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diff --git a/43431-0.txt b/43431-0.txt new file mode 100644 index 0000000..a54b70b --- /dev/null +++ b/43431-0.txt @@ -0,0 +1,25815 @@ +*** START OF THE PROJECT GUTENBERG EBOOK 43431 *** + + CAMBRIDGE BIOLOGICAL SERIES. + + GENERAL EDITOR:--ARTHUR E. SHIPLEY, M.A. + + FELLOW AND TUTOR OF CHRIST'S COLLEGE, CAMBRIDGE. + + + + + THE + + VERTEBRATE SKELETON. + + + + + London: C.J. CLAY AND SONS, + CAMBRIDGE UNIVERSITY PRESS WAREHOUSE, + AVE MARIA LANE, + + AND + + H.K. LEWIS, + 136, GOWER STREET, W.C. + + [Illustration] + + Glasgow: 50, WELLINGTON STREET. + + Leipzig: F.A. BROCKHAUS. + + New York: THE MACMILLAN COMPANY. + + Bombay and Calcutta: MACMILLAN AND CO., LTD. + + + [_All Rights reserved._] + + + + + THE + + VERTEBRATE SKELETON + + BY + + SIDNEY H. REYNOLDS, M.A., + + TRINITY COLLEGE, CAMBRIDGE; + LECTURER AND DEMONSTRATOR IN GEOLOGY AND ZOOLOGY AT UNIVERSITY + COLLEGE, BRISTOL. + + Cambridge: + + AT THE UNIVERSITY PRESS. + + 1897 + + [_All Rights reserved._] + + + + + =Cambridge=: + + PRINTED BY J. & C.F. CLAY, + AT THE UNIVERSITY PRESS. + + + + +PREFACE. + + +IN the following pages the term skeleton is used in its widest sense, +so as to include exoskeletal or tegumentary structures, as well as +endoskeletal structures. It was thought advisable to include some +account of the skeleton of the lowest Chordata--animals which are not +strictly vertebrates, but it seemed undesirable to alter the title of +the book in consequence. + +The plan adopted in the treatment of each group has been to give first +an account of the general skeletal characters of the group in question +and of its several subdivisions; secondly to describe in detail the +skeleton of one or more selected types; and thirdly to treat the +skeleton as developed in the group organ by organ. + +A beginner is advised to commence, not with the introductory chapter, +but with the skeleton of the Dogfish, then to pass to the skeletons of +the Newt and Frog, and then to that of the Dog. After that he might +pass to the introductory chapter and work straight through the book. I +have endeavoured to make the account of each type skeleton complete in +itself; this has necessitated a certain amount of repetition,--a +fault that I have found it equally difficult to avoid in other parts +of the book. + +Throughout the book generic names are printed in italics; and italics +are used in the accounts of the type skeletons for the names of +membrane bones. Clarendon type is used to emphasise certain words. In +the classificatory table the names of extinct genera only, are printed +in italics. + +In a book in which an attempt is made to cover to some extent such a +vast field, it would be vain to hope to have avoided many errors both +of omission and commission, and I owe it to the kindness of several +friends that the errors are not much more numerous. I cannot however +too emphatically say that for those which remain I alone am +responsible. Messrs C.W. Andrews, E. Fawcett, S.F. Harmer, J. Graham +Kerr, and B. Rogers have all been kind enough to help me by reading +proofs or manuscript, while the assistance that I have received from +Dr Gadow during the earlier stages and from Prof. Lloyd Morgan and Mr +Shipley throughout the whole progress of the work has been very great. +To all these gentlemen my best thanks are tendered. + +All the figures except 1, 35, 55, and 84 were drawn by Mr Edwin +Wilson, to whose care and skill I am much indebted. The majority are +from photographs taken by my sister Miss K.M. Reynolds or by myself in +the British Museum and in the Cambridge University Museum of Zoology, +and I take this opportunity of thanking Sir W.H. Flower and Mr S.F. +Harmer for the facilities they have afforded and for permission to +figure many objects in the museums respectively under their charge. I +have also to thank (1) Prof. von Zittel for permission to reproduce +figs. 27, 41, 52, 69, 70, 80, 106 A, and 107 C; (2) Sir W.H. Flower +and Messrs A. and C. Black for figs. 1 and 84; (3) Prof. O.C. Marsh +and Dr H. Woodward for fig. 35; (4) Dr C.H. Hurst and Messrs Smith, +Elder, and Co. for fig. 55. + +A few references are given, but no attempt has been made to give +anything like a complete list. The abbreviations of the titles of +periodicals are those used in the _Zoological Record_. + +I have always referred freely to the textbooks treating of the +subjects dealt with, and in particular I should like to mention that +the section devoted to the skeleton of mammals is, as it could hardly +fail to be, to a considerable extent based on Sir W.H. Flower's +_Osteology of the Mammalia_. + + SIDNEY H. REYNOLDS. + + _March 10, 1897._ + + + + + CONTENTS. + + + PAGE + CHAPTER I. + + Introductory account of the skeleton in general 1 + + + CHAPTER II. + + Classification 30 + + + CHAPTER III. + + Skeleton of Hemichordata, Urochordata and Cephalochordata 50 + + + CHAPTER IV. + + Skeletal characters of the Vertebrata. The skeleton in the + Cyclostomata 53 + + + CHAPTER V. + + Skeletal characters of the Ichthyopsida. Characters of the + several groups of Pisces 59 + + + CHAPTER VI. + + The skeleton of the Dogfish (Scyllium canicula) 71 + + + CHAPTER VII. + + The skeleton of the Codfish (_Gadus morrhua_) and the skull of + the Salmon (_Salmo salar_) 83 + + + CHAPTER VIII. + + General account of the skeleton in Fishes 104 + + + CHAPTER IX. + + Characters of the several groups of Amphibia 133 + + + CHAPTER X. + + The skeleton of the Newt (_Molge cristata_) 138 + + + CHAPTER XI. + + The skeleton of the Frog (_Rana temporaria_) 151 + + + CHAPTER XII. + + General account of the skeleton in Amphibia 168 + + + CHAPTER XIII. + + Skeletal characters of the Sauropsida. Characters of the + several groups of Reptiles 189 + + + CHAPTER XIV. + + The skeleton of the Green Turtle (_Chelone midas_) 214 + + + CHAPTER XV. + + The skeleton of the Crocodile (_Crocodilus palustris_) 237 + + + CHAPTER XVI. + + General account of the skeleton in Reptiles 270 + + + CHAPTER XVII. + + Characters of the several groups of Birds 295 + + + CHAPTER XVIII. + + The skeleton of the Wild Duck (_Anas boschas_) 302 + + + CHAPTER XIX. + + General account of the skeleton in Birds 328 + + + CHAPTER XX. + + Characters of the several groups of Mammalia 343 + + + CHAPTER XXI. + + The skeleton of the Dog (_Canis familiaris_) 374 + + + CHAPTER XXII. + + General account of the skeleton in Mammalia. The exoskeleton + and vertebral column 416 + + + CHAPTER XXIII. + + General account of the skeleton in Mammalia (_continued_). The + skull and appendicular skeleton 455 + + + + + LIST OF ILLUSTRATIONS. + + + FIG. PAGE + + 1 Diagrammatic sections of various forms of teeth 6 + + 2 Cervical vertebrae of an Ox (_Bos taurus_) 15 + + 3 Diagram of the skeleton of _Amphioxus lanceolatus_ 51 + + 4 Dorsal, lateral, and ventral views of the skull of + _Petromyzon marinus_ 56 + + 5 Skull of a male _Chimaera monstrosa_ 65 + + 6 Lateral view of the skull of a Dogfish (_Scyllium canicula_) 75 + + 7 Semidorsal view of the pectoral girdle and fins of a Dogfish + (_Scyllium canicula_) 80 + + 8 Dorsal view of the pelvic girdle and fins of a male Dogfish + (_Scyllium canicula_) 81 + + 9 Dorsal and ventral views of the cranium of a Salmon (_Salmo + salar_) from which most of the membrane bones have been + removed 88 + + 10 Lateral view of the chondrocranium of a Salmon (_Salmo + salar_) 90 + + 11 Lateral view of the skull of a Salmon (_Salmo salar_) 92 + + 12 Mandibular and hyoid arches of a Cod (_Gadus morrhua_) 99 + + 13 Right half of the pectoral girdle and right pectoral fin of + a Cod (_Gadus morrhua_) 102 + + 14 Diagram of a section through the jaw of a Shark + (_Odontaspis americanus_) showing the succession of + teeth 107 + + 15 Part of the lower jaw of a Shark (_Galeus_) 108 + + 16 Skulls of _Notidanus_ and _Cestracion_ 118 + + 17 Dorsal view of the branchial arches of _Heptanchus_ 120 + + 18 Lateral view of the skull of a Sturgeon + (_Acipenser sturio_) 122 + + 19 Dorsal and ventral views of the cranium of + _Ceratodus miolepis_ 125 + + 20 Lateral view of the skeleton of _Ceratodus miolepis_ 128 + + 21 Dorsal, ventral and lateral views of the skull of a Newt + (_Molge cristata_) 142 + + 22 Ventral and lateral views of the shoulder-girdle and + sternum of an old male Crested Newt (_Molge cristata_) 146 + + + 23 Right posterior and anterior limbs of a Newt (_Molge + cristata_) 148 + + 24 Dorsal and ventral views of the cranium of a Common Frog + (_Rana temporaria_) 155 + + 25 Dorsal and ventral views of the cranium of a Common Frog + (_Rana temporaria_) from which the membrane bones have + mostly been removed 157 + + 26 Lateral view of the skull and posterior view of the + cranium of a Common Frog (_Rana temporaria_) 159 + + 27 Dorsal view of the skull of a Labyrinthodont (_Capitosaurus + nasutus_) 176 + + 28 Ventral view of the cranium, and lateral view of the + cranium and mandible of _Siphonops annulatus_ 178 + + 29 Visceral arches of Amphibia: A, _Molge cristata_; B, _Rana + temporaria_, adult; C, Tadpole of _Rana_; D, _Siredon + pisciformis_ 181 + + 30 Shoulder-girdle and sternum of an adult male Common Frog + (_Rana temporaria_), and of an adult female + _Docidophryne gigantea_ 183 + + 31 A, Right antibrachium and manus of a larval Salamander + (_Salamandra maculosa_); B, Right tarsus and adjoining + bones of _Molge sp._ 186 + + 32 Lateral and dorsal views of the skull of an _Ichthyosaurus_ 196 + + 33 Lateral view and longitudinal section of the skull of a + Lizard (_Varanus varius_) 201 + + 34 Lateral view of the shoulder-girdle of a Lizard (_Varanus_) 202 + + 35 Restored skeleton of _Ceratosaurus nasicornis_ 206 + + 36 Dorsal and ventral views of the carapace of a Loggerhead + Turtle (_Thalassochelys caretta_) 216 + + 37 Plastron of a Green Turtle (_Chelone midas_) 218 + + 38 The skull of a Green Turtle (_Chelone midas_) 223 + + 39 Longitudinal vertical section through the cranium of a + Green Turtle (_Chelone midas_) 226 + + 40 Anterior limb of a young Hawksbill Turtle (_Chelone + imbricata_), and posterior limb of a large Green Turtle + (_Chelone midas_) 234 + + 41 The first four cervical vertebrae of a Crocodile + (_Crocodilus vulgaris_) 239 + + 42 Anterior view of a late thoracic and the first sacral + vertebrae of a Crocodile (_Crocodilus palustris_) 242 + + + 43 Palatal aspect of the cranium and mandible of an Alligator + (_Caiman latirostris_) 245 + + 44 Lateral view of the skull of an Alligator (_Caiman + latirostris_) 248 + + 45 Longitudinal section through the skull of an Alligator + (_Caiman latirostris_) 253 + + 46 Sternum and associated membrane bones of a Crocodile + (_Crocodilus palustris_) 261 + + 47 Left half of the pectoral girdle of an Alligator + (_Caiman latirostris_) 262 + + 48 Right anterior and posterior limbs of an Alligator + (_Caiman latirostris_) 264 + + 49 Pelvis and sacrum of an Alligator (_Caiman latirostris_) 267 + + 50 Preparation of part of the right mandibular ramus of + _Crocodilus palustris_ 274 + + 51 Dorsal and ventral views of the skull of a Common Snake + (_Tropidonotus natrix_) 279 + + 52 Skull of Hatteria (_Sphenodon punctatus_) 282 + + 53 Hyoids of an Alligator (_Caiman latirostris_), and of a + Green Turtle (_Chelone midas_) 285 + + 54 Ventral view of the shoulder-girdle and sternum of + _Loemanctus longipes_ 287 + + 55 Left half of the skeleton of a Common Fowl (_Gallus + bankiva_) 301 + 56 The wing of a Wild Duck (_Anas boschas_) 304 + + 57 Wings of a Wild Duck with the coverts removed (_Anas + boschas_) 305 + + 58 Dorsal and ventral views of the pelvis and sacrum of a Duck + (_Anas boschas_) 311 + + 59 Skull of a Duck (_Anas boschas_) 312 + + 60 A, Ventral view of the cranium of a Duck (_Anas boschas_); + B, Cranium and mandible seen from the left side 313 + + 61 Lateral view of the pelvis and sacrum of a Duck (_Anas + boschas_) 325 + 62 Third cervical vertebra of an Ostrich (_Struthio camelus_) 331 + + 63 Shoulder-girdle and sternum of A, Black Vulture (_Vultur + cinereus_); B, Peacock (_Pavo cristatus_); C, Pelican + (_Pelicanus conspicillatus_) 337 + + 64 Bones of the right wing of A, a Penguin; B, an Ostrich + (_Struthio camelus_) and C, a Gannet (_Sula alba_) 339 + + 65 Pelvic girdle and sacrum of A, Cassowary (_Casuarius + galeatus_); B, Owen's Apteryx (_A. oweni_); C, + Broad-billed Rhea (_R. macrorhyncha_); D, Ostrich + (_Struthio camelus_) 340 + + 66 Ventral view of the shoulder-girdle and sternum of a + Duckbill (_Ornithorhynchus paradoxus_) 347 + + 67 Cervical vertebrae of a Ca'ing Whale (_Globicephalus + melas_) 354 + + 68 Dentition of a Dog (_Canis familiaris_) 375 + + 69 Atlas and axis vertebrae of a Dog (_Canis familiaris_) 379 + + 70 Second thoracic and second lumbar vertebrae of a Dog + (_Canis familiaris_) 382 + + 71 Diagram of the relations of the principal bones in the + Mammalian skull 385 + + 72 Vertical longitudinal section through skull of a Dog + (_Canis familiaris_) 387 + + 73 Dorsal view of the cranium of a Dog (_Canis familiaris_) 389 + + 74 Diagram of the mammalian tympanic cavity and associated + bones 391 + + 75 Ventral view of the cranium of a Dog (_Canis familiaris_) 396 + + 76 Sternum and sternal ribs of a Dog (_Canis familiaris_) 403 + + 77 Bones of the left upper arm and fore-arm of a Dog (_Canis + familiaris_) 407 + + 78 Right innominate bone, A, of a full-grown Terrier; B, of a + Collie Puppy 410 + + 79 Left leg bones of a Dog (_Canis familiaris_) 411 + + 80 A, Right manus; B, Right pes of a Dog (_Canis familiaris_) 413 + + 81 Skull of a young Indian Rhinoceros (_R. unicornis_) showing + the change of the dentition 421 + + 82 Palatal aspect of the cranium and mandible of a Donkey + (_Equus asinus_) 431 + + 83 Skull of _Procavia (Dendrohyrax) dorsalis_ 433 + + 84 Carnassial or sectorial teeth of Carnivora 436 + + 85 Mandible of Isabelline Bear (_Ursus isabellinus_) 438 + + 86 Left mandibular ramus of the Sea Leopard (_Ogmorhinus + leptonyx_) 439 + + 87 Cervical vertebrae of a young Fin Whale (_Balaenoptera + musculus_) 444 + + 88 Atlas and axis vertebrae of an Ox (_Bos taurus_) 445 + + 89 First and second thoracic vertebrae of an Ox (_Bos taurus_) 449 + + 90 Skulls of Tasmanian Wolf (_Thylacinus cynocephalus_) and + Hairy-nosed Wombat (_Phascolomys latifrons_) 456 + + 91 Skull of Two-fingered Sloth (_Choloepus didactylus_) 458 + + 92 Skull of _Rhytina stelleri_ 460 + + + 93 Lateral view and longitudinal section of the skull of a + young Ca'ing Whale (_Globicephalus melas_) 463 + + 94 Cranium and mandible of a Pig (_Sus scrofa_) 466 + + 95 Mandible of a Hippopotamus (_Hippopotamus amphibius_) 467 + + 96 Skull of a young Indian Elephant (_Elephas indicus_) 474 + + 97 Longitudinal section of the skull of a young Indian + Elephant (_Elephas indicus_) 475 + + 98 Half-front view of the skull of a Porcupine (_Hystrix + cristata_) 477 + 99 Skulls of an old and of a young Gorilla (_Gorilla savagei_) 483 + + 100 Malleus, stapes, and incus of Man, Dog, and Rabbit 485 + + 101 Skeleton of a Cape Buffalo (_Bubalus caffer_) 492 + + 102 Lateral and dorsal views of the shoulder-girdle and part + of the sternum of the Spiny Anteater (_Echidna + aculeata_) 494 + + 103 Skeleton of a Llama (_Auchenia glama_) 496 + + 104 Dorsal view of the sternum and right half of the shoulder + girdle of _Mus sylvaticus_ 498 + + 105 Anterior surface of the right humerus of a Wombat + (_Phascolomys latifrons_) 500 + + 106 Manus of Perissodactyles: A, Left manus of _Tapirus_; B, + Right manus of _Titanotherium_; C, Left manus of + _Chalicotherium giganteum_ 508 + + 107 Left manus of A, _Coryphodon hamatus_; B, _Phenacodus + primaevus_; C, _Procavia (Dendrohyrax) arboreus_ 510 + + 108 Left anterior and posterior limbs and limb girdles of + _Uintatherium mirabile_ 516 + + 109 Left femur of an Ox (_Bos taurus_) and of a Sumatran + Rhinoceros (_Rhinoceros sumatrensis_) 518 + + 110 Pes of A, a Tapir (_Tapirus americanus_); B, a Rhinoceros + (_Rhinoceros sumatrensis_); C, _Hipparion gracile_; D, + a Horse (_Equus caballus_) 524 + + + + +ERRATA. + + + p. 172, note, _for_ 14 _read_ 15. + + p. 279, description of figure, _for_ Tropidinotus _read_ + Tropidonotus. + + p. 287, description of figure, _for_ shoulder-girdle of sternum + _read_ shoulder-girdle and sternum. + + p. 393, middle of page, _for_ VIII _read_ VII. + + p. 427, line 2, _for_ Grampus _read_ Killer. + + + + +CHAPTER I. + +INTRODUCTORY ACCOUNT OF THE SKELETON IN GENERAL. + + +BY the term =skeleton= is meant the hard structures whose function is +to support or to protect the softer tissues of the animal body. + +The skeleton is divisible into + +A. The EXOSKELETON, which is external; + +B. The ENDOSKELETON, which is as a rule internal; though in some +cases, e.g. the antlers of deer, endoskeletal structures become, as +development proceeds, external. + +In Invertebrates the hard, supporting structures of the body are +mainly =exoskeletal=, in Vertebrates they are mainly =endoskeletal=; +but the endoskeleton includes, especially in the skull, a number of +elements, the =dermal= or =membrane= bones, which are shown by +development to have been originally of external origin. These membrane +bones are so intimately related to the true endoskeleton that they +will be described with it. The simplest and lowest types of both +vertebrate and invertebrate animals have unsegmented skeletons; with +the need for flexibility however segmentation arose both in the case +of the invertebrate exoskeleton and the vertebrate endoskeleton. The +exoskeleton in vertebrates is phylogenetically older than the +endoskeleton, as is indicated by both palaeontology and embryology. +Palaeontological evidence is afforded by the fact that all the lower +groups of vertebrates--Fish, Amphibia, and Reptiles--had in former +geological periods a greater proportion of species protected by +well-developed dermal armour than is the case at present. +Embryological evidence tends the same way, inasmuch as dermal +ossifications appear much earlier in the developing animal than do the +ossifications in the endoskeleton. + +Skeletal structures may be derived from each of the three germinal +layers. Thus =hairs= and =feathers= are =epiblastic= in origin, +=bones= are =mesoblastic=, and the =notochord= is =hypoblastic=. + +The different types of skeletal structures may now be considered and +classified more fully. + + +A. EXOSKELETAL STRUCTURES. + +I. EPIBLASTIC (epidermal). + +Exoskeletal structures of epiblastic origin may be developed on both +the inner and outer surfaces of the Malpighian layer of the +epidermis[1]. Those developed on the outer surface include =hairs=, +=feathers=, =scales=, =nails=, =beaks= and =tortoiseshell=; and are +specially found in vertebrates higher than fishes. Those developed on +the inner surface of the Malpighian layer include only the =enamel= of +teeth and some kinds of scales. With the exception of feathers, which +are partly formed from the horny layer, all these parts are mainly +derived from the Malpighian layer of the epidermis. + +=Hairs= are slender, elongated structures which arise by the +proliferation of cells from the Malpighian layer of the epidermis. +These cells in the case of each hair form a short papilla, which sinks +inwards and becomes imbedded at the bottom of a follicle in the +dermis. Each hair is normally composed of an inner cellular pithy +portion containing much air, and an outer denser cortical portion of a +horny nature. Sometimes, as in Deer, the hair is mainly formed of the +pithy portion, and is then easily broken. Sometimes the horny part +predominates, as in the bristles of Pigs. A highly vascular dermal +papilla projects into the base of the hair. + +=Feathers=, like hairs, arise from epidermal papillae which become +imbedded in pits in the dermis. But the feather germ differs from the +hair germ, in the fact that it first grows out like a cone on the +surface of the epidermis, and that the horny as well as the Malpighian +layer takes part in its formation. + +=Nails=, =claws=, =hoofs=, and the =horns of Oxen= are also epidermal, +as are such structures as the =scales= of reptiles, of birds' feet, +and of _Manis_ among mammals, the =rattle= of the rattlesnake, the +=nasal horns= of _Rhinoceros_, and the =baleen= of whales. All these +structures will be described later. + +=Nails= arise in the interior of the epidermis by the thickening and +cornification of the stratum lucidum. The outer border of the nail +soon becomes free, and growth takes place by additions to the inner +surface and attached end. + +When a nail tapers to a sharp point it is called a =claw=. In many +cases the nails more or less surround the ends of the digits by which +they are borne. + +Horny =beaks= of epidermal origin occur casing the jaw-bones in +several widely distinct groups of animals. Thus among reptiles they +are found in Chelonia (tortoises and turtles) as well as in some +extinct forms; they occur in all living birds, in _Ornithorhynchus_ +among mammals, and in the larvae of many Amphibia. + +In a few animals, such as Lampreys and _Ornithorhynchus_, the jaws +bear horny tooth-like structures of epidermal origin. + +The =enamel= of teeth and of placoid scales is also epiblastic in +origin[2], and it may be well at this point to give some account of +the structure of teeth, though they are partly mesoblastic in origin. +The simplest teeth are those met with in sharks and dogfish, where +they are merely the slightly modified scales developed in the +integument of the mouth. They pass by quite insensible gradations into +normal placoid scales, such as cover the general surface of the body. +A =placoid scale=[3] is developed on a papilla of the dermis which +projects outwards and backwards, and is covered by the columnar +Malpighian layer of the epidermis. The outer layer of the dermal +papilla then gradually becomes converted into dentine and bone, while +enamel is developed on the inner side of the Malpighian layer, forming +a cap to the scale. The Malpighian and horny layers of the epidermis +get rubbed off the enamel cap, so that it comes to project freely on +the surface of the body. + +As regards their attachment teeth may be (1) attached to the fibrous +integument of the mouth, or (2) fixed to the jaws or other bones of +the mouth, or (3) planted in grooves, or (4) in definite sockets in +the jaw-bones (see p. 107). + +Teeth in general consist of three tissues, =enamel=, =dentine= and +=cement=, enclosing a central pulp-cavity containing blood-vessels and +nerves. Enamel is, however, often absent, as in all living Edentates. + +=Enamel= generally forms the outermost layer of the crown or visible +part of the tooth; it is the hardest tissue occurring in the animal +body and consists of prismatic fibres arranged at right angles to the +surface of the tooth. It is characterised by its bluish-white +translucent appearance. + +II. MESOBLASTIC (mesodermal). + +=Dentine= or =ivory= generally forms the main mass of a tooth. It is a +hard, white substance allied to bone. When examined microscopically +dentine is seen to be traversed by great numbers of nearly parallel +branching tubules which radiate outwards from the pulp-cavity. In +fishes as a rule, and sometimes in other animals, a variety of dentine +containing blood-vessels occurs, this is called =vasodentine=. + +=Cement= or =crusta petrosa= forms the outermost layer of the root of +the tooth. In composition and structure it is practically identical +with bone. In the more complicated mammalian teeth, besides enveloping +the root, it fills up the spaces between the folds of the enamel. + +The hard parts of a tooth commonly enclose a central pulp-cavity into +which projects the pulp, a papilla of the dermis including +blood-vessels and nerves. As long as growth continues the outer layers +of this pulp become successively calcified and added to the substance +of the dentine. In young growing teeth the pulp-cavity remains widely +open, but in mammals the general rule is that as a tooth gets older +and the crown becomes fully formed, the remainder of the pulp becomes +converted into one or more tapering roots which are imbedded in the +alveolar cavities of the jaws. The opening of the pulp-cavity is then +reduced to a minute perforation at the base of each root. A tooth of +this kind is called a =rooted= tooth. + +But it is not only in young teeth that the pulp-cavity sometimes +remains widely open; for some teeth, such as the tusks of Elephants +and the incisor teeth of Rodents, form no roots and continue to