Fig. 99. Half front view of the skulls,Aof an old,Bof ayoung Gorilla(Gorilla savagei) × ¼. (Camb. Mus.)1. parietal.5. squamosal.2. sagittal crest.6. maxillae.3. frontal.7. external auditory meatus.4. supra-orbital ridge.
Fig. 99. Half front view of the skulls,Aof an old,Bof ayoung Gorilla(Gorilla savagei) × ¼. (Camb. Mus.)1. parietal.5. squamosal.2. sagittal crest.6. maxillae.3. frontal.7. external auditory meatus.4. supra-orbital ridge.
Fig. 99. Half front view of the skulls,Aof an old,Bof a
young Gorilla(Gorilla savagei) × ¼. (Camb. Mus.)
In Man and in most monkeys, at any rate when young (fig. 99, B), the roof of the skull is smooth and rounded, but in many forms, such as the Baboons, in the adult the supra-orbital and occipital ridges are much developed. In the Gorilla this is also the case with the sagittal crest (fig. 99, A, 2). The bones of the upper surface of the cranium interlock with wavy outlines. The nasals vary much in length, being much shorter in man than in most monkeys; they commonly become early fused together, as do also the frontals. The vomer is well developed, and the ethmo-turbinal always forms part of the boundary of the orbit. There are frequently, as in many Lemuroidea, a pair of more or less well-marked ridges, crossing the roof of the skull from the postorbital processes of thefrontals to the occipital crest. The orbit is completely encircled by bone, and the alisphenoid assists the jugal and frontal in shutting it off from the temporal fossa, leaving however a communication between the two as the sphenomaxillary fissure. In most cases the frontals meet one another in the middle line between the mesethmoid and orbitosphenoid, but in Man, Simia, and some Cebidae this does not take place. In nearly all Cebidae the parietal and jugal meet one another, separating the frontal and alisphenoid on the skull wall; in Man and all Old World monkeys, on the other hand, the alisphenoid and frontal meet and separate the jugal and parietal. The premaxillae nearly always send back processes which meet the nasals. The palate is rather short and both the palatine and the premaxillae take a considerable part in its formation. The pterygoid plate of the alisphenoid is decidedly large, and there is no alisphenoid canal. There is never any great development either of the paroccipital process of the exoccipital, or of the postglenoid process of the squamosal. The periotic and tympanic are always fused together; in Cebidae they form a small bulla, but a bulla is not developed in any Old World forms. The periotic is large, especially the mastoid portion, which forms a distinct portion of the skull wall between the squamosal and exoccipital. In Man and still more in Old World monkeys, the external auditory meatus is drawn out into a definite tube, whose lower wall is formed by the tympanic; in the Cebidae the tympanic is ring-like. The perforation of the periotic by the carotid canal is always conspicuous.
The mandible is rather short and broad, and the angle formed by the meeting of the two rami is more obtuse than in most mammals. The coronoid process is fairly well developed, and the angle is more or less rounded. In most Primates the condyle is considerably widened, but this is not the case in Man. InMycetesthe mandible is very large, its ascending portions being specially developed. The hyoid of Primates isremarkable for the large expanded basi-hyal, which is generally concave above and convex below. The anterior cornu is never well ossified, but the thyro-hyal is always strong. InMycetesthe basi-hyal is enormously large, forming a somewhat globular thin-walled capsule.
Fig. 100. Malleus, stapes and incus ofA.Man.B.Dog.C.Rabbit.(AfterDoran) x 1.1. head of malleus.5. manubrium of malleus.2. canal of stapes.6. processus brevis.3. incus.7. lamella.4. processus longus (or gracilis).
Fig. 100. Malleus, stapes and incus ofA.Man.B.Dog.C.Rabbit.(AfterDoran) x 1.1. head of malleus.5. manubrium of malleus.2. canal of stapes.6. processus brevis.3. incus.7. lamella.4. processus longus (or gracilis).
Fig. 100. Malleus, stapes and incus of
A.Man.B.Dog.C.Rabbit.(AfterDoran) x 1.
Auditory ossicles.
There are in mammals four auditory ossicles forming a chain extending from the fenestra ovalis to the tympanic membrane. Three of these, themalleus,incusandstapes, are always distinct, while the fourth, thelenticular, is smaller than the others and is sometimes not distinct. The names are derived from human anatomy and indicate in the case of the first three a more or less fanciful resemblance respectively to a hammer, an anvil and a stirrup. The ossicles are homologousas a whole to the hyomandibular of fishes and to the columellar chain of Sauropsids and Amphibians. The malleus is homologous to the extra-columella of Crocodiles and the stapes to the columella. Themalleuswhen typically developed consists of a roundedhead(fig. 100, 1) which bears a surface articulating with the incus, and a shortneckcontinued into a process, themanubrium(fig. 100, 5), which comes into relation with the tympanic membrane. From the junction of the neck and manubrium two processes are given off, aprocessus longusorgracilis(fig. 100, 4), which in the embryo is continuous with Meckel's cartilage, and aprocessus brevis(fig. 100, 6). Theincusgenerally consists of a more or less anvil-shaped portion which articulates with the malleus, and of a process which is connected with the stapes by the smalllenticular. Thestapesis generally stirrup shaped, consisting of a basal portion from which arise twocruraseparated by a space the canal through which a branch of the pharyngeal artery runs The lenticular is frequently cartilaginous and sometimes is not developed at all.
