and Amphibia, especially in the Frog, with varietiesof insertion. Rudinger, Die Muekeln der vorderen Extremiatten derReptilien und Vogel.

is joined near its upper part by the pectoral as above stated, and descends behindthe pyramidalis to the pubes. There are no tendinous intersections in it or inthe pyramidalis. In the female the recti separate as they descend to the pubicbones, leaving the lower part of the abdomen uncovered by them.

Obliquus Externus, indigitating above with serratus, forms a thin expansion,covering the rectus and pyramidalis. Below, in the male Pteropus, it has a freeedge some distance above the pubes; and beneath this edge the large spermaticcord escapes, carrying with it some fibres of the muscle beneath, the obliquusexternus, which form a crernaster and are disposed nearly transversely uponthe cord.

In the female, where the pubic bones are widely apart and connected by afibrous band passing across in front of the vagina, the lower margin of the obl.ext. is less defined. Its expanded tendon passes to the pubes and to the sym-physial band.

The obliquus interns passes entirely behind the rectus, no part of its tendongoing in front. At the lowest part of the abdomen it is very thin.

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Biceps (B.) has two distinct heads, one from the summit ofthe long coracoid process, the other, larger, from the retiringangle at the root of that process and the ridge between thecoracoid and the margin of the glenoid surface. It cannot,therefore, be said to arise from the glenoid part of the scapulaat all. Neither does it pass through the shoulder-joint, but liesin a bursal canal, separate from and over the joint. It thentraverses the bicipital grove. The two heads come into con-tact in the middle of the arm still preserving their distinctness,though the smaller portion (that from the extremity of thecoracoid) derives some fibres from the other larger and longerportion. They are inserted on the inner or ulnar side of theradius, the smaller portion a little in front of the other. Abursa intervenes between their tendons and the bone.

The smaller, or shorter, or coracoid portion of the biceps is notunfrequently inserted in front of the other. It is so in man and in aGreen Monkey now before me. It is the "part also chiefly connectedwith the fascia ofthe forearm in man.

Brcachialis Anticus (Br. a.) small, arises by muscular fibresfrom the inner side of the shaft of the humerus just beneaththe coraco-brachialis. Its tendon, which is long and thin,travels, in company with those of the biceps, to the internallateral ligament'. There it separates from them, passing onthe inner side of the ligament to the fore part of the ulnawhere it is inserted. In the female some fibres of this muscle-not a distinct muscle, but fibres arising in common with thebrachialis anticus-join the smaller portion of the biceps.

The distinctness of these three flexors and particularly of thetwo portions of the biceps is remarkable, especially as the latterarise so near together. The small size of the brachialis anticus hasrelation to the small size of the ulna. Meckel describes the biceps asarising in the Bat from the humerus and the coracoid. So far as myobservation has gone in the human subject portions of the brachialisanticus which join the biceps join its coracoid portion and terminatechiefly, if not entirely, in its fascial expansion.

Coraco-brachialis (C. b.), from the extremity of the coracoidprocess and the upper part of the tendon of the biceps, which

1 This ligament is long and very strong. It passes from the lower and forepart of the internal condyle, between the tendons of the biceps and the brachi-alis anticus, to the ulnar side of the radius, a little below the joint.

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arises from the same point, to along the inner side of the hu-merus between the lesser tubercle and the origin of the bra-chialis anticus. The portion arising from the tendon of thebiceps can easily be separated from the remainder, there beinga stratum of areolar tissue between them; and it is insertedhigher than the remainder of the muscle. A few fibres of thebrachialis anticus arise from the lowermost part of its tendon ofinsertion in front.

Trapezius (Trap.). The cervical and occipital portions areabsent'. The upper edge is very defined, and extends acrossbetween the shoulders. The remainder of its origin is asusual. It is attached to the back of the acromion and theouter fifth of the clavicle.

Cervico-humeral (C. h.) (omo- ou acromio-trachdlien), fromthe second and third transverse cervical spines to about a fifthof the clavicle immediately internal to the trapezium. In itsorigin this muscle is a continuation of the levator scapulae inits insertion of the trapezium. I am disposed to agree withMeckel (1. c. 481) in assigning it to the latter muscle ratherthan to the levator scapulav.

Levator scapulce (Lev. so.) is large from the cervical transversespines, beneath the cervico-humeral, to the edge of the scapulabetween the spine and the superior angle.

Rhomboideus, one strong muscle from the upper dorsal spinesto the spine and edge beneath it of the scapula. There is nocervical or occipital portion.

Latissimus dorsi (Lat. d.) is narrow, having no attachmentto the ribs. It arises from two or three anterior lumbar andthree hinder dorsal spines, beneath the trapezium, overlaps, buthas no attachment to, the angle of the scapula', and is insertedby a narrow tendon into the inner tubercle of the humerus.