grow +throughout the animal's life. Such teeth are said to be rootless or to +have persistent pulps. + +An intermediate condition is seen in some teeth, such as the grinding +teeth of Horses. These teeth grow for a very long time, their crowns +wearing away as fast as their bases are produced; finally however +definite roots are formed and growth ceases. + +[Illustration FIG. 1. DIAGRAMMATIC SECTIONS OF VARIOUS FORMS OF TEETH +(from FLOWER). + +I. Incisor or tusk of elephant, with pulp-cavity persistently open at +base. II. Human incisor during development with root imperfectly +formed, and pulp-cavity widely open at base. III. Completely formed +human incisor, with pulp-cavity contracted to a small aperture at the +end of the root. IV. Human molar with broad crown and two roots. V. +Molar of Ox, with the enamel covering the crown, deeply folded and the +depressions filled with cement. The surface is worn by use, otherwise +the enamel coating would be continuous at the top of the ridges. In +all the figures the enamel is black, the pulp white, the dentine +represented by horizontal lines, and the cement by dots.] + +The teeth of any animal may be =homodont=, that is, all having the +same general character, or =heterodont=, that is, having different +forms adapted to different functions. The dentition is heterodont in a +few reptiles and the majority of mammals. + +SUCCESSION OF TEETH. In most fishes, and many amphibians and reptiles +the teeth can be renewed indefinitely. In sharks, for example, +numerous rows of reserve teeth are to be seen folded back behind those +in use (see fig. 15). The majority of mammals have only two sets of +teeth, and are said to be =diphyodont=; some have only a single series +(=monophyodont=). + +DEVELOPMENT OF TEETH. A brief sketch of the method in which +development of teeth takes place in the higher vertebrates may here be +given. Along the surface of the jaws a thickening of the epiblastic +epithelium takes place, giving rise to a ridge, which sinks inwards +into the tissue of the jaw, and it is known as the primary enamel +organ. At the points where teeth are to be developed special ingrowths +of this primary enamel organ take place, and into each there projects +a vascular dental papilla from the surrounding mesoblast of the jaw. +Each ingrowth of the enamel organ forms an =enamel cap=, which +gradually embraces the dental papilla, and at the same time appears to +be pushed on one side, owing to the growth not being uniform. The +external layer of the dental papilla is composed of long nucleated +cells, the =odontoblasts=, and it is by these that the dentine is +formed. Similarly the internal layer of the enamel organ is formed of +columnar enamel cells, which give rise to the enamel. The mesoblastic +cells surrounding the base of the tooth give rise to the cement. + +=Bone= is in many cases exoskeletal, but it will be most conveniently +described with the endoskeleton. + +The =scales of fish= are wholly or in part mesoblastic in origin, +being totally different from those of reptiles. The =cycloid= and +=ctenoid= scales of Teleosteans (see p. 105) are thin plates coated +with epidermis. They are sometimes bony, but as a rule are simply +calcified. =Ganoid= scales are flat plates of bone coated with an +enamel-like substance, and articulating together with a peg and socket +arrangement; they are probably identical with enlarged and flattened +placoid scales. + +The =armour plates= of fossil Ganoids, Labyrinthodonts, and Dinosaurs, +and of living Crocodiles, some Lizards and Armadillos, are composed of +bone. They are always covered by a layer of epidermis. + +The =antlers of deer= are also composed of bone; they will be more +fully described in the chapter on mammals. It may perhaps be well to +mention them here, though they really belong to the endoskeleton, +being outgrowths from the frontal bones. + + +B. ENDOSKELETAL STRUCTURES. + +I. HYPOBLASTIC. + +(_a_) The =notochord= is an elastic rod formed of large vacuolated +cells, and is surrounded by a membranous sheath of mesoblastic origin. +It is the primitive endoskeleton in the Chordata, all of which possess +it at some period of their existence; while in many of the lower forms +it persists throughout life. Even in the highest Chordata it is the +sole representative of the axial skeleton for a considerable part of +the early embryonic life. A simple unsegmented notochord persists +throughout life in the Cephalochordata, Cyclostomata, and some Pisces, +such as Sturgeons and Chimaeroids. + +(_b_) The enamel of the pharyngeal teeth of the Salmon and many other +Teleosteans is hypoblastic in origin. The epiblast of the stomodaeum, +in which the other teeth are developed, passes into the hypoblast of +the mesenteron in which these pharyngeal teeth are formed. + + +II. MESOBLASTIC. + +The most primitive type of a mesoblastic endoskeleton consists of a +membranous sheath surrounding the notochord, as in _Myxine_ and its +allies. The first stage of complication is by the development of +cartilage in the notochordal sheath, as in _Petromyzon_. Often the +cartilage becomes calcified in places, as in the vertebral centra of +_Scyllium_ and other Elasmobranchs. Lastly, the formation of bone +takes place; it generally constitutes the most important of the +endoskeletal structures. + +=Bone= may be formed in two ways:-- + +(1) by the direct ossification of pre-existing cartilage, when it is +known as =cartilage bone= or =endochondral bone=; + +(2) by independent ossification in connective tissue; it is then known +as =membrane= or =dermal= or =periosteal bone=. + +With the exception of the _clavicle_[4] all the bones of the trunk and +limbs, together with a large proportion of those of the skull, are +preformed in the embryo in cartilage, and are grouped as cartilage +bones; while the clavicle and most of the roofing and jaw-bones of the +skull are not preformed in cartilage, being developed simply in +connection with a membrane. Hence it is customary to draw a very +strong line of distinction between these two kinds of bone; in reality +however this distinction is often exaggerated, and the two kinds pass +into one another, and as will be shown immediately, the permanent +osseous tissue of many of those which are generally regarded as +typical cartilage bones, is really to a great extent of periosteal +origin. The palatine bone, for instance, of the higher vertebrates in +general is preceded by a cartilaginous bar, but is itself almost +entirely a membrane bone. + +Before describing the development of bone it will be well to briefly +describe the structure of adult bone and cartilage. + +The commonest kind of =cartilage=, and that which preforms so many of +the bones of the embryo, is =hyaline= cartilage. It consists of oval +nucleated cells occupying cavities (=lacunae=) in a clear +intercellular semitransparent matrix, which is probably secreted by +the cells. Sometimes one cell is seen in each lacuna, sometimes +shortly after cell-division a lacuna may contain two or more cells. +The free surface of the cartilage is invested by a fibrous membrane, +the =perichondrium=. + +=Bone= consists of a series of lamellae of ossified substance between +which are oval spaces, the =lacunae=, giving rise to numerous fine +channels, the =canaliculi=, which radiate off in all directions. The +lacunae are occupied by the =bone cells= which correspond to cartilage +cells, from which if the bone is young, processes pass off into the +canaliculi. It is obvious that the ossified substance of bone is +intercellular in character, and corresponds to the matrix of +cartilage. + +Bone may be compact, or loose and spongy in character, when it is +known as =cancellous bone=. In compact bone many of the lamellae are +arranged concentrically round cavities, the =Haversian canals=, which +in life are occupied by blood-vessels. Each Haversian canal with its +lamellae forms a =Haversian system=. In spongy bone instead of +Haversian canals there occur large irregular spaces filled with +marrow, which consists chiefly of blood-vessels and fatty tissue. The +centre of a long bone is generally occupied by one large continuous +marrow cavity. The whole bone is surrounded by a fibrous connective +tissue membrane, =the periosteum=. + +THE DEVELOPMENT OF BONE. + +=Periosteal ossification.= An example of a bone entirely formed in +this way is afforded by the parietal. The first trace of ossification +is shown by the appearance, below the membrane which occupies the +place of the bone in the early embryo, of calcareous spicules of bony +matter, which are laid down round themselves by certain large cells, +the =osteoblasts=. These osteoblasts gradually get surrounded by the +matter which they secrete and become converted into bone cells, and in +this way a mass of spongy bone is gradually produced. Meanwhile a +definite periosteum has been formed round the developing bone, and on +its inner side fresh osteoblasts are produced, and these with the +others gradually render the bone larger and more and more compact. +Finally, the middle layer of the bone becomes again hollowed out and +rendered spongy by the absorption of part of the bony matter. + +=Endochondral ossification=[5]. This is best studied in the case of a +long bone like the femur or humerus. Such a long bone consists of a +shaft, which forms the main part, and two terminal portions, which +form the =epiphyses=, or portions ossifying from centres distinct from +that forming the shaft or main part of the bone. + +In the earliest stage the future bone consists of hyaline cartilage +surrounded by a vascular sheath, the perichondrium. + +Then, starting from the centre, the cartilage becomes permeated by a +number of channels into which pass vessels from the perichondrium and +osteoblasts. In this way the centre of the developing shaft becomes +converted into a mass of cavities separated by bands or trabeculae of +cartilage. This cartilage next becomes calcified, but as yet is not +converted into true bone. The osteoblasts in connection with the +cavities now begin to deposit true endochondral spongy bone, and then +after a time this becomes absorbed by certain large cells, the +osteoclasts, and resolved into marrow or vascular tissue loaded with +fat. So that the centre of the shaft passes from the condition of +hyaline cartilage to that of calcified cartilage, thence to the +condition of spongy bone, and finally to that of marrow. At the same +time beneath the perichondrium osteoblasts are developed which also +begin to give rise to spongy bone. The perichondrium thus becomes the +periosteum, and the bone produced by it, is periosteal or membrane +bone. So that while a continuous marrow cavity is gradually being +formed in the centre of the shaft, the layer of periosteal bone round +the margin is gradually thickening, and becoming more and more compact +by the narrowing down of its cavities to the size of Haversian canals. +The absorption of endochondral and formation of periosteal bone goes +on, till in time it comes about that the whole of the shaft, except +its terminations, is of periosteal origin. At the extremities of the +shaft, however, and at the epiphyses, each of which is for a long time +separated from the shaft by a pad of cartilage, the ossification is +mainly endochondral, the periosteal bone being represented only by a +thin layer. + +Until the adult condition is reached and growth ceases, the pad of +cartilage between the epiphysis and the shaft continues to grow, its +outer (epiphysial) half growing by the formation of fresh cartilage as +fast as its inner half is encroached on by the growth of bone from the +shaft. The terminal or articular surfaces of the bone remain +throughout life covered by layers of articular cartilage. + +Even after the adult condition is reached the bone is subject to +continual change, processes of absorption and fresh formation going on +for a time and tending to render the bone more compact. + +METHODS IN WHICH BONES ARE UNITED TO ONE ANOTHER. + +The various bones composing the endoskeleton are united to one another +either by =sutures= or by movable =joints=. + +When two bones are suturally united, their edges fit closely together +and often interlock, being