The above is the arrangement of the auditory ossicles met with in the higher Mammalia, but in the lower Mammalia the characters approach more nearly to those met with in Sauropsids.
InMonotremesthe ossicles, though distinctly mammalian in character, show a very low type of development. The incus is articulated, or often fused, with an outgrowth from the head of the malleus. The stapes is very much like a reptilian columella, having a single crus with no perforation.
InMarsupialsthe ossicles are of a low type, but not so low as the rest of the skeleton might have led one to expect, and all or almost all the points showing a low grade of development may be paralleled among the Monodelphia. The lowest Marsupials as regards the ossicles are the Peramelidae, whose ossicles are of a frail papery consistence. The Didelphyidae on the other hand have the most highly developedossicles, the malleus much resembling that of many Insectivores, and the stapes having two definite crura separated by a canal.
InEdentatesthe character of the ossicles varies much. In Sloths the stapes approaches that of Sauropsids in its narrowness and the slight trace of a canal; this character is however still more marked inManis, whose stapes is as Sauropsidan as that of Monotremes, and consists of a nearly circular basal plate bearing a column which does not show any sign of division into crura. The stapes of other Edentates, such as ant-eaters, aard varks, and most armadillos, is of a high type and has well-developed crura.Priodonhas a lower type of stapes thanDasypusandTatusia.
The ossicles of theSireniadiffer widely from those of all other mammals in their great density and clumsy form.
InCetaceathe ossicles are solid, though not so solid as in Sirenia, and their details vary much. The malleus is always firmly fused to the tympanic by means of the processus longus, and the manubrium is very little if at all developed. The incus has the stapedial end greatly developed, and the stapes has very thick crura with hardly any canal. The ossicles of the Mystacoceti are apparently less specialised than are those of the Odontoceti.
The auditory ossicles of theUngulatado not present any characters common to all the members of the group.
Among Ruminants they are chiefly remarkable for the development of a broad lamellar expansion between the head and the processus longus of the malleus. In some cases the malleus of the foetus differs strikingly from that of the adult. Among Perissodactyla the Rhinoceros and Tapir have the malleus of a low type, recalling those of Marsupials; while in the Horse the head is well developed, and the malleus is of a higher type.
The ossicles ofProcavia, which recall those of the Equidae, are chiefly remarkable for the small size of the body of the incus. In Elephants the ossicles are large and massive.
In theRodentia(fig. 100, C) the malleus is generally characterised by a very broad manubrium. In many genera such asBathyergus, and most of the Hystricomorpha such asHystrix,ChinchillaandDasyprocta, the malleus and incus are ankylosed together.
Carnivora.In Carnivora vera the most striking feature of the malleus is the occurrence of a broad lamellar expansion between the head and neck and the processus longus. This however does not occur in some Viverridae. In the Carnivora vera the incus and stapes are small as compared with the malleus, but in the Pinnipedia they are large. In the Pinnipedia the auditory ossicles have a very dense consistence, and except in the Otariidae are very large. The stapes frequently has no canal, or only a very small one.
InInsectivorathe characters of the auditory ossicles are very diverse. Many forms such as shrews, moles, hedgehogs, and the Centetidae have a low type of malleus resembling that of Edentates.Chrysochlorishas very extraordinary auditory ossicles. The head of the malleus is drawn out into a great club-shaped process, the incus is long and narrow, and differs much from the ordinary type.
InChiropterathe ossicles and especially the malleus much resemble those of shrews. The stapes is always normal in character, never becoming at all columelliform.
Primates.In Man and the Anthropoid Apes the malleus has a rounded head, a short neck, and the manubrium, a processus longus and a processus brevis. The incus consists of an anvil-shaped portion from which arises a long tapering process. The stapes has diverging crura and consequently a wide canal. The crura in other monkeys do not diverge so much as in man and anthropoid apes. The New World monkeys have no neck to the malleus.
The Sternum[168].
InMonotremesand mostMarsupialsthe sternum does not present any characters of special importance. The presternum is strongly keeled inNotoryctes.