Deltoid (D.), from the outer half of the clavicle, extending,that is, for some distance internal to the region of attachmentof the trapezius and cervico-humeral, and from the acromion.The fibres of the clavicular portion take nearly the same di-rection as those of the clavicular portion of the pectoral muscle,

1 Cuvier suggests that the deficient cervical and occipital portions may berepresented by the ' dorso-occipitien.' Vide supra.

2 The angle (A. sc.) of the scapula in P1. vii. has been raised up from beneaththe cover of the trapezium.

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and are partly blended with theni at their insertion1. Theacromial portion, on the contrary, which is separate from theclavicular, meets the pectoral muscle and is inserted, opposite toit, into the same ridge. The fibres of the deltoid are insertedinto the same length of the bicipital ridge with the pectoral.The hindmost fibres of the deltoid, arising from the outer partof the acromion, join the outer part of the triceps and becomeblended with it.

Teres Major as usual. It has a broad musculo-tendinoustermination, which can scarcely be said to be inserted with thelatissimus dorsi, but into the inner bicipital ridge beneath it.

Teres Minor (T. mn.), large, from a considerable part of thehinder edge of the spine as well as from the lower costa of thescapula to the outer part of the great tubercle of the humerus.It nearly covers the Infra-spinatus (Inf sp.), which is small.

It is represented by Cuvier as a part of the deltoid; and certainlyit has the usual origin of part of the deltoid. Yet it is quite distinctfrom the deltoid and crosses it nearly at a right angle, running be-neath it to the tubercle of the humerus. Meckel considers that it iswanting, and, I conclude, takes it for the infra-spinatus which he saysis large. But that muscle is distinct, lying beneath the teres, and ofrather small size.

Supra-spinatus is situated as usual.Subscapularis presents nothing peculiar. It is thick, and

arises from the whole of the under surface of the scapula, passesover the shoulder-joint quite free from the capsule, and is in-serted into the inner tubercle of the humerus.

Triceps (Tr.). The long head has a broad musculo-tendinousorigin from the scapula behind the glenoid cavity. The re-mainder of the muscle arises from the posterior surface of theupper half of the humerus quite up to the shoulder-joint, de-riving, on the outer side, fibres arising with those of the deltoidfrom the hindmost part of the acromion; and, internally, itsfibres arise from the posterior surface of the tendon of insertionof the coraco-brachialis. The long, strong, well-defined tendonincludes a sesamoid bone (s), the homologue of the patella, as itpasses over the trochlea to the projecting edge of the ulna,

Pronator teres (Pr. t.) has no connection with the ulna.Arising from the inner condyle in conjunction with the four

1 Meckel finds it difficult to decide whether this clavicular portion belongs tothe pectoral or the deltoid.

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following muscles, it passes across to the oblique line on thepalmar surface of the upper third of the radius 1

Flexor carpi radials (Fl. c. r.) from the inner condyle to theulnar side of the base of the metacarpal of index.

Flexor carpi ulnaris (Fl. c. u.) from the ulna as well as fromthe inner condyle to the distal margin of the transverse processof the os magnum', opposite the interval between the third andfourth metacarpals.

Flexor digitorum sublimis (Fl. d. s.)5 from the condyle andthe upper part of the radius. As it passes through a tendinoussheath in front of the carpus it divides into two tendons, ofwhich one passes to the sesamoid bone on the ulnar side of themetacarpo-phalangeal joint of the polled, and the other, havinga slight attachment to the metacarpal bone of index, passes onto be attached to the base of the second phalanx of index.

Flexor digitorum profundus (Ft. d. p.) arises from the innercondyle together with the preceding, but soon becomes distinctfrom it, and situated beneath and to the ulnar side of it. Itarises, in addition, from the anterior surface of the ulna andfrom the contiguous margin of the surface of the radius, passes.over the carpus in a pulley beneath the preceding and beneaththe process of the os magnum, and divides into two tendons.Of these, one passes between the bellies of the short flexorpollicis and the sesamoid bones to the ungual phalanx ofpollex. The other again divides into two, which pass to theterminal phalanges of the index and middle digits. Theseslender tendons are accompanied, just below the carpus, bysome muscular fibres, like lumbricales, which seem to terminatein as well as to arise from the tendons. In the female Pteropusthe tendon to the third digit is quite interrupted by the mus-

1 Cuvier says there is an absence of all pronators and supinators. Meckelfinds a long pronator and a short supinator. I find in addition the long supina-tor very distinct, but no short pronator.

2 The os magnum is the largest in the distal row of carpal bones; and itspalmar surface is expanded into a broad process extending tranversely across thebases of metacarpals ii, iiI, Iv, and reaching partly across metacarpal I. Thereis no distinct hooklike process to the unciform bone. The trapezoid and tra-pezium are of moderate size. The proximal part of the carpus is formed almostentirely by the thick transversely elongated scapho-lunar bone, in which a slightoblique groove on the palmar surface indicates the distinction between its com-ponent parts. The cuneiform is very small and articulated with the distal surfaceof the lunar. I do not find any pisiform.

3 Meckel describes one flexor muscle in the bat dividing to each of the fin-gers as well as to the thumb.

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cular belly. That to the second digit retains its continuity,the muscular fibres being superadded.