also bound together by the periosteum. + +In many cases this sutural union passes into fusion or =ankylosis=, +ossification extending completely from one bone to the other with the +obliteration of the intervening suture. This feature is especially +well marked in the cranium of most birds. + +The various kinds of joints or articulations[6] may be subdivided into +imperfect joints and perfect joints. + +In =imperfect joints=, such as the intervertebral joints of mammals, +the two contiguous surfaces are united by a mass of fibrous tissue +which allows only a limited amount of motion. + +In =perfect joints= the contiguous articular surfaces are covered with +cartilage, and between them lies a synovial membrane which secretes a +viscid lubricating fluid. + +The amount of motion possible varies according to the nature of the +articular surfaces; these include-- + +_a._ =ball and socket joints=, like the hip and shoulder, in which the +end of one bone works in a cup provided by another, and movements can +take place in a variety of planes. + +_b._ =hinge joints=, like the elbow and knee, in which as in +ball-and-socket joints one bone works in a cup provided by another, +but movements can take place in one plane only. + + +THE ENDOSKELETON. + +The endoskeleton is divisible into =axial= and =appendicular= parts; +and the =axial= skeleton into-- + + 1. the spinal column, + + 2. the skull {_a._ the cranium, + {_b._ the jaws and visceral skeleton, + + 3. the ribs and sternum[7]. + + +I. THE AXIAL SKELETON. + +1. THE SPINAL COLUMN. + +The spinal column in the simplest cases consists of an unsegmented +rod, the notochord, surrounded by the =skeletogenous layer=, a sheath +of mesoblastic origin, which also envelops the nerve cord. Several +intermediate stages connect this simple spinal column with the +vertebral column characteristic of higher vertebrates. A typical +vertebral column may be said to consist of (1) a series of +cartilaginous or bony blocks, the vertebral =centra=, which arise in +the sheath surrounding the notochord. They cause the notochord to +become constricted and to atrophy to a varying extent, though a +remnant of it persists, either permanently or for a long period, +within each centrum or between successive centra. (2) From the dorsal +surface of each centrum arise a pair of processes which grow round the +spinal cord and unite above it, forming a =dorsal= or =neural arch=. +(3) A similar pair of processes arising from the ventral surface of +the centrum form the =ventral= or =haemal arch=. To the ventral arch +the ribs strictly belong, and it tends to surround the ventral +blood-vessels and the body cavity with the alimentary canal and other +viscera. + +A =neural spine= or spinous process commonly projects upwards from the +dorsal surface of the neural arch, and a pair of =transverse +processes= project outwards from its sides. When, as is commonly the +case, the two halves of the haemal arch do not meet, the ventral +surface of the centrum often bears a downwardly-projecting +=hypapophysis=. + +The character of the surfaces by which vertebral centra articulate +with one another varies much. Sometimes both surfaces are concave, and +the vertebra is then said to be =amphicoelous=; sometimes a centrum is +convex in front and concave behind, the vertebra is then +=opisthocoelous=, sometimes concave in front and convex behind, when +the vertebra is =procoelous=. Again, in many vertebrae both faces of +the centra are flat, while in others they are saddle-shaped, as in the +neck vertebrae of living birds, or biconvex, as in the case of the +first caudal vertebra of crocodiles. + +In the higher vertebrates pads of fibrocartilage--the =intervertebral +discs=--are commonly interposed between successive centra, these or +parts of them often ossify, especially in the trunk and tail, and are +then known as =inter centra=. + +[Illustration FIG. 2. CERVICAL VERTEBRAE OF AN OX (_Bos taurus_). + +A, is the fifth; B, the fourth; C, the third. x 1/4 (Camb. Mus.) + + 1. neural spine. + 2. transverse process. + 3. hypapophysis. + 4. convex anterior face of the centrum. + 5. concave posterior face of the centrum. + 6. prezygapophysis. + 7. postzygapophysis. + 8. vertebrarterial canal. + 9. neural canal. + 10. inferior lamella of transverse process.] + +The vertebrae of the higher forms can generally be arranged in the +following five groups, each marked by certain special characteristics: + +1. The =cervical= or =neck vertebrae=. These connect the skull with +the thorax, and are characterised by relatively great freedom of +movement. They often bear small ribs, but are distinguished from the +succeeding thoracic vertebrae by the fact that their ribs do not reach +the sternum. The first cervical vertebra which articulates with the +skull is called the =atlas=, but a study of the nerve exits shows that +the first vertebra is not serially homologous throughout the +Ichthyopsida, so that it is best to reserve the term atlas for the +first vertebra in Sauropsida and Mammalia. + +2. The =thoracic vertebrae= (often called dorsal) bear movably +articulated ribs which unite ventrally with the sternum. + +3. The =lumbar vertebrae= are generally large, and are often more +movable on one another than are the thoracic vertebrae. They bear no +ribs. + +4. The =sacral vertebrae= are characterised by the fact that they are +firmly fused together, and are united with the pelvic girdle by means +of their transverse processes and rudimentary ribs. + +5. The =caudal= or =tail vertebrae= succeed the sacral. The anterior +ones are often fused with one another and with the sacrals, but they +differ from true sacral vertebrae in that there are no rudimentary +ribs between their transverse processes and the pelvic girdle. They +often bear V-shaped =chevron bones=. + +In fish and snakes the vertebral column is divisible into only two +regions, an anterior trunk region, whose vertebrae bear ribs, and a +posterior tail region, whose vertebrae are ribless. + + +2. THE SKULL. + +Before giving a general account of the adult skull it will be well to +briefly describe its development. + + +GENERAL DEVELOPMENT OF THE CRANIUM[8]. + +Shortly after its appearance, the central nervous system becomes +surrounded by a membranous mesodermal investment which in the region +of the spinal cord is called the =skeletogenous layer= or =perichordal +sheath=, while in the region of the brain it is called the +=membranous cranium=. Ventral to the central nervous system is the +notochord, which extends far into the region of the future cranium, +and like the nervous system, is enclosed by the skeletogenous layer. +The primitive cartilaginous cranium is formed by histological +differentiation within the substance of the membranous cranium and +always consists of the following parts: + +(_a_) the =parachordals=. These are a pair of flat curved plates of +cartilage, each of which has its inner edge grooved where it comes in +contact with the notochord. The parachordals, together with the +notochord, form a continuous plate, which is known as the =basilar +plate=. The basilar plate is the primitive floor below the hind- and +mid-brain. In front the parachordals abut upon another pair of +cartilaginous bars, the trabeculae, the two pairs of structures being +sometimes continuous with one another from the first; + +(_b_) the =trabeculae= which meet behind and embrace the front end of +the notochord. Further forwards they at first diverge from one +another, and then converge again, enclosing a space, the =pituitary +space=. After a time they generally fuse with one another in the +middle line, and, with the parachordals behind, form an almost +continuous basal plate. The trabeculae generally appear before the +parachordals. They form the primitive floor below the fore-brain; + +(_c_) the cartilaginous =capsules= of the three pairs of =sense +organs=. At a very early stage of development involutions of the +surface epiblast give rise to the three pairs of special sense +organs--the olfactory or nasal organs in front, the optic in the +middle, and the auditory behind. The olfactory and auditory organs +always become enclosed in definite cartilaginous capsules, the eyes +often as in the Salmon, become enclosed in cartilaginous sclerotic +capsules, while sometimes, as in mammals, their protecting capsules +are fibrous. + +Each pair of sense capsules comes into relation with part of the +primitive cranium, and greatly modifies it. Thus the auditory or +periotic capsules press on the parachordals till they come to be more +or less imbedded in them. Perhaps owing to the pressure of the nasal +capsules the trabeculae fuse in front, and then grow out into an +anterior pair of processes, the =cornua trabeculae=, and a posterior +pair, the =antorbital processes=, which together almost completely +surround the nasal capsules. The sclerotic capsules of the eyes +greatly modify the cranium, although they never become completely +united with it. + +The cartilaginous cranium formed of the basal plate, together with the +sense capsules, does not long remain merely as a floor. Its sides grow +vertically upwards, forming the =exoccipital= region of the cranium +behind, and the =alisphenoidal= and =orbitosphenoidal= regions further +forwards. In many forms, such as Elasmobranchs, all these upgrowths +meet round the brain, roofing it in and forming an almost complete +cartilaginous cranium. But in most vertebrata, while in the occipital +region, the cartilaginous cranium is completed dorsally, in the +alisphenoidal and orbitosphenoidal regions the cartilage merely forms +the lateral walls of the cranium, the greater part of the brain having +dorsal to it a wide space, closed by merely membranous tissue in +connection with which the large frontal and parietal bones are +subsequently formed. + + +The SKULL includes + +_a._ the cranium, + +_b._ the jaws and visceral skeleton. + +The =cranium= can be further subdivided into + +(1) an axial portion, the =cranium proper= or =brain case=; + +(2) =the sense capsules.= The capsules of the auditory and olfactory +sense organs are always present, and as has been already mentioned, +in many animals the eye likewise is included in a cartilaginous +capsule. + + +(1) THE CRANIUM PROPER OR BRAIN CASE. + +The cranium varies much in form and structure. In lower vertebrates, +such as Sharks and Lampreys, it remains entirely cartilaginous and +membranous, retaining throughout life much of the character of the +embryonic rudiment of the cranium of higher forms. The dogfish's +cranium, described on pp. 73 to 76, is a good instance of a cranium of +this type. But in the majority of vertebrates the cartilage becomes +more or less replaced by cartilage bone, while membrane bones are also +largely developed and supplant the cartilage. + +The cranium of most vertebrates includes a very large number of bones +whose arrangement varies much, but one can distinguish a definite +=basicranial axis= formed of the basi-occipital, basisphenoid, and +presphenoid bones, which is a continuation forwards of the axis of the +vertebral column. From the basicranial axis a wide arch arises, +composed of a number of bones, which form the sides and roof of the +brain-case These bones are arranged in such a manner that if both +cartilage and membrane bones are included they can be divided into +three rings or segments. The hinder one of these segments is the +occipital, the middle the parietal, and the anterior one the frontal. + +The occipital segment is formed of four cartilage bones, the +=basi-occipital= below, two =exoccipitals= at the sides, and the +=supra-occipital= above. The parietal segment is formed of the +=basisphenoid= below, two =alisphenoids= at the sides and two membrane +bones, the _parietals_ above, and the frontal segment in like manner +consists of the =presphenoid= below, the two =orbitosphenoids= at the +sides, and two membrane bones, the _frontals_, above. The parietals +and frontals, being membrane bones, are not comparable to the +supra-occipital, in the way that the presphenoid and basisphenoid are +to the basi-occipital. + +The cartilage bones of the occipital segments are derived from the +parachordals of the embryonic skull, those of the parietal and frontal +segments from the trabeculae. + +In front of the presphenoid the basicranial axis is continued by the +=mesethmoid=. + + +(2) THE SENSE CAPSULES. + +These enclose and protect the special sense organs. + +(_a_) =Auditory capsule.