The sternum inEdentatesis very variable: in the Sloths it is very long, the mesosternum ofCholoepushaving twelve segments. In the ant-eaters and armadillos the presternum is broad and sometimes as inPriodonstrongly keeled. InManis macrurathe xiphisternum is drawn out into a pair of cartilaginous processes about nine inches long.
In theSireniathe sternum is simple and elongated, and of fairly equal width throughout, in the adult it shows no sign of segmentation. Its origin from the union of two lateral portions can be well seen inManatus.
Two distinct types of sternum are met with in theCetacea. In the Odontoceti the sternum consists of a broad presternum followed by three or four mesosternal segments, but with no xiphisternum. Indications of the original median fissure can be traced, and are very evident inHyperoödon. In the Mystacoceti, on the other hand, the sternum consists simply of a broad flattened presternum which is sometimes more or less heart-shaped, sometimes cross-shaped. Only a single pair of ribs are united to it.
The sternum inUngulatais generally long and narrow and formed of six or generally seven segments. The presternum is as a rule small and compressed, often much keeled, especially in the horse and tapir. The segments of the mesosternum gradually widen as followed back and the xiphisternum is often terminated by a cartilaginous plate.
In theRodentiathe sternum is long and narrow and generally has a large presternum, and a xiphisternum terminated by a broad cartilaginous plate.
In theCarnivora, too, the sternum (fig. 76) is long and narrow and formed of eight or nine pieces, all of nearly the same size. The xiphisternum generally ends in an expanded plate of cartilage.
InInsectivorathe sternum is well developed but variable. The presternum is commonly large and is sometimes as in the Hedgehog (Erinaceus) bilobed in front, sometimes as in the Shrew (Sorex) trilobed. It is especially large in the Mole (Talpa) and is expanded laterally and keeled below.
In theChiropterathe presternum is strongly keeled and so is sometimes the mesosternum.
AmongPrimates, in Man and the Anthropoid Apes the sternum is rather broad and flattened; the mesosternum consists of four segments which are commonly fused together and the xiphisternum is imperfectly ossified.
The Ribs.
Free ribs are borne as a rule only by the thoracic vertebrae; ribs may be found in other regions, especially the cervical and sacral, but these are almost always ankylosed to the vertebrae. As a general rule the first thoracic rib joins the presternum, while the succeeding ones are attached between the several segments of the mesosternum. Some of the posterior ribs frequently do not reach the sternum; they may then be attached by fibrous tissue to the ribs in front, or may end freely (floating ribs). There are generally thirteen pairs of ribs, and in no case do they have uncinate processes.
InMonotremes(fig. 102, B) each rib is divided not into two but into three parts, an intermediate portion being interposed between the vertebral and sternal parts. The sternal ribs are well ossified, and some are very broad and flat. The intermediate portions are unossified, those of the anterior ribs are short and narrow, but they become longer and wider further back.
InMarsupialsthere are almost always thirteen pairs of ribs, whose sternal portions are very imperfectly ossified.Notorycteshas fourteen pairs of ribs, eight of which are floating: the first rib is very stout, and is abruptly bent on itself to join the sternum. It has no distinct sternal portion. All the other ribs are slender.
Of theEdentatesthe Sloths have very numerous ribs; twenty-four pairs occur inCholoepus, and half of these reach the sternum. In the Armadillos there are only ten or twelve pairs of ribs, but the sternal portions are very strongly ossified. The first rib is remarkably broad and flat, and is not divisible into vertebral and sternal portions.
In theSireniathere are a very large number of ribs noticeable for their great thickness and solidity, but not more than three are attached to the sternum.
Cetacea.In the Whalebone whales the ribs are remarkable for their very loose connection both with the vertebral column and with the sternum. The capitula are scarcely developed, and the attachment of the tubercula to the transverse processes is loose. The first rib is the only one connected with the sternum. In the Toothed whales the anterior ribs have capitula articulating with the centra, as well as tubercula articulating with the transverse processes; in the posterior ones, however, only the tubercula remain. Seven pairs of well-ossified sternal ribs generally meet the sternum. In the Physeteridae most of the ribs are connected to the vertebrae by both capitula and tubercula.
In theUngulatathe ribs are generally broad and flattened, and this is especially the case in the generaBosandBubalus(fig. 101, 6). The anterior ribs are short and nearly straight, and sternal ribs are well developed. The Artiodactyla have twelve to fifteen pairs of ribs, the Perissodactyla eighteen or nineteen, andProcaviatwenty to twenty-two. The Elephant has nineteen to twenty-one pairs, seven of which may be floating ribs.
Fig. 101. Skeleton of a Cape Buffalo(Bubalus caffer).The left scapula is omitted for the sake of clearness ×1/17. (Brit. Mus.)1. premaxillae.7. femur.2. nasal.8. patella.3. orbit.9. tibia.4. neural spine of first thoracic10. metatarsals.vertebra.11. radius.5. scapula.12. metacarpals.6. rib.