Interossei are well marked and regular, two at least for eachdigit passing in pairs along each metacarpal and attached tothe bases of the several first phalanges constituting 'interossealflexors.' In the case of digit v, they arise from the trans-verse process of the os magnum; and, in addition to thetwo interosseal flexors (Ft. b.), one, separate from and ulnad ofthem, becomes attached to the palniar surface of the meta-carpal about one quarter from the carpus, and may representan 'interosseal abductor' (Abd.), while a slip from the in-terosseal flexor on the radial side extends beyond the rest,passes along the radial side of the first phalanx and joins theextensor tendon, so that it may be regarded as an 'interossealadductor.' The interossei of digits I, II, and Iv. arise from theproximal ends of the metacarpals and the contiguous carpalbones, therefore on a plane deeper than those of digit v. Theflexor tendons of digits II. and III. run between the interossealtendons; all the interossei are free in the whole, or nearly thewhole, of their length from the metacarpals, and are roundmuscular bellies like lumbricales. In the case of the pollex, ashort, thick, muscular belly descends from the carpus, on eitherside of the metacarpal, to the sesamoid bones and the adjacentbase of the first phalanx, constituting interosseal, or short,flexors, as in the case of the other digits; and an additionalmuscle arising from the os magnum terminates in a thin tendon,which passes along the ulnar side of the first phalanx to itsdorsum, and is inserted into the back part of the terminalphalanx, constituting the 'interosseal adductor,' and provingthe sole extensor of that phalanx.

The interossei constitute the only flexors of digits iv. and v,and extend only to their first phalanges; the flexion of the secondor terminal phalanges being, apparently, effected to the requiredextent by the muscular tissue in the covering of the wing.The above muscles, with the exception of the fl. c. u. and the inter-

ossei, have the effect, in addition to their other purposes, of flexingthe metacarpals upon the carpus; and these are so articulated thatthey are at the same time inclined towards the ulnar side andapproximated to one another, that is to say, they are folded togetheras they are bent upon the forearm. The metacarpals indeed arearticulated to the carpus Di such a manner as to permit the opposite

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movements of flexion with inclination ulnad or folding together, andof extension with inclination radiad or separation from one another.

Supinator longus (Sup. 1.), from the outer surface of thehumerus above the condyle to the outer surface of the radiusjust external to the insertion of the pronator radii teres. Is aflexor of the forearm.

Supinator brevis (Sup. br.), from the outer condyle and thetendons of the extensors of the digits to the oblique ridge onthe front of the radius to which the pronator teres is attached.Is an extensor of the forearm.

Extensores Carpi radiates longior et brevior (Ex. c. r. 1. et. br.),from the outer condyle together. The longior, the anterior andrather the smaller, passes to the process on the back of the proxi-mal end of the second metacarpal bone. The brevior passes tothe process on the radial side of the end of the third metacarpal.

In the male Pteropus a slip passes from this tendon to the fourthand fifth metacarpals. I found a similar slip to the fourth metacarpalin the Chimpanzee.

These muscles extend and abduct the metacarpals on the carpusas well as extend the carpus.

Extensor Carpi u1naris (Ex. a. u.), chiefly from the outersurface of the ulna and slightly from a fascia connected withthe outer condyle to the ulnar surface of metacarpal v. justbelow the carpus.

Extensor digitorum (Ex. dig.), from the fascia connected withthe outer condyle and the olecranon, and divides into three sepa-rate tendons, which pass to the terminal phalanges of the threeulnar digits. They lie on the ulnar sides of the metacarpals,pass over the dorsal surfaces of the metacarpo-phalangeal joints,expand there, become connected with the joints, and narrowslips pass on to the terminal phalanges. The tendon to digit v.runs in a separate sheath, and is joined by a slip from that todigit IV.

Extensor primus (ossis metacarpi) pollicis (Ext. poll. pr.),from the radius and ulna high up, runs over the edge of thelower end of radius above the base of the pollex, having here asmall ossicle in it, to the radial extremity of the transverse pro-cess of the os magnum.

Secundus (Ext.poll. see.), just beneath the preceding, througha groove on the ulnar side of the end of the radius, in company

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with the succeeding muscle to the base of the dorsum of thefirst phalanx of polled. The 'interosseal adductor' is theextensor of the third or terminal bone of the pollex.

Extensor indices, beneath preceding, and in company with itto the terminal phalanx of index'.

Thus each of the terminal phalanges of the digits is suppliedwith an extensor, though those of iv. and v. are devoid of flexors.

The differences from the dorsal muscles of the human forearmand hand are in the insertion of the supinator and in the extensorespollicis.

MUSCLES OF THE HIND-LIMB.

Psoas parvus (Ps. p.), from the two or three lower dorsal andthe three upper lumbar vertebrae, a small muscle. Its slendertendon is inserted into the outer extremity of the projectingspine of the pubes which overhangs the hip-joint2. Some ofits fibres are continued on into the pectineus.