= + +The basisphenoid is always continuous with the basi-occipital, but the +alisphenoid is not continuous with the exoccipital as the =periotic= +or =auditory capsule= is interposed between them. Each periotic +capsule has three principal ossifications; an anterior bone, the +=pro-otic=, a posterior bone, the =opisthotic=, and a superior bone, +the =epi-otic=. + +These bones may severally unite, or instead of uniting with one +another they may unite with the neighbouring bones. Thus the epi-otic +often unites with the supra-occipital, and the opisthotic with the +exoccipital. + +Two other bones developed in the walls of the auditory capsule are +sometimes added, as in Teleosteans; these are the =pterotic= and +=sphenotic=. + +(_b_) =Optic capsule.= + +The eye is frequently enclosed in a cartilaginous sclerotic capsule, +and in this a number of scale-like bones are often developed. + +Several membrane bones are commonly formed around the orbit or cavity +for the eye. The most constant of these is the _lachrymal_ which lies +in the anterior corner; frequently too, as in Teleosteans, there is a +_supra-orbital_ lying in the upper part of the orbit, or as in many +Reptiles, a _postorbital_ lying in the posterior part of the orbit. + +(_c_) =Nasal capsule.= + +In relation to the nasal capsules various bones occur. + +The basicranial axis in front of the presphenoid is ossified, as the +=mesethmoid=, dorsal to which there sometimes, as in Teleosteans, +occur a _median ethmoid_ and a pair of =lateral ethmoids=[9]. Two +pairs of membrane bones very commonly occur in this region, viz. the +_nasals_ which lie dorsal to the mesethmoid, and the _vomers_ +(sometimes there is only one) which lie ventral to it. + +The part of the skull lying immediately in front of the cranial cavity +and in relation to the nasal capsules constitutes the =ethmoidal +region=. + +There remain certain other membrane bones which are often found +connected with the cranium. Of these, one of the largest is the +_parasphenoid_ which, in Ichthyopsids, is found underlying the +basicranial axis. _Prefrontals_ often, as in most reptiles, occur +lying partly at the sides and partly in front of the frontal, and +_postfrontals_ similarly occur behind the orbit lying partly behind +the frontals and partly at their sides. Lastly a _squamosal bone_ is, +as in Mammals, very commonly developed, and lies external and partly +dorsal to the auditory capsules. + +THE JAWS AND VISCERAL SKELETON. + +In the most primitive fish these consist of a series of cartilaginous +rings or arches placed one behind another and encircling the anterior +end of the alimentary canal. Originally they are mainly concerned with +branchial respiration. + +The first or =maxillo-mandibular= arch forms the upper jaw and the +lower jaw or mandible. + +The second or =hyoid= arch bears gills and often assists in attaching +the jaws to the cranium. The remaining arches may bear gills, though +the last is commonly without them. + +The above condition is only found in fishes, in higher animals the +visceral skeleton is greatly reduced and modified. + +The first or maxillo-mandibular arch is divisible into a dorsal +portion, the =palato-pterygo-quadrate bar=, which forms the primitive +upper jaw and enters into very close relations with the cranium, and a +ventral portion, =Meckel's cartilage=, which forms the primitive lower +jaw. The cartilaginous rudiments of both these portions disappear to a +greater or less extent and become partly ossified, partly replaced by +or enveloped in membrane bone. + +The posterior part of the palato-pterygo-quadrate bar becomes ossified +to form the =quadrate=, the anterior part to form the palatine and +pterygoid, or the two latter may be formed partially or entirely of +periosteal bone, developed round the cartilaginous bar. Two pairs of +important membrane bones, the _premaxillae_ and _maxillae_ form the +anterior part of the upper jaw, and behind the maxilla lies another +membrane bone, the _jugal_ or _malar_, which is connected with the +quadrate by a _quadratojugal_. The premaxillae have a large share in +bounding the external nasal openings or anterior nares. + +In lower vertebrates the nasal passage leads directly into the front +part of the mouth cavity and opens by the posterior nares. In some +higher vertebrates, such as mammals and crocodiles, processes arise +from the premaxillae and palatines, and sometimes from the pterygoids, +which meet their fellows in the middle line and form the palate, +shutting off the nasal passage from the mouth cavity and causing the +posterior nares to open far back. + +The cartilage of the lower jaw is in all animals with ossified +skeletons, except the Mammalia, partly replaced by cartilage bone +forming the =articular=, partly overlain by a series of membrane bones +the _dentary_, _splenial_, _angular_, _supra-angular_ and _coronoid_. +In many sharks large paired accessory cartilages occur at the sides of +the jaws; and in a few reptiles and some Amphibia, such as the Frog, +the ossified representative of the anterior of these structures occurs +forming the =mento-meckelian= bone. In mammals the lower jaw includes +but a single bone. + +The quadrate in all animals with ossified skeletons, except the +Mammalia, forms the suspensorium of the mandible or the skeletal link +between the jaw and the cranium; in the Mammalia, however, the +mandible articulates with the squamosal, while the quadrate is greatly +reduced, and is now generally considered to be represented by the +tympanic ring of the ear. + +The second visceral or hyoid arch in fishes consists of two pieces of +cartilage, a proximal[1] piece the =hyomandibular=, and a distal[10] +piece the =cerato-hyal=. The cerato-hyals of the two sides are commonly +united by a median ventral plate, the =basi-hyal=. The hyoid arch +bears gills on its posterior border, but its most important function +in most fishes is to act as the suspensorium. In higher vertebrates +the representative of the hyomandibular is much reduced in size, and +comes into relation with the ear forming the =auditory ossicles=; the +cerato-hyal looses its attachment to the hyomandibular and becomes +directly attached to the cranium, forming a large part of the hyoid +apparatus of most higher vertebrates. + +Behind the hyoid arch come the branchial arches. They are best +developed in fishes, in which they are commonly five in number and +bear gills. Their ventral ends are united in pairs by median pieces, +the =copulae=. + +In higher vertebrates they become greatly reduced, and all except the +first and second completely disappear. In the highest vertebrates, the +mammals, the second has disappeared, but in birds and many reptiles it +is comparatively well developed. + +3. THE RIBS AND STERNUM. + +The =ribs= are a series of segmentally arranged cartilaginous or bony +rods, attached to the vertebrae; they tend to surround the body +cavity, and to protect the organs contained within it. Ribs are very +frequently found attached to the transverse processes of the +vertebrae, but a study of their origin in fish shows that they are +really the cut off terminations of the ventral arch, not of the +transverse processes which are outgrowths from the dorsal arch. In the +tail their function is to surround and protect structures like the +ventral blood-vessels which do not vary much in size, consequently +they meet one another, and form a series of complete ventral or haemal +arches. But the trunk contains organs like the lungs and stomach which +are liable to vary much in size at different times, consequently the +halves of the haemal arch do not meet ventrally, and then the ribs +become detached from the rest of the haemal arch. Having once become +detached, they are able to shift about and unite themselves to various +points of the vertebra. They frequently, as has been already +mentioned, become entirely attached to the transverse process, or they +may be attached to the transverse process by a dorsal or =tubercular= +portion and to the centrum or to the ventral arch by a ventral or +=capitular= portion. + +In all animals above fishes the distal ends of the thoracic ribs unite +with a median breast bone or sternum which generally has the form of a +segmented rod. The =sternum= is really formed by the fusion of the +distal ends of a series of ribs. In many animals elements of the +shoulder girdle enter into close relation with the rib elements of the +sternum. + +II. THE APPENDICULAR SKELETON. + +This consists of the skeleton of the anterior or =pectoral=, and the +posterior or =pelvic= limbs, and their girdles. In every case (except +in Chelonia) the parts of the appendicular skeleton lie external to +the ribs. + +1. THE LIMB GIRDLES. + +=The Pectoral girdle=[11]. In the simplest case the pectoral or +shoulder girdle consists of a hoop of cartilage incomplete dorsally. +It is attached by muscle to the vertebral column, and is divided on +either side into dorsal and ventral portions by a cavity, the =glenoid +cavity=, at the point where the anterior limb articulates. In higher +fishes this hoop is distinctly divided into right and left halves; it +becomes more or less ossified, and a pair of important bones, the +clavicles, are developed in connection with its ventral portion. + +In higher vertebrates ossification sets up in the cartilage and gives +rise on each side to a dorsal bone, the =scapula=, and frequently to +an anterior ventral bone, the =precoracoid=, and a posterior ventral +bone, the =coracoid=. The precoracoid is often not ossified, and upon +it is developed the clavicle which more or less replaces it. In some +forms a =T= shaped _interclavicle_ occurs, in others =epicoracoids= are +found in front of the coracoids. In all vertebrata above fish, except +the great majority of mammals, the coracoids are large and articulate +with the sternum. But in mammals the coracoids are nearly always quite +vestigial, and the pectoral girdle is attached to the axial skeleton +by the clavicle or sometimes by muscles and ligaments only. + +The =Pelvic girdle=[12] like the pectoral consists primitively of a +simple rod or hoop of cartilage, which in vertebrata above fishes is +divided into dorsal and ventral portions, by a cavity, the +=acetabulum=, with which the posterior limb articulates. In the pelvic +girdle as in the pectoral one dorsal, and (commonly) two ventral +ossifications take place. The dorsal bone is the =ilium= and +corresponds to the scapula. The posterior ventral bone is the +=ischium= corresponding to the coracoid. The anterior ventral bone is +the =pubis= and is generally compared to the precoracoid, but in some +cases a fourth pelvic element, the =acetabular= or =cotyloid= bone is +found, and this may correspond to the precoracoid. + +The pelvic girdle differs from the pectoral in the fact that the +dorsal bones--the ilia--are nearly always firmly united to transverse +processes of the sacral vertebrae, by means of rudimentary ribs. The +pubes and ischia generally meet in ventral symphyses. + +2. THE LIMBS. + +It will be most convenient to defer a discussion of the limbs of +fishes to chap. VIII. + +All vertebrates above fishes have the limbs divisible into three main +segments:-- + + =Anterior or Fore limb.= =Posterior or Hind limb.