Fig. 101. Skeleton of a Cape Buffalo(Bubalus caffer).The left scapula is omitted for the sake of clearness ×1/17. (Brit. Mus.)1. premaxillae.7. femur.2. nasal.8. patella.3. orbit.9. tibia.4. neural spine of first thoracic10. metatarsals.vertebra.11. radius.5. scapula.12. metacarpals.6. rib.
Fig. 101. Skeleton of a Cape Buffalo(Bubalus caffer).
The left scapula is omitted for the sake of clearness ×1/17. (Brit. Mus.)
In theRodentiathere are generally thirteen pairs of ribs,which do not present any marked peculiarities.
TheCarnivorahave thirteen to fifteen pairs of ribs, whose vertebral portions are slender, nearly straight and subcylindrical, while their sternal portions are long and imperfectly ossified (fig. 76, 5). There is nothing that calls for special remark about the ribs, in eitherInsectivoraorChiroptera.
Primates.In Man and the Orang (Simia) there are generally twelve pairs of ribs; in the Gorilla and Chimpanzee (Anthropopithecus), and Gibbons (Hylobates), there are thirteen, in the Cebidae twelve to fifteen, and in the Lemuroidea twelve to seventeen pairs. The first vertebral rib is shorter than the others, and the sternal ribs generally remain cartilaginous throughout life, though in man the first may ossify.
Appendicular Skeleton.
The Pectoral Girdle.
By far the most primitive type of the pectoral or shoulder girdle is found in theMonotremata. The scapula (fig. 102, A, 1) is long and recurved, and has only two surfaces, one corresponding to the prescapular[1] fossa, the other to the postscapular[1] and subscapular[169]fossae. The coracoid is a short bone attached above to the scapula and below to the presternum; it forms a large part of the glenoid cavity. In front of the coracoid there is a fairly large flattened epicoracoid (fig. 102, 6);there is also a largeT-shaped interclavicle (fig. 102, 4), which is expanded behind and rests on the presternum. The clavicles rest on and are firmly united to the anterior border of the interclavicle. This shoulder girdle differs greatly from that of any other mammals, and recalls that of some Lacertilia.
Fig. 102.A, Side view,B, Dorsal view of the shoulder girdle andpart or the sternum of the Spiny Anteater(Echidna aculeata)× 1. (AfterParker.)1. scapula.6. epicoracoid.2. suprascapula.7. glenoid cavity.3. clavicle.8. presternum.4. interclavicle.9. second sternal rib.5. coracoid.10. second vertebral rib.
Fig. 102.A, Side view,B, Dorsal view of the shoulder girdle andpart or the sternum of the Spiny Anteater(Echidna aculeata)× 1. (AfterParker.)1. scapula.6. epicoracoid.2. suprascapula.7. glenoid cavity.3. clavicle.8. presternum.4. interclavicle.9. second sternal rib.5. coracoid.10. second vertebral rib.
Fig. 102.A, Side view,B, Dorsal view of the shoulder girdle and
part or the sternum of the Spiny Anteater(Echidna aculeata)
× 1. (AfterParker.)
InMarsupials, as in all mammals except the Monotremes, the shoulder girdle is much reduced; there are no epicoracoids and interclavicle, and the coracoid forms simply a small process on the scapula, ossifying from a centre separate from that giving rise to the rest of the bone. The scapula has a long acromion, and a clavicle is always present except inPerameles. Unossified remains of the precoracoids are found at either end of the clavicle. The scapula ofNotorycteshas a very highoverhanging spine, and there is a second strong ridge running along the proximal part of the glenoid border.
The shoulder girdle of theEdentatashows some very curious variations. InOrycteropusthe scapula is of very normal form and the clavicle is well developed. In the Pangolins and Anteaters the scapula is very broad and rounded; there is no clavicle in the Pangolins, and generally only a vestigial one in Anteaters. In Armadillos, Sloths, and Megatheriidae, the acromion is very long and the clavicle is well developed. In the Sloths,Megatherium, andMyrmecophaga, a connection is formed between the coracoid, which is unusually large, and the coracoid border of the scapula, converting the coraco-scapula notch into a foramen. InBradypusthe clavicle is very small, and is attached to the coracoid, which sometimes forms a distinct bone[170].
In theSireniathe scapula is somewhat narrow and curved backwards: the spine, acromion, and coracoid process are moderately developed, and there is no clavicle.