Psoas magnus (Ps. in.), from the bodies of all the lumbarvertebrae, lying external to the preceding, as well as from thefront of the ilium and the adjacent surface of the sacrum, passesbeneath the overhanging pubic spine, external to and behindthe hip, to the outer or tibial trochanter.

Iliacus internus (Ii. i.), from the outer surface of the iliumclose to the crest, has no further connection with the ilium, isdistinct from the psoas, and passes beneath it to the outer sur-face of the femur quite beneath the tibial trochanter. Itcrosses external to the extensor cruris.

Sartorius, not clearly distinguishable. Cuvier describes un-der this name some fibres passing from the ilium to the lateralpart of the pannicle. Meckel says it is absent in cheiroptera.

Pectineus (Pect.), a round muscle in its upper part, arises bya strong well-defended tendon from the most projecting pointof the pubic crest just beneath the insertion of the psoas par-vus, and from the tendon of that muscle. Becoming widerand flatter it crosses anterior and external to the psoas andiliacus, and is inserted, beneath the latter muscle, into the

1 Meckel does not mention this muscle, but describes the ext. dig. as passingto the four fingers.

2 The mistake in the Leqons respecting the absence of this muscle as well asof the psoas magnus and the iliacus is rectified in the Anatomie Comnparie,Planches of Cuvier and Laurillard.

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310 PROFESSOR HUMPHRY.

middle of the outer (tibial) side of the femur. It is the ' pec-tineus' of Cuvier and Meckel; still it is a question whether itmay not be the adductor longus of which there is no other re-presentative; or it may be the representative of both thesemuscles.

Adductor brevis, from the outer surface of the pubes undercover of the gracilis, between it and the obturator externus,crosses the psoas to the fore and outer surface of the femurimmediately in front of the insertion of the iliacus.

Adductor magnus. There is a considerable mass of muscle,consisting of three or more parts, arising from the outer surfaceof the pubes and ischium, appearing superficially between thegracilis and the semi-tendinosus and semi-membranosus, andextending beneath the latter muscles to the middle line. It isinserted along the fore part of the femur from the outer, ortibial, trochanter three fourths down the bone. An upper por-tion, easily separated, may represent the Quadratus femoris,and another portion, separated in great measure from the restby the sciatic nerve and the origins of the semi-tendinosus andsemi-membranosus and connected with its fellow in the middleline and inserted lower and rather more on the fibular aspectof the femur than the remainder of the muscle, may possiblyrepresent the Biceps which, owing to the absence of the fibulain the upper part of the leg, may have stopped short in thethigh and acquired an attachment to the femur'.

Gluteus Maximus (GI.) arises, muscular, from the hindermargin of the crest of the ilium and from the sacral spines, andis inserted, tendinous, into the upper half of the inner, or fibu-lar, side of the shaft of the femur between the adductor massand the vastus externals which is here internal.

Glutamus Medius arises from the outer and hinder surface ofthe ilium, under cover of the gluteus maximus and iliacus, andis inserted, tendinous, into the summit of the inner, or fibular,trochanter. There is no glutwacs minimus distinct from themedium.

Obturator Externus (Obt. e.) is a large muscle arising fromthe whole of the exterior of the obturator ligament and the

1 Cuvier and Meckel mention only one adductor besides the pectineus andmake no mention of a quadratus.


surrounding margin of bone. It passes immediately over thefront of the hip-joint and is inserted into the outer part of thetibial trochanter and into the adjacent part of the capsule ofthe hip. It reminds one strongly of the supra-spinatus.

Gernelli. A small short muscle arises from the deeper partof the tuber ischii near to and in front of the acetabulum andimmediately in front of the preceding muscle. It is insertedinto the inner surface of the inner or fibular trochanter. ThisI conclude may be regarded as the representative of the ge-melli. None of its fibres entered into the pelvis.

There is no trace of a distinct obturator interns either inthe interior or on the exterior of the pelvis.

Three flexor muscles of the knee pass from the fore andinner part of the pelvis to the middle of the plantar or anteriorsurface of the tibia, instead of being attached as usual to thesides of the leg, bones; in this respect resembling the flexors ofthe forearm. They are the three following.

Gracilis (Gr.) from the edge of the spine of the pubes, inits whole length, also from the symphysis and from the roundedangle between the two. Its tendon joins that of the followingmuscle;

Seemi-tendinosus from almost the hindmost part of the is-chium (the back part of the tuber ischii). The tendons ofthese two are united in the lower third of the thigh, and areinserted into a projecting ridge in the middle of the back ofthe tibia, .Ith from the upper end, beneath therefore the spacethat would usually be occupied by the popliteus.

Semi-menbranosua from the tuber ischii just in front ofthe preceding. Its tendon runs clear of the other two andabove them to the same ridge a little higher up.