= + + Proximal segment. upper arm or _brachium_. thigh. + Middle segment. fore-arm or _antibrachium_. shin or _crus_. + Distal segment. _manus_. _pes_. + +The proximal segments each contain one bone, the =humerus= in the case +of the upper arm, and the =femur= in the case of the thigh. The middle +segments each contain two bones, the =radius= and =ulna= in the case +of the fore-arm, and the =tibia= and =fibula= in the case of the shin. + + * * * * * + +The manus and pes are further subdivided into + +(_a_) two or three proximal rows of bones forming the wrist or +=carpus= in the case of the manus, and the ankle or =tarsus= in the +case of the pes. + +(_b_) a middle row called respectively the =metacarpus= and +=metatarsus=. + +(_c_) a number of distal bones called the =phalanges= which form the +skeleton of the fingers and toes, or =digits=. + +Typically the manus and pes both have five digits (pentedactylate). +The first digit of the manus is commonly called the =pollex=, and the +first digit of the pes the =hallux=. + +In a very simple =carpus= such as that of _Chelydra_, there are nine +bones. They are arranged in a proximal row of three, the radiale, +intermedium, and ulnare,--the first being on the radial side of the +limb, and a distal row of five called respectively carpale 1, 2, 3, 4, +5, beginning on the radial side. Between these two rows is a single +bone the centrale, or there may be two. + +Similarly there are nine bones in a simple =tarsus= such as that of +_Salamandra_. They form a proximal row of three, the tibiale, +intermedium and fibulare, and a distal row of five, called +respectively tarsale 1, 2, 3, 4, 5, beginning on the tibial side. +Between the two rows there is a centrale as in the carpus, or there +may be two. + + * * * * * + +The following names derived from human anatomy are commonly applied to +the various carpal and tarsal bones: + + =Carpus.= + + radiale = scaphoid + intermedium = lunar + ulnare = cuneiform + centrale = central + carpale 1 = trapezium + " 2 = trapezoid + " 3 = magnum + " 4 } = unciform + " 5 } + + =Tarsus.= + + tibiale } + intermedium } astragalus + fibulare = calcaneum + centrale = navicular + tarsale 1 = internal cuneiform + " 2 = middle " + " 3 = external " + " 4 } = cuboid + " 5 } + + NOTE. The above is the view commonly accepted concerning the + homology of the carpal and tarsal bones. But with regard to the + proximal row of tarsal bones there is difference of opinion. + All anatomists are agreed that the calcaneum is the fibulare + and that the intermedium is contained in the astragalus, but + while the majority regard the astragalus as the fused tibiale + and intermedium, Baur considers that a small bone found on the + tibial side of the tarsus in _Procavia_, many Rodents, + Insectivores, and the male _Ornithorhynchus_, is the vestigial + tibiale, and regards the astragalus as the intermedium + alone[13]. He also considers that the mammalian scaphoid + represents a centrale. + + +MODIFICATIONS IN THE POSITIONS OF THE LIMBS[14]. + +In their primitive position the limbs are straight and are extended +parallel to one another at right angles to the axis of the trunk. Each +limb then has a dorsal surface, a ventral surface, an anterior or +=pre-axial= edge, and a posterior or =postaxial= edge. + +In the anterior limb the radius and the pollex are pre-axial, the ulna +and the fifth finger are postaxial. In the posterior limb the tibia +and the hallux are pre-axial, the fibula and the fifth toe are +postaxial. The Cetacea and various extinct reptiles, such as +_Ichthyosaurus_ and _Plesiosaurus_, have their limbs in practically +this primitive position. + +The first modification from it is produced by the bending ventrally of +the middle segments of both limbs upon the proximal segments, while +the distal segment is bent in the opposite direction on the middle +segment. Then the ventral surfaces of the antibrachium and crus come +to look inwards, and their dorsal surfaces to look outwards. The +brachium and manus, thigh and pes still have their dorsal surfaces +facing upwards and their ventral surfaces facing downwards as before, +and the relations of their pre-and postaxial borders remain as they +were. Many Amphibians and Reptiles, such as tortoises, carry their +limbs in this position. + +In all higher vertebrates, however, a further change takes place, each +limb is rotated as a whole from its proximal end, the rotation taking +place in opposite directions in the fore and hind limbs respectively. +The anterior limb is rotated backwards from the shoulder, so that the +brachium lies nearly parallel to the body, and the elbow points +backwards, the antibrachium downwards, and the manus backwards; the +pre-axial surface of the whole limb with the radius and pollex now +faces outwards, and the postaxial surface with the ulna and fifth +finger now faces inwards. In the Walrus and, to a certain extent, in +the Sea lions the anterior limb remains throughout life in this +position. The posterior limb is also rotated, but the rotation in this +case takes place forwards, so that the thigh lies nearly parallel to +the body, the knee-joint pointing forwards; the crus downwards and the +pes forwards. The pre-axial surface of the whole limb with the tibia +and hallux looks towards the middle of the body, the postaxial surface +with the fibula and fifth toe looks outwards. This is the position in +which the hind limb is carried in nearly all mammals. + +In nearly all mammals a further change takes place in the position of +the anterior limb. The radius and ulna have hitherto been parallel to +one another, but now the lower end of the radius, carrying with it the +manus, comes to be rotated forwards round the ulna, so that the manus, +as well as the pes, comes to be forwardly-directed, and its pre-axial +surface faces inwards. + +In the majority of mammals the radius and ulna are permanently fixed +in this, which is known as the =prone= position, but in man and some +other mammals the manus can be pronated or turned into this position +at will. When the radius and ulna are parallel throughout their whole +length the manus is said to be in the =supine= position. + +The =extensor= side of a limb is that to which the muscles which +straighten it are attached, the =flexor= side is that to which the +muscles which bend it are attached. + + +FOOTNOTES: + +[1] The skin consists of an outer layer of epiblastic origin, the +epidermis, and an inner layer of mesoblastic origin, the dermis. The +epidermis is divided into two principal layers, an outer one, the +horny layer or _stratum corneum_, and an inner one, the _stratum +Malpighii_. The innermost part of the stratum corneum is distinguished +as the _stratum lucidum_, and the outermost part of the stratum +Malpighii as the _stratum granulosum_. + +[2] The enamel of the pharyngeal teeth of some Teleosteans is +hypoblastic in origin. + +[3] See also p. 71. + +[4] It is usual to regard the clavicle as a membrane bone, but +Kölliker has shown that in rabbit embryos of about the 17th day it is +cartilaginous. + +[5] In compiling these paragraphs on Histology, free use has been made +of Klein and Noble Smith's _Atlas of Histology_, the small Histologies +of Klein and Schäfer, Huxley's _Elementary Physiology_, and Lloyd +Morgan's _Animal Biology_. + +[6] See Huxley's _Elementary Physiology_, Revised edition, London, +1886, p. 180. + +[7] Strictly speaking the jaws, visceral skeleton, ribs and sternum do +not form part of the axis, but it is convenient to group them as parts +of the axial skeleton. + +[8] F.M. Balfour, _Comparative Embryology_, vol. II., London, 1881, p. +465. W.K. Parker and G.T. Bettany, _The Morphology of the Skull_, +London, 1877. + +[9] Sometimes also called ectethmoids or parethmoids. + +[10] The _proximal_ end of anything is the one nearest the point of +origin or attachment, the _distal_ end is the one furthest from the +point of origin or attachment. + +[11] W.K. Parker, _A Monograph of the Shoulder Girdle and Sternum_, +Ray Soc. London, 1868. + +[12] See R. Wiedersheim, _Zeitschr. wiss. Zool._ vol. LIII. suppl. p. +43, 1892. + +[13] G. Baur, _Beiträge zur Morphogenie des Carpus und Tarsus der +Vertebraten_, Theil 1. Batrachia. Jena, 1888, and _Amer. Natural._, +vol. XIX. 1885 (several papers). + +[14] This account is based on Chapter XX. of Flower's _Osteology of +the Mammalia_. London 1876. + + + + +CHAPTER II. + +CLASSIFICATION. + + +THE following classification includes _only the forms mentioned in the +succeeding pages_. The relative value of some of the terms employed in +classification is not identical throughout the book. This remark +applies specially to the term _group_, which is a convenient one, +owing to its not having such a hard and fast zoological meaning as has +the term _family_, for instance. The term _group_ is applied in this +book to divisions of the animal kingdom of very different +classificatory importance. + + PHYLUM CHORDATA. + + SUBPHYLUM A. HEMICHORDATA. + + Balanoglossus. + Cephalodiscus. + Rhabdopleura. + ? Phoronis. + (? Actinotrocha--larval Phoronis). + + SUBPHYLUM B. UROCHORDATA (TUNICATA). + + Group LARVACEA and others. + + SUBPHYLUM C. CEPHALOCHORDATA. + + Amphioxus--lancelet. + + NOTE. In this chapter all the generic names printed in italics + are those of extinct animals. + + SUBPHYLUM D. VERTEBRATA. + + DIVISION (I). CYCLOSTOMATA. + + Order 1. MARSIPOBRANCHII. + + Family =Myxinoidei=. Myxine--hag-fish. + Bdellostoma. + + Family =Petromyzontidae=. Petromyzon--lamprey. + (Ammocoetes--larval lamprey.) + + Family =Palaeospondylidae=. _Palaeospondylus._ + + Order 2. OSTRACODERMI. + + Suborder 1. HETEROSTRACI. + + Family =Pteraspidae=. _Pteraspis._ + + Suborder 2. OSTEOSTRACI. + + Family =Cephalaspidae=. _Cephalaspis._ + + Suborder 3. ANTIARCHA. + + Family =Asterolepidae=. _Pterichthys._ + _Asterolepis._ + + + DIVISION (II). GNATHOSTOMATA. + + A. ICHTHYOPSIDA. + + CLASS I. PISCES. + + Order 1. ELASMOBRANCHII. + + Suborder (1). ICHTHYOTOMI. + + Family =Pleuracanthidae=. _Xenacanthus._ + + Suborder (2). PLEUROPTERYGII. + _Cladoselache._ + + Suborder (3). SELACHII. + + Group SQUALIDAE. + + Family =Notidanidae=. Heptanchus. + Hexanchus. + Chlamydoselache--frill-gilled shark. + + Family =Cochliodontidae=. _Cochliodus._ + + Family =Cestraciontidae=. Cestracion--Port Jackson shark. + _Acrodus._ + + Family =Scylliidae=. Scyllium--spotted dogfish. + + Family =Lamnidae=. Odontaspis. + + Family =Carcharidae=. Galeus--tope. + + Family =Spinacidae=. Acanthias--spiny dogfish. + Scymnus. + + Family =Squatinidae=. Squatina (Rhina)--angel fish. + + Group BATOIDEI. + + Family =Pristidae=. Pristis--saw-fish. + + Family =Raiidae=. Raia--skate. + + Family =Myliobatidae=. Myliobatis--eagle ray. + + Family =Trygonidae=. Trygon--sting ray. + + Family =Torpedinidae=. Torpedo--electric ray. + + Suborder (4). ACANTHODII. + + Family =Acanthodidae=. _Acanthodes._ + + Family =Diplacanthidae=. _Diplacanthus._ + + Order 2. HOLOCEPHALI. + + Family =Chimaeridae=. Chimaera--rabbit fish. + Harriotta. + Callorhynchus. + _Ischyodus._ + + Order 3. GANOIDEI. + + Suborder (1). CHONDROSTEI. + + Family =Palaeoniscidae=. _Palaeoniscus._ + _Trissolepis._ + + Family =Acipenseridae=. Acipenser--sturgeon. + Scaphirhynchus. + + Family =Polyodontidae=. Polyodon (Spatularia)--spoon-beaked sturgeon. + Psephurus--slender-beaked sturgeon. + + Suborder (2). CROSSOPTERYGII. + + Family =Holoptychiidae=. _Holoptychius._ + + Family =Rhizodontidae=. _Rhizodus._ + + Family =Osteolepidae=. _Osteolepis._ + + Family =Polypteridae=. Polypterus--bichir. + Calamoichthys--reed-fish. + + Suborder (3). HOLOSTEI. + + Family =Lepidosteidae=. Lepidosteus--gar pike. + + Family =Semionotidae=. _Lepidotus._ + + Family =Amiidae=. Amia--bow-fin. + + Order 4. TELEOSTEI. + + Suborder (1). PLECTOGNATHI. + + Family =Balistidae=. Balistes--file-fish. + + Family =Gymnodontidae=. Diodon--globe-fish. + + Family =Ostracionidae=. Ostracion--coffer-fish. + + Suborder (2). PHYSOSTOMI. + + Family =Siluridae=.--cat-fishes. + + Family =Cyprinidae=. Cyprinus--carp. + + Family =Esocidae=. Esox--pike. + + Family =Salmonidae=. Salmo--salmon. + + Family =Clupeidae=. Clupeus--herring. + Exocaetus--'flying fish'. + + Family =Muraenidae=. Anguilla--eel. + + Suborder (3). ANACANTHINI. + + Family =Gadidae=. Gadus--cod, haddock, whiting. + + Family =Pleuronectidae=. Solea--sole. + + Suborder (4). PHARYNGOGNATHI. + + Family =Labridae=. Labrus--wrasse. + Scarus--parrot fish. + + Suborder (5). ACANTHOPTERYGII. + + Family =Cataphracti=. Dactylopterus--flying gurnard. + + Family =Percidae=. Perca--perch. + + Order 5. DIPNOI. + + Suborder (1). SIRENOIDEI. + + Family =Dipteridae=. _Dipterus._ + + Family =Monopneumona=. Ceratodus--barramunda. + + Family =Dipneumona=. Protopterus--African mud-fish. + Lepidosiren. + + Suborder (2). ARTHRODIRA. + + Family =Coccosteidae=. _Coccosteus._ + _Dinichthys._ + + NOTE. Palaeontological research has disclosed the existence of + a great number of forms which seem to connect with one another + almost all the orders of fishes as usually recognised. Forms + connecting the living Ganoids with the Teleosteans have been + especially numerous, so that these terms Ganoid and Teleostean + can hardly be any longer used in a precise and scientific + sense. This has rendered the subject of the classification of + fishes a very difficult one. Though unsuitable for adoption in + a work like the present, by far the most natural classification + hitherto proposed seems to be that of Smith Woodward[15]. He + considers that the course of development of fishes has followed + two distinct lines, the autostylic and hyostylic (see p. 119), + and groups the various forms as follows: + + HYOSTYLIC. AUTOSTYLIC. + Subclass 1. ELASMOBRANCHII. Subclass 3. HOLOCEPHALI. + 1. Ichthyotomi. 