Cetacea.In nearly all the Odontoceti the scapula is broad and somewhat fan-shaped; the prescapular fossa is much reduced, and the acromion and coracoid process form flattened processes, extending forwards nearly parallel to one another. Some of the Mystacoceti, such asBalaenoptera, have a broad, fan-shaped scapula, with a long acromion and coracoid process, extending parallel to one another. Others, such asBalaena, have a higher and narrower scapula, with a smaller coracoid process.
InUngulatathe scapula is always high and rather narrow, and neither acromion nor coracoid process is ever much developed. In no adult Ungulate exceptTypotheriumis there any trace of a clavicle, but a vestigial clavicle has been described in early embryos of sheep[171].
Fig. 103. Skeleton of a Llama(Auchenia glama) ×1/18.(Brit. Mus.)1. hyoid.6. olecranon process of ulna.2. atlas vertebra.7. metacarpals.3. seventh cervical vertebra.8. ilium.4. scapula.9. patella.5. imperfectly ossified10. calcaneum.suprascapula.
Fig. 103. Skeleton of a Llama(Auchenia glama) ×1/18.(Brit. Mus.)1. hyoid.6. olecranon process of ulna.2. atlas vertebra.7. metacarpals.3. seventh cervical vertebra.8. ilium.4. scapula.9. patella.5. imperfectly ossified10. calcaneum.suprascapula.
Fig. 103. Skeleton of a Llama(Auchenia glama) ×1/18.
(Brit. Mus.)
Ungulata vera.In the Ruminantia the suprascapularregion (fig. 103, 5) is very imperfectly ossified, and when this is removed the upper border of the scapula is very straight (fig. 101, 5). The spine is prominent, and generally has a fairly well-marked acromion. InHippopotamusthe acromion is fairly prominent, but in the other Suina, though the spine is prominent, the acromion is not developed. The Perissodactyla have no acromion, but while the Equidae andHyracotheriumhave the scapula long and slender, with the spine very slightly developed, the other living Perissodactyla have the spine prominent and strongly bent back at about the middle of its length.
Subungulata.Typotherium(Toxodontia) differs from all other known Ungulates in having well-developed clavicles; its scapula has a strong backwardly-projecting process, much like that inRhinoceros.
Phenacodus(Condylarthra), has a curiously rounded scapula, with the coracoid and suprascapular borders passing imperceptibly into one another. The scapula resembles that of a carnivore more than does that of any existing Ungulate.
Procaviahas a triangular scapula with a prominent spine and no acromion; there is a large unossified suprascapular region.
The scapula in the Proboscidea has a large rounded suprascapular border and a narrow, slightly concave glenoid border. The spine is large, and has a prominent process projecting backwards from about its middle. The spine lies towards the front end of the scapula, so that the postscapular fossa is much larger than the prescapular fossa.
InRodentiathe shoulder girdle is of a rather primitive type. The scapula is generally high and narrow, somewhat as in Ruminantia; it differs, however, from the Ruminant scapula in having a high acromion, which is often, as in the Hares and Rabbits, terminated by a long metacromion. The development of the clavicle varies, and sometimes it is altogether absent. It is frequently connected by cartilaginous bands or ligaments (fig. 104, 7 and 9), on the one hand with the scapula, and on the other with the sternum. These unossified bands are remains of the precoracoid. Epicoracoidal vestiges of the sternal ends of the coracoids (fig. 104, 11) are also often present.
In theCarnivora verathe scapula is large, and generally has rather rounded borders. The spine and acromion are well developed, and the prescapular and postscapular fossae are nearly equal in size. The coracoid is very small, and the clavicle is never completely developed, being often absent, as in the Bears and most of their allies. In the Seals (Phocidae)the scapula is elongated and curved backwards, and has a very concave glenoid border. In the Eared Seals (Otariidae) the scapula is proportionally much larger and wider, the prescapular fossa being specially large, and being traversed by a ridge, which converges to meet the spine.
Fig. 104. Dorsal view of the sternum and right half of theshoulder-girdle ofMus sylvaticus× 4. (AfterParker.)1. postscapular fossa.precoracoid at sternal end of2. prescapular fossa.clavicle.3. spine.10. omosternum.4. suprascapular border11. epicoracoid.unossified.12. presternum.5. coracoid process.13. first segment of mesosternum.6. acromion.14. xiphisternum.7. cartilaginous vestige of15. cartilaginous termination ofprecoracoid at scapular end ofxiphisternum.clavicle.16. 2nd sternal rib.8. clavicle.17. 1st vertebral rib.9. cartilaginous vestige of
Fig. 104. Dorsal view of the sternum and right half of theshoulder-girdle ofMus sylvaticus× 4. (AfterParker.)1. postscapular fossa.precoracoid at sternal end of2. prescapular fossa.clavicle.3. spine.10. omosternum.4. suprascapular border11. epicoracoid.unossified.12. presternum.5. coracoid process.13. first segment of mesosternum.6. acromion.14. xiphisternum.7. cartilaginous vestige of15. cartilaginous termination ofprecoracoid at scapular end ofxiphisternum.clavicle.16. 2nd sternal rib.8. clavicle.17. 1st vertebral rib.9. cartilaginous vestige of
Fig. 104. Dorsal view of the sternum and right half of the
shoulder-girdle ofMus sylvaticus× 4. (AfterParker.)