This accords with the description of the above three flexors byCuvier and Meckel, neither of whom admit the presence of theBiceps in Cheiroptera. The tendons of all these three lie entirely onthe tibial side of the gastrocnemius which indicates them to be tibialmuscles. The insertion of these flexors into the middle of the frontof the tibia, instead of on either side of the leg bones, accords withthe fact that, owing to the transversely excavated or grooved con-dition of the upper surface of the tibia, no rotatory motion-no pro-nation and supination-of leg can be effected. These movementsare limited to flexion and extension as in the case of the two bones ofthe forearm upon the humerus; and the disposition of the flexor

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muscles is accordingly the same. It is also interesting to observethat there are no interarticular cartilages in the knee-joint.

Extensor cruris (Ext. cr.), not large, from the margin of theilium, external to and behind the acetabulum; also from theupper half of the hinder surface of the femur, on one sidebeing prolonged up to the tibial, or outer, and on the other upto the fibular, or inner, trochanter. The two portions-theiliac and the femoral portion-are quite distinct, though statedby Meckel not to be so in the Bat. Its tendon descends alongthe back of the thigh, over the projecting knee, to a ridge-likeprocess on the middle of the posterior and upper edge of thetibia, which is not so large as that in the bird, but rises rathermore directly upon the joint. There is no patella.

The absence of the patella may be associated with the circum-stance that little or no weight is borne upon the hind limbs in theseanimals and that, consequently, the pressure of the extensor tendonupon the femoial condyles is proportionately small.

The view of parts obtained by a dissection of the outer region ofthe left thigh and haunch, given in PI. VI. figs. 3 and 4 is highlyinstructive with reference to their comparison with correspondingparts in a fore-limb. The angle formed by the femur with theilium resembles that of the humerus with the lower or hinder marginof the scapula. The pubes is directed inwards overhanging the jointlike the acromion. The ischium is directed backwards and inwardsfrom the joint. The ligamentum teres ascends, from the dimple onthe outer and fore-part of the head of the femur, beneath the edge ofthe cotyloid ligament covering the notch between the pubes andisehium, to the edge of the acetabulum between these two bones,reminding us of the gleno-humeral ligament'. The extensor muscle(Ext. cr.) passes from behind the acetabulum, down the back of thethigh, over the projecting back of the knee, to the hinder edge of thetibia. Theflexors, semni-tendinosus and semi-membranosus, pass from theischium in front and on the inner side of the joint, over the fiont ofthe thigh, on the tibial side of the flexors of the foot, to the middleof the fore-part of the tibia. The gracilis passes from the outer andanterior projecting edge of the pubes-like part of the delto-pectoralgroup in the fore-limb from the clavicle and sternum-and joins thesemitendinosus, just as the part of the deltoid in Orycteropus (Vol.ii. p. 299) joins the biceps and becomes a flexor or (which is moreappropriate for comparison) as the clavicular or the upper sternalpart of the pectoral not unfrequently extends to the forearm or theelbow. The pectineits, adductor brevis, and adductor magnus passfrom the crest of the pubes to the outer surface of the femur, just asseveral parts of the deltoid pass from the acronion and clavicle to the

1 See my Treatise on the Human Skeleton, p. 411.

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outer surface of the humerus. The obturator externus passes from theobturator ligament, between the pubes and ischium, over the front ofthe hip, to the capsular ligament covering the interval between thetrochanters and to the inner or fibular trochanter, reminding usforcibly of the 8upraspinatt which passes from between the acromionand the coracoid, over the front of the shoulder, to the capsulebetween the tubercles. The difference is that this muscle inclines tothe outer or radial tubercle and extends beyond the interval betweenthe acromion and coracoid to the dorsum of the scapula; whereas theobturator inclines to the fibular trochanter and is limited by theobtuiator ligament. The psoas magnus and iliacus interns, passingfrom the anterior and outer surface of the ilium, beneath the pec-tineus and gracilis, to the outer trochanter, remind us of the infra-spinatus and teres minor passing from the outer surface of the scapulabeneath the deltoid, to the outer tubercle of the humerus. Theglutceus passing from the spines of the sacrum, on the inner side ofthe extensor muscle, to the inner side of the femur beneath theinner trochanter, is the reminder of the lati'S&'Mu8 dorsi, whichtakes a nearly similar direction and has a similar attachment in thefore-limb.

Gastrocnemius, a delicate muscle, arises from the condyles ofthe femur in the usual manner, except that it is attached onlyto the fibular side of the outer or tibial condyle. The innerhead (from the fibular condyle) is tendinous; the outer head(from the tibia] condyle) is muscular. There is a minute ossiclein the outer head; I cannot find one in the inner head. Boththese heads lie on the inner, or fibular, side of the fiexores curis.They unite and terminate in a thin tendon, which runs down tobe inserted into the os calcis. A little above its insertion thebase of the styliform bone (St. b.) is closely connected with it.The bone takes its curved course inwards towards the oppositelimb in the connecting cutaneous fold, having no direct bonyconnection. A slip of muscular fibres (S. m. styliform muscle)passes from the proximal end of the fifth metacarpal andradiates out to be inserted along the lower margin of thisbone.