In theInsectivorathe shoulder girdle is well developed and, as in Rodents, remains are met with of various parts not generally seen in mammals. In the Shrews the scapula is long and narrow, and has a well-marked spine, whose end bifurcates, forming the acromion and metacromion. The clavicle is long and slender, and is connected with the sternum and acromion by vestiges of the precoracoid. Considerable remains of the sternal end of the coracoid are also found. InPotamogale, however, there are no clavicles. In the Mole the shoulder girdle is greatly developed, and of very remarkable form. The scapula is high and very narrow, with the spine and acromion very little developed. The other shoulder girdle element is an irregular bone, which articulates with the humerus and presternum, and is connected by ligaments with the scapula. This bone appears to represent both the coracoid and the clavicle, being formed partly of cartilage bone, partly of membrane bone.
In theChiropterathe scapula is large and oval, and has a moderately high spine and a large acromion. The coracoid process is well developed and is often forked. The clavicles are also well developed, and vestiges of the precoracoid and of the sternal end of the coracoid are often found.
InPrimatesthe clavicle and coracoid process are always well developed. In Man and the Gorilla the scapula has a long straight suprascapular border, a well-developed coracoid process and spine, and a large curved acromion. Vestiges of the precoracoid occur at each end of the clavicle. The shape of the scapula varies much in the lower Primates.
The Upper arm and Fore-arm.
In theMonotrematathe humerus is short, very broad at each end and contracted in the middle. The radius and ulna are stout and of nearly equal size, while the ulna has a greatly expanded olecranon.
In theMarsupialiathe humerus is generally a strong bone,broad at the distal end and having well marked deltoid and supinator ridges, which are specially large inNotoryctes. An ent-epicondylar or supracondylar foramen (fig. 105, 5) is almost always present except inNotoryctes. The radius and ulna are always distinct and well developed, and a certain amount of rotation can take place between them. The ulna ofNotorycteshas an enormous hooked olecranon which causes the bone to be nearly twice as long as the radius.
Fig. 105. Anterior surface of the right humerus of a Wombat(Phascolomys latifrons). (AfterOwen.)1. head.6. supinator ridge.2. greater tuberosity.7. external condyle.3. lesser tuberosity.8. internal condyle.4. deltoid ridge.9. articular surface for radius.5. ent-epicondylar10. articular surface for ulna.(supracondylar) foramen.
Fig. 105. Anterior surface of the right humerus of a Wombat(Phascolomys latifrons). (AfterOwen.)1. head.6. supinator ridge.2. greater tuberosity.7. external condyle.3. lesser tuberosity.8. internal condyle.4. deltoid ridge.9. articular surface for radius.5. ent-epicondylar10. articular surface for ulna.(supracondylar) foramen.
Fig. 105. Anterior surface of the right humerus of a Wombat
(Phascolomys latifrons). (AfterOwen.)
Edentata.The Sloths have long slender arm bones; the humerus is nearly smooth and has a very large ent-epicondylar foramen inCholoepus, but not inBradypus. The radiusand ulna can be rotated on one another to a considerable extent. The humerus in all other Edentates is very strong and has the points for the attachment of muscles much developed, especially in the Armadillos and Megatheriidae. An ent-epicondylar foramen is found in all living forms. The radius and ulna are well developed, but are not capable of much rotation.
In theSireniathe humerus is well developed and of a normal character. It is expanded at each end and has a prominent internal condyle, a small olecranon fossa, and no ent-epicondylar foramen. In the Dugong andRhytinathere is a bicipital groove and the tuberosities are distinct, but in the Manatee there is no bicipital groove, and the tuberosities coalesce. The radius and ulna are about equally developed and ankylosed together at both ends.
In theCetaceathe arm bones are very short and thick. The humerus has a globular head, and a distal end terminated by two equal flattened surfaces to which the radius and ulna are united. There is no bicipital groove, and the tuberosities coalesce. The radius and ulna are flat expanded bones fixed parallel to one another, but the ulna has a definite olecranon. Scarcely any movement can take place between them and the humerus, and in old animals the three bones are often ankylosed together.