I do not find any representative of the Solceus or of thePopliteus. The absence of the latter accords with the want ofpronatory and supinatory movement mentioned above (Meckelnotes the absence of this muscle in the bat). Neither do Ifind any Plantaris.

Tibialis posticus (Tib. p.), small, from the lower half of thenner (fibular) surface of the back of the tibia and from the

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adjacent surface of the fibula to the tarsal bone supportingthe second metatarsal. Meckel thinks it is absent in the bat.

Flexores digitorum long, two long and large muscles fromthe two sides of the leg and the leg bones, separated above bythe flexores cruris. One, Tibiatlis (Fl. d. tib.), arises by a largemuscular belly from the outer or tibial condyle of the femur,external to and encroaching upon the space usually occupied bythe gastrocnemius, from the posterior and outer surface of thetibia, and by a muscular slip from the tendon of the semi-tendinosus. A muscular slip also is derived from the back ofthe tibia above the flexor tendons in the position usuallyoccupied by the popliteus. This extends to the inner (fibular)articular surface of the tibia, but does not pass over the jointto the condyle of the femur. The other, fibularis (Fl. d. fib.),corresponding with the FL. 1. h. of man, arises tendinous fromthe tubercular projection on the fibular side of the head of thetibia, and muscular from the tibia beneath it and from theupper half of the fibula'. Their strong tendons pass throughdistinct sheaths in front of the ankle; the tibial flexor sendsa tendon to the ungual phalanx of the hallux; the fibularflexor sends one to the ungual phalanx of digit v; and thetendons of the two muscles combine to send tendons to theungual phalanges of the three middle digits. The tibial lies,in the sole, superficial to the fibular; but their three middletendons are intimately blended.

By this blending of the tendons from the two flexor musclesand the non-division of the extensor tendon till it has neared thephalanges, synchronism in the movements of the digits is insured, anyone not being moveable alone. The greater individuality in themovement of the digits, especially in the hand, which is so markedin ruan, is in him provided for by the greater separation of the flexorand extensor tendons passing to the fingers and toes.

The origin of the tibial flexor of the toes from the condyle of thefemur' and from the flexor tendon of the leg is interesting. It hasthe effect of associating flexion of the toes with that of the leg', and

1 The fibula, though deficient above, is pretty well developed in the lowerpart of the leg, and has a broad surface at the lower end which is jointed withthe os calcis.

2 It is worthy of remark that in birds the flexors of the toes derive originsfrom the condyles of the femur.

3 Just as, in the bird, the insertion of part of the tensor plicae alaris, fromthe clavicle, into the supinator longus, which is inserted into the metacarpal of

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it has the further effect of causing the forced extension of the legwhen the animal is hanging by its curved pedal claws, to give tensionto the flexor muscle of the toes. and so increases the grasp of theclaws. In short the weight of the animal is thus made to conduceto the firmness of the hold by which the weight is maintained. Thisconnection of the flexors of the digits with the femur is very rarelymet with in mammals.

Lumbricales are eight in all, and large. One from the tibialor superficial flexor tendon going to digit i. is inserted into thefibular side of that digit; one from the tendon of the fibularor deep flexor going to digit v. is inserted into the tibial side ofthat digit; the others, arising apparently from the combinedtendons of the tibial and the fibular flexors, are inserted one onthe tibial and one on the fibular side of each of the other digits.They are inserted, not into the extensor tendons, but into thesides of the shafts of the first phalanges about the middle.

The imperfect complement of lumbricales in digits I. and v. shownby the absence of the marginal or distal ones, whereas two are presentin each of the other digits, may be regarded as among the many evi-dences, of which small size is the most obvious, of the incompletenessof these lateral and least constant, last segmented and, as it were,least cared for elements of the terminal part of the limb. The lesscomplete blending of the tibial and fibular components of the flexormuscle in the tendons passing to these digits as compared with thosepassing to the other digits, as well as the absence of the flexor brevison the fibular side of digit v. and its total absence in digit I, and theabsence of a long extensor in digit i, are features of similar import.

Accessories I do not find.Flexor brevis digitorum arises from the os calcis and the

lower part of the tendo Achillis on its tibial side-that is, inthe situation usually occupied by the plantaris tendon whikh,as I have said, is absent. It passes to the second phalanges ofdigits ii, III, and iv, splitting to permit the long flexors to passjust as the corresponding tendons do in man, and reunitingabove them. It passes also to digit v, but only to its fibularside.

Interossei constitute a pair of muscles on the plantar surfaceof each of the metatarsals, arising also from the tarsals, ten inall. One is on the tibial and one on the fibular side of each

the pollex, serves to associate extension of the carpus and digits with that ofthe forearm.

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metatarsal. They pass to the tibial and fibular sides of thepalmar surface of the first phalanges of their respective digits alittle beyond their proximal ends. I could not trace them tothe extensor tendons. They act chiefly as flexors of the firstjoints of the digits'.

Abductor minimi digiti, a short, thick muscle extending fromthe os calcis, near the insertion of the tendo Achillis, to thefibular side of the fifth metatarsal.

Transversus pedis (Tr.) is a distinct muscle arising from theinner cuneiform bone, from the fibular side of the proximal endof the first phalanx of the hallux, and from a tendinous archbetween these two points, beneath which passes the fibular in-terosseous of the hallux. From this broad origin the fibresconverge to the tibial side of the first phalanx of the fifth digit,where they are blended with the interosseous of that side. Itcorresponds, apparently, with the 'adducteur transverse' ofCuvier, who does not mention its presence in this animal.

The metatarsals admit of a certain amount of lateral movement,abduction and addiction, which is effected by the alternate action ofthis transversus and the tibialis anticus and the peroneus brevisaidpd somewhat by the lateral interossei. The peroneus longus notpassing to the metatarsal of hallux and playing little or no part inabduction may be a reason for the presence of this transversus.Moreover the interossei are more parallel than usual, and those tothe lateral digits (i. and iv.) do not extend across so as to serve asadductors, which is an additional reason for the presence of a dis-tinct and well developed transverse muscle. It will be observed thatthe separators of the metatarsals-tibialis anticus and peroneus brevis-are on the dorsal or extensor aspect of the limb, and the approxi-mator of them-the transversus-is on the plantar or flexor as-pect. With regard to this association of abduction with extensionand of adduction with flexion, see preceding Vol. p. 306.

Tibialis anticus (Tib. a.) descends as usual from the fibularsurface of the tibia, over the tibial side of the tarsus, and is in-serted into the tibial side of the first metatarsal, abducting itand turning the foot outwards.

Extensor digitorum (Ext. dig.), from the fore part and thefibular side of the head of the tibia. Its long tendon passesover the tarsus and metatarsus and, near the phalanges, radiatesinto four slips to the four small toes. It is a delicate muscle;

1 See remarks on the arrangement of the interossei in preceding Vol. p. 305.

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Extensor brevis, comparatively strong, arises from the frontof the os calcis and the back of the metatarsals. It passes asusual to the four, in this instance, outer digits (i, II, III, Iv.),being the only extensor of the hallux. Its four divisions aredistinct, each lying along its metatarsal bone. The two middle,to digits ii. and iii, present a longitudinal separation, somewhatreminding us of the interossei on the plantar aspect. Thesedivisions join the long extensor tendons, one portion of eachjoining the fibular and the other the tibial edge of a tendon.The division to digit iv. is developed, or well developed, onlyalong the fibular side of the metacarpal, and joins only thefibular side of the extensor longus tendon, which is the usualdisposition of the extensor brevis in the case of all the digits inother animals. The division to digit I. is large, is inserted partlyinto the back of the first phalanx, and sends a delicate tendonalong the back of this phalanx to the base of the secondphalanx.

The distribution of the extensor brevis to the two sides of theextensor tendons of digits ii. and ini. is interesting in connection withthe anatomy of the lumbricales. Usually the tendons of the extensorbrevis lie only on the fibular sides of those of the extensor longs; andto balance them, as it were, the lumbricales pass to the tibial sides ofthe tendons ofthe extensor longus. But in this instance there is agreater completeness and separation of the extensor brevis and thelumbricales: the tendons of the former pass to both sides of the ten-dons of the extensor longus and the lumbricales have no connectionwith the extensor tendons, pass on both sides of the phalanges, andare inserted into them.

Peroneus longus, from the fibular side of the head of thetibia and the septum between the extensor and the flexordigitorum descends, in front of the lower end of the fibula,through a groove on the outer side of the tarsus, to the underpart of the proximal end of the second metatarsal bone. Icannot trace it to the metatarsal of hallux.

Peroneus tertius, rather large, from front of fibula, beneathand internal to extensor digitorum, to metatarsal bone of digit v.In the male it sends a tendon to join the extensor tendon of thisdigit; and there is also a small slip to this digit from the ex-tensor brevis. In the female this slip is wanting; but theperoneal tendons to the extensor and to the metatarsal are from

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distinct muscles. Thus the extensor tendon of digit v. is sup-plied from two or three sources, whereas that to hallux is derivedonly from the Ext. br. No other peroneus present. Meckelfinds only one peroneus in Cheiroptera.

PLATE VI.

Fig. 1. Muscles on the abdominal aspect of male Pteropus.The right hind-limb has been partially and the left has been muchturned or rotated from the natural position for the purpose of show-ing the muscles on the two surfaces.

1. 2. 3. 4. The facial, occipital, cervical and sternal portions ofplatysma uniting to form 5 the tensor plicse alaris, which is elasticfrom near the convergence of the muscular bundles to a, between aand b is again muscular, and between b and its insertion is tendinous.

St. h., sterno-hyoid. Digr., Iigastric. Tr. in., trachelo-mastoid.St. ct. m., sterno-cleido-mastoid, from the anterior extremity of thekeel, from the upper edge of the keel (this is rather in shade) andfrom the clavicle. Subcl., subclavius. C. A., cervico-humeral. R. a.,rectus-al)dominis. The sternal portion of the first rib is seen betweenthis and the subelavius.