In theUngulata verathe humerus is stout and rather short. The great tuberosity is always large and often overhangs the bicipital groove, it is especially large inTitanotherium(Brontops). There is never an ent-epicondylar foramen. The radius is always large at both ends, but the condition of the ulna is very variable. Sometimes, as inTapirus,Rhinoceros,Macrauchenia, Suina and Tragulina, the ulna is well developed, and quite distinct from the radius; but in most forms, although complete, it is much reduced distally, and is fused to the radius. Sometimes, as in the Horse and Giraffe, it is reduced to the olecranon and to a very slender descending processwhich does not nearly reach the carpus. In the Tylopoda, though the ulna is complete and its distal end is often distinct, it has coalesced with the radius throughout its whole length; the olecranon is generally very large.
Subungulata.In the large Condylarthra the humerus has an ent-epicondylar foramen, and the radius and ulna are stout bones nearly equal in size.
InProcaviathe humerus is rather long, and has a very prominent greater tuberosity, and a large supra-trochlear fossa, but no ent-epicondylar foramen.
In the Proboscidea the humerus is marked by a greatly developed supinator ridge, and is very long, longer than the radius and ulna. The ulna has a remarkable development, having its distal end larger than that of the radius, it has also a larger articular surface for the humerus than has the radius.
InRodentiathe humerus varies much in its development according to the animal's mode of life. In the Hares it is long and straight, with a small distal end, and a slight deltoid ridge. In the Beaver on the other hand the deltoid and supinator ridges are considerably developed. There is generally a large supra-trochlear fossa, but no ent-epicondylar foramen.
Carnivora.In the Carnivora vera the humerus has large tuberosities, a prominent deltoid ridge and a deep olecranon fossa. The shaft is generally curved, and an ent-epicondylar foramen is often found, though not in the Canidae, Hyaenidae, and Ursidae. The radius and ulna are never united. The radius (fig. 77, B) has a very similar development throughout its whole length, while the ulna has a large olecranon (fig. 77, C, 11) and a shaft tapering somewhat towards the distal end.
In the Pinnipedia the arm bones are very strongly developed. The humerus has a very prominent deltoid ridge, and the proximal end of the ulna and distal end of the radius are much expanded.
In theInsectivorathe arm bones are well developed, and the radius and ulna, though sometimes united, are generally distinct; as a rule there is an ent-epicondylar foramen, but this is absent in the Hedgehog. The Mole has an extraordinary humerus, very short and curved, and much flattened and expanded at both ends. It articulates both with the scapula and coraco-clavicle. The ulna has a greatly developed olecranon.
In theChiropteraboth humerus and radius are exceedingly long and slender; the ulna is reduced to little more than the proximal end and is fused to the radius. There is no ent-epicondylar foramen.
AllPrimateshave the power of pronation and supination of the fore-arm, by the rotation of the distal end of the radius round that of the ulna.
In Man and the Anthropoid Apes the humerus is long and straight, and has a globular head; neither of the tuberosities, nor the deltoid nor supinator ridges are much developed. The olecranon fossa is deep and there is no ent-epicondylar foramen. The radius is curved and has a narrow proximal, and expanded distal end, the ulna is straighter than the radius and has the distal end much smaller than the proximal; the olecranon is not much developed.
In the lower Primates, although the radius and ulna are always quite separate, the power of pronation and supination is not nearly so great as in the higher forms. In most of the Cebidae and Lemurs an ent-epicondylar foramen occurs.
The Manus.
The Manus is divisible into two parts, viz. the carpus or wrist, and the hand which is composed of the metacarpals and phalanges. The carpal bones are always modified from their primitive arrangement, sometimes more, sometimes less. One modification however is always found in mammals, viz. theunion of carpalia, 4 and 5 to form theunciformbone. Two sesamoid bones are commonly developed, one on each side of the carpus, thepisiformor one on the ulnar side being much the larger and more constant: it has been suggested that these represent respectively vestiges of a prepollex and a post-minimus digit[172].
One or more of the five digits commonly present may be lost, and sometimes all are lost except the third. The terminal or ungual phalanges of the digits are commonly specially modified to support nails, claws, or hoofs. There are as a rule two small sesamoid bones developed on the ventral or flexor side of the metacarpo-phalangeal articulations, and sometimes similar bones occur on the dorsal or extensor side.
Monotremata.InEchidnathe carpus is broad, the scaphoid and lunar are united and there is no centrale. The pisiform is large and several other sesamoid bones occur. Each of the five digits is terminated by a large ungual phalanx. InOrnithorhynchusthe manus is more slender, but the general arrangement is the same as inEchidna.
Marsupialia.The carpus has no centrale and the lunar is generally small or absent. Five digits are almost always present. InChoeropushowever the only two functional digits are the second and third, which have very long closely apposed metacarpals; the fourth digit is vestigial, but has the normal number of phalanges, while the first and fifth are absent. The manus inNotoryctesis extraordinarily modified, the scaphoid and all the distal carpalia are apparently fused, the first, second, and fifth digits are very small, the third and fourth, though having only one phalanx apiece, bear each an enormous claw. Lying on and obscuring the ventral surface of the manus is a large bone, probably a sesamoid.