RIGHT FORE-LIMB. Pect., sternal portion of pectoralis major dividedand reflected from its origin. Pect. a., abdominal portion of pectoral.The clavicular portion has been removed to show subclavius. B.,biceps. C. b., coraco-brachialis. Br. a., brachialis anticus. Ext. c. r.,extensor carpi radialis. Sup. 1., supinator longus. Pr. t., pronatorteres. El. c. r., flexor carpi radialis. Fl. d. s., flexor digitorumsublimis. Fl. c. u., flexor carpi ulnaris.

RIGHT HIND-LIMNIB. Ps. p., psoas parvus. Ps., psoas magnus It. i.,iliacus internus. P., pectineus. Gr., gracilis. F. c., femoro cutaneous.Fl. d., flexor dig. long. P. I., peroneus longus. Ext. d., extensordigitorum. P. 3, peroneus tertius. fib. a., tibialis anticus.

LEFT HIND-LIMB. Gr., gracilis. Ad., adductor muscles. S. in.,semi-membranosus. S. t., semi-tendinosus. I. c., ischio-cutaneous.St. m., styliform muscle. St. b., styliform bone. 1. Ach., tendoAchillis. Fl. d.fib., flexor digitorum fibularis. Tib. p., tibialis posticus.Fl. d. tib., flexor digitorum tibialis.

LEFT FORE-LIMB. Fl. br., flexor brevis, ' interosseal flexors' ofdigit v. Abd., abductor of digit v. Fl. c. r., flexor carpi radialis.Fl. d. pr., flexor digitorum profundus. Fl. d. s., flexor digitorumsublimis. Fl. c. u., flexor carpi ulnaris. Pr. t., pronator teres. Sup. I.,supinator longus.

Fig. 2. The right side of pelvis femur and upper part of tibia,of male Pteropus. It., upper part or crest of ilium. C., crest of pubeswhere psoas parvus and pectineus are attached. S., symphysis pubis.ech., ischium.

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Fig. 3. View of superficial muscles on the outer side of the righthip in female Pteropus the bones being in the same position as in pre-ceding figure. S. t., semi-tendinosus. S. in., semimembranosus. P., pec-tineus. Cr., gracilis. Ad. br., adductor brevis. Ps. p., psoas parvus.Ps., psoas magnus. II. i., iliacus internus. Gl., glutaeus maximus.Ext. er., extensor cruris. F. c., origin of femoro-cutaueous.

Fig. 4. View of deep muscles about right hip of the same animal.Gr., gracilis divided and reflected upwards and downwards. S. t. andS. mn., semi-tendinosus and semi-membranosus. P., pectineus dividedand reflected upwards and downwards. Ad. in., adductor magnusdivided and upper part reflected upwards. Ad. br., adductor brevisreflected upwards and downwards. L. t., ligamentum teres. Obt. e.,obturator externus. Ps. p., psoas parvus. Ps., psoas magnus. 11. i.,iliacus internus.

Fig. 5. Dissection of foot showing the transversalis muscle (Tr.).Rl. br., flexor brevis divided and reflected to the os calcis. Inteross.,the interossei of hallux, and digit v. S. m., muscle from metacarpalv. to styliform bone.

PLATE VII.

Fig. 6. Muscles on the dorsal aspect of male Pteropus.FORE-LIMBS. EL.C. U., extensor carpi ulnaris. Ex. dig., extensor

digitorum. Ex. p.sec., extensor pollicis secundus. EX. i., extensorindicis. Ex. p. pr., extensor pollicis primus. Ex. c. r. br., extensorcarpi radialis brevior. Ex. c. r. I., extensor carpi radialis longior.Sup. I., supinator longus. Sup. br., supinator brevis. B., biceps. D.,deltoid. Trap., trapezius. C. h., cervico-humeral. S., sesamoid bonein tendon of (Tr.) triceps. A. se., angle of scapula projected frombeneath (Lat. d.) latissimus dorsi. H., humerus. Lev. se., levatorscapulae. S. 8p., supra-spinatus. L. sp., infra-spinatus. T. min., teresminor. T., teres major.

HIND-LIMBS. Il. i., iliacus internus. Gl. med., gluteus medius.Ext. c., extensor cruris. S. in., semi-membranosus; S. t. and Gr., semi-tendinosus and gracilis. Ad., abductors. Gl., glutseus maximus.Q. 1. quadratus lumborum.

Fig. 7. Side view of muscles of shoulder and hip for pur-pose of comparison. D'., portion of deltoid passing to humerus.D"., portion of deltoid continued into outer part of triceps. Pect.,pectoralis major. B., biceps. Br. a., brachialis anticus. Gr., gracilis.P., pectineus. Ad. adductor. S. in., semi-membranosus. S. t. semi-tendinosus. Ext. cr., quadriceps extensor cruris. Il. i., iliacus in-ternus. Ps., psoas magnus. Ps. p., psoas parvus.

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