Among theEdentatathere is a great diversity in the structure of the manus, the centrale is however always wanting, and except inManisthe scaphoid and lunar are distinct.In the Sloths the manus is very long, narrow, and curved, and terminated by two or three long hooked claws, borne by the second and third, or the second, third and fourth digits. The fifth digit is absent, and the fourth is represented only by a small metacarpal. In the Anteaters the third digit is greatly developed and bears a long hooked claw. InMyrmecophagaall five digits are fairly well though irregularly developed, inCycloturusthe first, fourth, and fifth, are vestigial. In the Armadillos the manus is broad, and has strongly developed ungual phalanges. The digits, though almost always five in number, vary much in their relative arrangement. InDasypusthey are regular, but are remarkably irregular in Priodon. The pollex is absent in Glyptodonts and inMegatherium.InMegatheriumthe fifth digit is clawless while the second, third, and fourth bear enormous claws. In the Manidae the scaphoid and lunar are united; five digits are present, the third and fourth being very large, and all being terminated by deeply cleft ungual phalanges. InOrycteropusthe pollex is absent, while the other digits are terminated by pointed ungual phalanges.
InSireniathe general structure of the manus is quite of the ordinary mammalian type. InManatusmost of the bones of the carpus are distinct, but inHalicoremany, especially those of the distal row, have coalesced. The digits are always five in number and have the normal number of flattened phalanges.
In theCetacea, on the other hand, the manus is much modified by the fact that the number of phalanges may be greatly increased above the normal number of three, thirteen or fourteen sometimes occurring in each digit. These are believed to be duplicated epiphyses. In the Mystacoceti the manus remains largely cartilaginous, in the Odontoceti it is better ossified, and the phalanges commonly have epiphyses at both ends. InPhyseterthe carpal bones also have epiphyses. The carpus generally consists of six bones arranged in tworows of three each. Five digits are generally present, but sometimes as inBalaenoptera musculus, there are four, the third being suppressed. Their relative development varies much. The Sperm Whale which till recently was placed in the entrance hall of the Natural History Museum at South Kensington has one phalanx to the first digit, four to the second, five to the third, four to the fourth, and three to the fifth. Generally the manus is short and broad, but sometimes, as inGlobicephalus, it is much elongated owing to the great development of the second and third digits.
Ungulata[173]. The manus of the members of this great order is of very great classificatory and morphological importance. All the members agree in having the scaphoid and lunar distinct, and in almost every case the ends of the digits are either encased in hoofs or provided with broad flat nails. It is by means of characters derived from the manus and pes that the group is subdivided into the Ungulata vera and the Subungulata.
In theUngulata verathe manus is never plantigrade, and there are not more than four digits, the pollex being almost always completely suppressed: inCotylopsamong extinct Artiodactyla however a vestigial pollex is found. The centrale is absent, and the magnum articulates freely with the scaphoid, and is separated from the cuneiform by the unciform and lunar. All the bones of the carpus interlock strongly, and the axis of the third digit passes through the magnum and between the scaphoid and lunar.
There is a very strong distinction between the manus of the suborders Artiodactyla and Perissodactyla. In the Artiodactyla the axis of the manus passes between the third and fourth digits, which are almost equally developed and, except in the Hippopotami and some extinct forms such asAnoplotherium,have their ungual phalanges flattened on their contiguous surfaces.
In allArtiodactylathe third and fourth digits are large, but a gradual reduction in the second and fifth can be well traced. Thus in the Suina the second and fifth digits, though smaller than the third and fourth, are well developed and all four metacarpals are distinct. In the Tragulina too all four metacarpals are developed, and inDorcatheriumthe third and fourth commonly remain distinct as in the Suina. In the other Artiodactyla however the third and fourth metacarpals are almost always united, though indications of their separate origin remain. In some Ruminantia, such as many Deer, the second and fifth digits are reduced to minute splint bones attached to the proximal end of the fused third and fourth metacarpals, and to small hoof-bearing phalanges, sometimes attached to splint-like distal vestiges of the metacarpals, sometimes altogether unconnected with any other skeletal structures. In some other Ruminants, such as the Sheep and Oxen, the only remnants of the second and fifth digits are nodules of bone supporting the hoofs, and in others, such as the Giraffe,Anoplotherium commune, some Antelopes and the Tylopoda, all traces of these digits have disappeared. The Camels differ from all living Ungulata vera in not having the distal phalanges completely encased in hoofs, and from all except the Hippopotami in placing a considerable amount of the manus on the ground in walking.