1. (unknown). + 2. Selachii. 2. Chimaeroidei. + 3. Acanthodii. 3. (unknown). + + Subclass 2. TELEOSTOMI. Subclass 4. DIPNOI. + 1. Crossopterygii (Palaeozoic 1. Sirenoidei. + and Mesozoic). + 2. Crossopterygii (Cainozoic). 2. (unknown). + 3. Actinopterygii. 3. Arthrodira. + + The primitive forms in each of these four subclasses have the + fins archipterygia (see p. 127). + + CLASS II. AMPHIBIA. + + Order 1. URODELA. + + Suborder (1). ICHTHYOIDEA. + + Group A. PERENNIBRANCHIATA. + + Family =Menobranchidae=. Menobranchus. + + Family =Proteidae=. Proteus--olm. + + Family =Sirenidae=. Siren. + + Group B. DEROTREMATA. + + Family =Amphiumidae=. Megalobatrachus. + Cryptobranchus (Menopoma). + Amphiuma. + + Suborder (2). SALAMANDRINA. + + Family =Salamandridae=. Salamandra--salamander. + Molge--newt. + Onychodactylus. + Amblystoma. + (Siredon--axolotl, larval Amblystoma). + Batrachoseps. + Spelerpes (Gyrinophilus). + + Order 2. LABYRINTHODONTIA. + + Group =Lepospondyli=. _Branchiosaurus._ + + Group =Temnospondyli=. _Archegosaurus._ + _Nyrania._ + _Euchirosaurus._ + + Group =Stereospondyli=. _Capitosaurus._ + _Mastodonsaurus._ + + Order 3. GYMNOPHIONA. + + Family =Caeciliidae=. Siphonops. + Epicrium. + + Order 4. ANURA. + + Suborder (1). AGLOSSA. + + Family =Xenopidae=. Xenopus. + + Family =Pipidae=. Pipa--Surinam toad. + + Suborder (2). PHANEROGLOSSA. + + Group ARCIFERA. + + Family =Discoglossidae=. Discoglossus--painted frog. + Bombinator--fire-bellied frog. + Alytes--midwife frog. + + Family =Pelobatidae=. Pelobates--toad frog. + + Family =Hylidae=. Hyla--green tree-frog. + + Family =Bufonidae=. Bufo--toad. + Docidophryne. + + Family =Cystignathidae=. Ceratophrys--horned frog. + + Group FIRMISTERNIA. + + Family =Ranidae=. Rana--common and edible frogs. + + Family =Engystomatidae=. Brachycephalus. + + B. SAUROPSIDA. + + CLASS I. REPTILIA[16]. + + Order 1. THEROMORPHA. + + Group =Anomodontia=. _Dicynodon._ + _Udenodon._ + + Group =Placodontia=. _Placodus._ + + Group =Pariasauria=. _Pariasaurus._ + _Elginia._ + + Group =Theriodontia=. _Dimetrodon._ + _Galesaurus._ + _Cynognathus._ + + Order 2. SAUROPTERYGIA. + + Family =Mesosauridae=. _Mesosaurus._ + + Family =Nothosauridae=. _Nothosaurus._ + + Family =Plesiosauridae=. _Plesiosaurus._ + _Pliosaurus._ + + Order 3. CHELONIA. + + Suborder (1). TRIONYCHIA. + + Family =Trionychidae=. Trionyx--snapping turtle. + + Suborder (2). CRYPTODIRA. + + Family =Dermochelydidae=. Dermochelys (Sphargis)--leathery + turtle. + + Family =Chelonidae=. Chelone--green turtle. + + Family =Chelydridae=. Chelydra--terrapin. + + Family =Chersidae=. Testudo--tortoise. + + Suborder (3). PLEURODIRA. + + Family =Chelydae=. Chelys. + + Order 4. ICHTHYOSAURIA. + + Family =Ichthyosauridae=. _Ichthyosaurus._ + + Order 5. RHYNCHOCEPHALIA. + + Suborder (1). RHYNCHOCEPHALIA VERA. + + Family =Sphenodontidae=. Sphenodon (Hatteria). + + Family =Rhynchosauridae=. _Hyperodapedon._ + + Suborder (2). PROGANOSAURIA. + + Family =Proterosauridae=. _Proterosaurus._ + + Order 6. SQUAMATA. + + Suborder (1). LACERTILIA. + + Group =Lacertilia vera=. + + Family =Geckonidae=. Gecko. + + Family =Pygopodidae=. Lialis--scale-foot. + + Family =Agamidae=. Draco--flying lizard. + Agama. + + Family =Iguanidae=. Iguana. + + Family =Anguidae=. Ophisaurus (Bipes, Pseudopus). + Anguis--blindworm. + + Family =Varanidae=. Varanus--monitor. + + Family =Amphisbaenidae=. Chirotes. + Amphisbaena. + + Family =Scincidae=. Tiliqua (Cyclodus). + Scincus--skink. + Chalcides (Seps). + + + Group =Rhiptoglossa=. + + Family =Chamaeleonidae=. Chamaeleon. + + Suborder (2). OPHIDIA. + + Family =Typhlopidae=. Typhlops--blind snake. + + Family =Boidae=. Python. + + Family =Colubridae=. Tropidonotus--ringed snake. + + Family =Hydrophidae=--sea snakes. + + Family =Crotalidae=. Crotalus--rattlesnake. + + Suborder (3). PYTHONOMORPHA. + + Family =Mosasauridae=. _Mosasaurus._ + + Order 7. DINOSAURIA. + + Suborder (1). SAUROPODA. + + Family =Atlantosauridae=. _Brontosaurus._ + + Family =Cetiosauridae=. _Morosaurus._ + + Suborder (2). THEROPODA. + + Family =Megalosauridae=. _Megalosaurus_ (_Ceratosaurus_). + + Family =Compsognathidae=. _Compsognathus._ + + Suborder (3). ORTHOPODA. + + Section (_a_). STEGOSAURIA. + + Family =Scelidosauridae=. _Polacanthus._ + + Family =Stegosauridae=. _Stegosaurus._ + + Section (_b_). CERATOPSIA. + + Family =Ceratopsidae=. _Polyonax_ (_Ceratops_). + + Section (_c_). ORNITHOPODA. + + Family =Camptosauridae=. _Hypsilophodon._ + + Family =Iguanodontidae=. _Iguanodon._ + + Family =Hadrosauridae=. _Hadrosaurus._ + + Order 8. CROCODILIA. + + Suborder (1). PARASUCHIA. + + Family =Phytosauridae=. _Phytosaurus_ (_Belodon_). + + Suborder (2). EUSUCHIA. + + Family =Teleosauridae=. _Teleosaurus._ + _Metriorhynchus._ + + Family =Goniopholidae=. _Goniopholis._ + + Family =Alligatoridae=. Alligator. + Caiman. + Jacare. + + Family =Crocodilidae=. Crocodilus. + + Family =Garialidae=. Garialis (Gavialis). + + Order 9. PTEROSAURIA. + + Family =Pterodactylidae=. _Pterodactylus._ + + Family =Rhamphorhynchidae=. _Rhamphorhynchus._ + + Family =Pteranodontidae=. _Pteranodon._ + + +FOOTNOTES: + +[15] A. Smith Woodward, _Catalogue of Fossil Fishes in the British +Museum_, Part II., Introduction, p. xii. + +[16] This classification of reptiles is mainly based on that of +Lydekker (_Catalogue of Fossil Reptiles in the British Museum_) but in +some respects that of von Zittel has been followed. + + + + + CLASS II. AVES[17]. + + + Subclass (I). ARCHAEORNITHES. + _Archaeopteryx._ + + Subclass (II). NEORNITHES. + + Order 1. RATITAE. + + Group =Æpyornithes=. _Æpyornis._ + + Group =Apteryges=. Apteryx--kiwi. + + Group =Dinornithes=. Moas. + + Group =Megistanes=. Casuarius--cassowary. + Dromaeus--emeu. + + Group =Rheornithes=. Rhea--American ostrich. + + Group =Struthiornithes=. Struthio--ostrich. + + Order 2. ODONTOLCAE. + _Hesperornis._ + + Order 3. CARINATAE. + + Group =Ichthyornithiformes=. + _Ichthyornis._ + _Apatornis._ + _Odontopteryx._ + + Group =Colymbiformes=. + Subgroup Colymbi--divers. + + Group =Sphenisciformes=. + Subgroup Sphenisci--penguins. + + + Group =Ciconiiformes=. + + Subgroup Steganopodes. Sula--gannet. + Pelicanus--pelican. + Phaëthon--frigate bird. + Phalacrocorax--cormorant. + + Subgroup Ardeae. Ardea--heron + + Subgroup Ciconiae. Leptoptilus--adjutant. + Ciconia--white stork. + + Group =Anseriformes=. + + Subgroup Palamedeae. Palamedea } + } screamers. + Chauna } + + Subgroup Anseres. Anas--wild duck. + Anser--goose. + Plectropterus--spur-winged goose. + Cygnus--swan. + Mergus--merganser. + + Group =Falconiformes=. + + Subgroup Cathartae. Cathartes--American vulture. + + Subgroup Accipitres. Falco--falcon. + Vultur--vulture. + Harpagus. + Gypogeranus--secretary bird. + + Group =Tinamiformes=. + + Subgroup Tinami. Tinamus. + + Group =Galliformes=. + + Subgroup Galli. Gallus--fowl. + Pavo--peacock. + + Subgroup Opisthocomi. Opisthocomus--hoatzin. + + Group =Gruiformes=. + Gruidae--cranes. + + Group =Stereornithes=. _Phororhacos._ + + Group =Charadriiformes=. + + Subgroup Limicolae. Charadriidae--plovers. + Parra--jacana. + + Subgroup Lari. Laridae--gulls. + Alcidae--auks. + + Subgroup Pteroclidae. Pterocles--sandgrouse. + + Subgroup Columbidae. Columbae--pigeons. + _Didus_--dodo. + _Pezophaps_--solitaire. + + Group =Cuculiformes=. + + Subgroup Cuculi. Scythrops. + + Subgroup Psittaci. Stringops--owl-parrot. + + Group =Coraciiformes=. + + Subgroup Coraciae. Coracias--roller. + Buceros--hornbill. + Upupa--hoopoe. + + Subgroup Striges. Owls. + + Subgroup Cypseli. Cypselidae--swifts. + Trochilidae--humming-birds. + + Subgroup Trogonidae. Trogons. + + Subgroup Pici. Rhamphastos--toucan. + Picus--woodpecker. + + Group =Passeriformes=. Crows, finches, larks, warblers, + and many others. + +C. MAMMALIA[18]. + +Class MAMMALIA. + +Subclass (I). ORNITHODELPHIA or PROTOTHERIA. + +Order. MONOTREMATA. + + Family =Ornithorhynchidae=. Ornithorhynchus--duck-bill. + + Family =Echidnidae=. Echidna--spiny ant-eater. + + Group =Multituberculata=. _Tritylodon._ + + Subclass (II). DIDELPHIA or METATHERIA. + + Order. MARSUPIALIA. + + Suborder (1). POLYPROTODONTIA. + + Family =Amphitheriidae=. _Phascolotherium._ + + Family =Didelphyidae=. Didelphys--opossum. + + Family =Dasyuridae=. Thylacinus--Tasmanian wolf. + Sarcophilus--Tasmanian devil. + Dasyurus. + + Family =Peramelidae=. Perameles--bandicoot. + Choeropus. + + Family =Notoryctidae=. Notoryctes--marsupial mole. + + Suborder (2). DIPROTODONTIA. + + Family =Phascolomyidae=. Phascolomys--wombat. + + Family Phalangeridae. Tarsipes. + Phalanger--cuscus. + Phascolarctus--koala. + _Thylacoleo._ + + Family =Diprotodontidae=. _Diprotodon._ + + Family =Nototheriidae=. _Nototherium._ + + Family =Macropodidae=. Macropus--kangaroo. + + Family =Epanorthidae=. Coenolestes. + + Subclass (III). MONODELPHIA or EUTHERIA. + + Order 1. EDENTATA. + + Family =Bradypodidae=. Bradypus } + }--sloths. + Choloepus } + + Family =Megatheriidae=. _Megatherium_--ground sloth. + + Family =Myrmecophagidae=. Myrmecophaga--great ant-eater. + Cycloturus--two-toed ant-eater. + + Family =Dasypodidae=. Chlamydophorus } + Dasypus }--armadillos. + Priodon } + Tatusia } + + Family =Glyptodontidae=. _Glyptodon._ + + Family =Manidae=. Manis--pangolin. + + Family =Orycteropodidae=. Orycteropus--aard-vark. + + Order 2. SIRENIA. + + Family =Manatidae=. Manatus--manatee. + + Family =Rhytinidae=. _Rhytina_--Steller's sea-cow. + + Family =Halicoridae=. Halicore--dugong. + + Family =Halitheriidae=. _Halitherium._ + + Order 3. CETACEA. + + Suborder (1). ARCHAEOCETI. + + Family =Zeuglodontidae=. _Zeuglodon._ + + Suborder (2). MYSTACOCETI or BALAENOIDEA. + + Family =Balaenidae=. Balaena--right whale. + Megaptera--humpbacked whale. + Balaenoptera--rorqual. + + Suborder (3). ODONTOCETI. + + Family =Physeteridae=. Physeter--sperm whale. + Hyperoödon--bottlenose. + Ziphius. + Mesoplodon. + + Family =Physodontidae=. _Physodon._ + + Family =Squalodontidae=. _Squalodon._ + + Family =Platanistidae=. Platanista--Gangetic dolphin. + Inia. + Pontoporia. + + Family =Delphinidae=. Monodon--narwhal. + Phocaena--porpoise. + Orca--killer. + Globicephalus--Ca'ing whale. + Grampus. + Lagenorhynchus. + Delphinus--dolphin. + Tursiops. + Prodelphinus. + + Order 4. UNGULATA. + + Division A. UNGULATA VERA. + + Suborder (1). ARTIODACTYLA. + + Section (_a_). SUINA. + + Family =Hippopotamidae=. Hippopotamus. + + Family =Suidae=. Sus--pig. + Babirussa. + Phacochaerus--wart hog. + _Hyotherium._ + + Family =Cotylopidae=. _Cotylops (Oreodon)._ + _Cyclopidius._ + + Family =Agriochoeridae=. _Agriochoerus._ + + Family =Anoplotheriidae=. _Anoplotherium._ + + Section (_b_). TYLOPODA. + + Family =Camelidae=. Camelus--camel. + Auchenia--llama. + + Section (_c_). TRAGULINA. + + Family =Tragulidae=. Dorcatherium (Hyomoschus)--chevrotain. + + Section (_d_). RUMINANTIA or PECORA. + + Family =Cervidae=. Moschus--musk deer. + Cervus--deer. + Cervulus--muntjac. + Hydropotes--Chinese water deer. + + Family =Giraffidae=. Giraffa--giraffe. + _Sivatherium._ + + Family =Antilocapridae=. Antilocapra--prongbuck. + + Family =Bovidae=. Tetraceros--four-horned antelope. + Gazella--gazelle. + Bos--ox. + Bison. + Bubalus--buffalo. + + Suborder (2). PERISSODACTYLA. + + Family =Tapiridae=. Tapirus--tapir. + + Family =Lophiodontidae=. _Lophiodon._ + _Hyracotherium._ + + Family =Palaeotheriidae=. _Palaeotherium._ + + Family =Equidae=. _Hipparion._ + Equus--horse. + + Family =Rhinocerotidae=. Rhinoceros. + _Elasmotherium._ + + Family =Titanotheriidae=. _Titanotherium (Brontops)._ + _Palaeosyops._ + + Family =Chalicotheriidae=. _Chalicotherium._ + + Family =Macraucheniidae=. _Macrauchenia._ + + Division B. SUBUNGULATA. + + Suborder (1). TOXODONTIA. + + Family =Astrapotheriidae=. _Astrapotherium._ + + Family =Nesodontidae=. _Nesodon._ + + Family =Toxodontidae=. _Toxodon._ + + Family =Typotheriidae=. _Typotherium._ + + + Suborder (2). CONDYLARTHRA. + + Family =Phenacodontidae=. _Phenacodus._ + + Suborder (3). HYRACOIDEA. + + Family =Hyracidae=. Procavia (Hyrax). + + Suborder (4). AMBLYPODA. + + Family =Coryphodontidae=. _Coryphodon._ + + Family =Uintatheriidae=. _Uintatherium_ (_Dinoceras_). + + Suborder (5). PROBOSCIDEA. + + Family =Dinotheriidae=. _Dinotherium._ + + Family =Elephantidae=. _Mastodon._ + Elephas--elephant. + + Group Tillodontia. + + Order 5. RODENTIA. + + Suborder (1). SIMPLICIDENTATA. + + Section SCIUROMORPHA. + + Family =Castoridae=. Castor--beaver. + + Section MYOMORPHA. + + Family =Lophiomyidae=. Lophiomys. + + Family =Muridae=. Hydromys. + Acanthomys--spiny mouse. + Mus--mouse. + + Family =Spalacidae=. Bathyergus. + + Family =Dipodidae=. Dipus--jerboa. + Pedetes--Cape jumping-hare. + + Section HYSTRICOMORPHA. + + Family =Hystricidae=. Hystrix--porcupine. + + Family =Chinchillidae=. Chinchilla. + Lagostomus--viscacha. + + Family =Dasyproctidae=. Coelogenys--paca. + Dasyprocta--agouti. + + Family =Caviidae=. Cavia--guinea-pig. + Hydrochaerus--capybara. + + Suborder (2). DUPLICIDENTATA. + + Family =Leporidae=. Lepus--hare and rabbit. + + Order 6. CARNIVORA. + + Suborder (1). CREODONTA. + + Family =Hyaenodontidae=. _Hyaenodon._ + + Suborder (2). CARNIVORA VERA or FISSIPEDIA. + + Section ÆLUROIDEA. + + Family =Felidae=. Felis--cat, lion, tiger. + _Machaerodus_--sabre-toothed lion. + + Family =Viverridae=. Viverra--civet. + Paradoxurus--palm civet. + + Family =Protelidae=. Proteles--aard wolf. + + Family =Hyaenidae=. Hyaena. + + Section CYNOIDEA. + + Family =Canidae=. Canis--dog, wolf, fox. + + Section ARCTOIDEA. + + Family =Ursidae=. Ursus--bear. + + Family =Mustelidae=. Latax--sea otter. + + Suborder (3). PINNIPEDIA. + + Family =Otariidae=. Otaria--sea lion. + + Family =Trichechidae=. Trichechus--walrus. + + Family =Phocidae=. Ogmorhinus--sea leopard. + + Order 7. INSECTIVORA. + + Suborder (1). DERMOPTERA. + + Family =Galeopithecidae=. Galeopithecus--'flying lemur'. + + Suborder (2). INSECTIVORA VERA. + + Family =Macroscelidae=. Macroscelides--jumping shrew. + + Family =Erinaceidae=. Erinaceus--hedgehog. + Gymnura. + + Family =Soricidae=. Sorex--shrew. + + Family =Talpidae=. Talpa--mole. + + Family =Potamogalidae=. Potamogale. + + Family =Solenodontidae=. Solenodon. + + Family =Centetidae=. Microgale. + Centetes--tenrec. + + Family =Chrysochloridae=. Chrysochloris--golden mole. + + Order 8. CHIROPTERA. + + Suborder (1). MEGACHIROPTERA. + + Family =Pteropidae=. Pteropus--flying fox. + + Suborder (2). MICROCHIROPTERA. + + Family =Rhinolophidae=. Horseshoe bats. + + Family =Phyllostomatidae=. Desmodus--vampire. + + Order 9. PRIMATES. + + Suborder (1). LEMUROIDEA. + + Family =Tarsiidae=. Tarsius--tarsier. + + Family =Chiromyidae=. Chiromys--aye aye. + + Suborder (2). ANTHROPOIDEA. + + Family =Hapalidae=. Hapale--marmoset. + + Family =Cebidae=. Mycetes--howling monkey. + Ateles--spider monkey. + + Family =Cercopithecidae=. Cynocephalus--baboon. + Macacus. + Colobus. + + Family =Simiidae=. Hylobates--gibbon. + Simia--orang. + Gorilla. + Anthropopithecus--chimpanzee. + + Family =Hominidae=. Homo--man. + + +FOOTNOTES: + +[17] This classification of birds is essentially that of Gadow and +Selenka in Bronn's _Classen und Ordnungen des Thierreichs_, Band VI., +Abth. IV., Vögel. Leipzig, 1891. + +[18] The classification adopted is almost entirely that given in +Flower and Lydekker's _Mammals Living and Extinct_. London, 1891. + + + + +CHAPTER III. + +SKELETON OF HEMICHORDATA, UROCHORDATA, AND CEPHALOCHORDATA. + + +SUBPHYLUM A. HEMICHORDATA. + +THE subphylum includes three genera, _Balanoglossus_[19], +_Cephalodiscus_ and _Rhabdopleura_; and perhaps a fourth, _Phoronis_. + +The skeletal structures found in _Balanoglossus_[20] are all +endoskeletal. They include: + +(1) The =notochord=. This arises as a diverticulum from the alimentary +canal which grows forwards into the proboscis and extends beyond the +front end of the central nervous system. It is hypoblastic in origin +and arises in the same way as does the notochord of _Amphioxus_. Its +cells become highly vacuolated and take on the typical notochordal +structure[21]. The cavity of the primitive diverticulum becomes +obliterated in front, but behind it opens throughout life into the +alimentary canal. + +(2) The =axial skeletal rods=. These are a pair of chitinous rods +which lie ventral to the notochord and in the collar region unite to +form a single mass. + +(3) The =branchial skeleton=. The gill bars separating the gill slits +from one another are strengthened by chitinous rods in a way closely +similar to that in _Amphioxus_. But between one primary forked rod and +the next there are two secondary unforked rods--not one, as in +_Amphioxus_. + +(4) The =chondroid tissue=. This is of mesoblastic origin and may be +regarded as an imperfect sheath for the notochord. + +In _Cephalodiscus_ and _Rhabdopleura_ as in _Balanoglossus_ the +notochord forms a small diverticulum growing forwards from the +alimentary canal into the proboscis stalk. + +Recent researches on _Phoronis_[22] show the existence in the collar +region of the larva (_Actinotrocha_) of a paired organ, which is +regarded by its discoverer as representing a double notochord. + + +SUBPHYLUM B. UROCHORDATA (TUNICATA). + +Skeletal structures of epiblastic and hypoblastic origin occur in the +Urochordata. Most Tunicates are invested by a thick gelatinous test +which often contains calcareous spicules, and serves as a supporting +organ for the soft body. The cells of this test are mesodermal in +origin. + +In larval Tunicata and in adults of the group Larvacea the tail is +supported by a typical notochord, which is confined to the tail. In +all Tunicata except Larvacea all trace of the notochord is lost in the +adult. + + +SUBPHYLUM C. CEPHALOCHORDATA. + +[Illustration FIG. 3. DIAGRAM OF THE SKELETON OF _Amphioxus +lanceolatus_ × 3 (after a drawing in the Index collection at the Brit. +Mus.). + + 1. skeleton of dorsal fin. + 2. notochord. + 3. neural tube. + 4. buccal skeleton. + 5. branchial skeleton. + 6. septa separating the myotomes. + 7. skeleton of ventral fin.] + +This subphylum includes the well-known genus _Amphioxus_[23]. In +_Amphioxus_ the skeleton is very simple. It contains no trace of +cartilage or bone and remains throughout life in a condition +corresponding to a very early stage in Vertebrata. The skeleton of +_Amphioxus_ is partly hypoblastic, partly mesoblastic in origin. + +(_a_) =Hypoblastic skeleton.= + +The =notochord= (fig. 3, 2) is an elastic rod extending along the +whole length of the body past the anterior end of the nerve cord. It +lies ventral to the nerve cord, and shows no trace of segmentation. It +is chiefly made up of greatly vacuolated cells containing lymph, but +near the dorsal and ventral surfaces the cells are less vacuolated. +The notochord is immediately surrounded by a structureless cuticular +layer, the _chordal sheath_, and outside this comes the mesoblastic +_skeletogenous layer_, which also surrounds the nerve cord. + +The =branchial skeleton=. This consists of a series of chitinous +elastic rods which strengthen the gill bars and are alternately forked +and unforked ventrally. The forked rods are primary, and are U-shaped +in section, the unforked rods are secondary, and are circular in +section. All these rods are united at intervals by transverse rods. + +(_b_) =Mesoblastic skeleton.= + +The =buccal skeleton=. On each side of the mouth there is a curved bar +resembling the notochord in structure. The bars are segmented, and +each segment bears a smaller rod which supports a tentacle, the whole +forming the buccal skeleton (fig. 3, 4). + +The notochord is enclosed in a thick =sheath= of connective tissue +continuous with a thinner sheath round the nerve cord. The sheaths of +the notochord and nerve cord together form the skeletogenous layer, +and prolongations of it form the myomeres or septa between the +myotomes or segments of the great lateral muscles of the body. + +The =skeleton of each median fin= consists of small cubical masses of +a gelatinous substance arranged in rows (fig. 3, 1 and 7), and serving +to strengthen the fins. + + +FOOTNOTES: + +[19] The name _Balanoglossus_ is used here in its widest sense to +include all the Enteropneusta. + +[20] See W. Bateson, _Quart. J. Micr. Sci._ n. s. vol. XXIV. 1884, p. +208 and later; also E.W. Macbride, _Ibid._ vol. XXXVI. 1894, p. 385. + +[21] See p. 52. + +[22] A.T. Masterman, _P.R. Soc. Edinb._ 1895-96, p. 59; and _Anat. +Anz._ 1896, p. 266. + +[23] See E. Ray Lankester, _Quart. J. Micr. Sci._ vol. XXIX. n. s. +1889, p. 365. W.B. Benham, _Ibid._ vol. XXXV. n. s. 1893, p. 97. J.W. +Kirkaldy, _Ibid_. vol. XXXVII. n. s. 1895, p. 303. The last-named +writer divides the genus into three subgenera. + + + + +CHAPTER IV. + +SUBPHYLUM D. VERTEBRATA. + + +THE animals included in this great group all possess an internal axial +skeleton forming the vertebral column or back-bone; and a dorsal +spinal cord. The vertebral column is developed from the skeletogenous +layer, which surrounds the spinal cord together with the notochord and +its sheath; and in the great majority of cases the notochord becomes +more or less modified and reduced in the adult. In some cases the +notochord remains unmodified and the skeletogenous layer surrounding +it is not segmented to form vertebrae, but in every case the neural +arches which protect the spinal cord are segmented. The notochord +never extends further forwards than the mid-brain. + +All true vertebrates possess a cranium or skeletal box enclosing the +brain. + + +(I.) CYCLOSTOMATA. + +The mouth in living forms is suctorial and is not supported by jaws. +In some fossil forms the character of the mouth is unknown. + + +_Order I._ MARSIPOBRANCHII[24]. + +In these animals limbs and limb girdles are always completely absent. +They have no exoskeleton except horny teeth. + +The endoskeleton, excluding the notochord, is entirely cartilaginous +or membranous. The axial skeleton consists of a cartilaginous cranium +without jaws, succeeded by a thick persistent notochord enveloped in +a sheath. The notochord in living forms is unsegmented, but segmented +cartilaginous neural arches are present in some cases. A complicated +series of cartilaginous elements occurs in relation to the mouth, +gills, and sense organs. The median fins are supported by +cartilaginous pieces, the radiale. The order includes the Lampreys and +Hags. + + +_Order II._ OSTRACODERMI[25]. + +The forms included in this group have long been extinct, being known +only from beds of Upper Silurian and Lower Devonian age. They differ +much from all other known animals. The exoskeleton is always greatly +developed and includes (1) large bony plates covering the anterior +region; (2) scales covering the posterior region. The plates are +deeply marked by canals belonging to dermal sense organs. Jaws are +unknown, and arches for the support of the appendicular skeleton are +rudimentary or absent. The tail is heterocercal (see p. 60). + + +_Suborder_ (1). HETEROSTRACI. + +The exoskeleton consists principally of calcifications forming dorsal +and ventral shields which cover the head and abdominal region; the +dorsal shield is formed of a few plates firmly united, the ventral +shield of a single plate. The shields are composed of three layers, +the middle layer being traversed by canals belonging to the dermal +sense organs which open to the exterior by a series of pores. The tail +is sometimes covered by scales. The orbits are widely separated and +laterally placed. Paired appendages are absent. These curious forms +are found in beds of Upper Silurian and Lower Devonian age. One of the +best known genera is _Pteraspis_. + + +_Suborder_ (2). OSTEOSTRACI. + +The exoskeleton as in the Heterostraci consists of shields and scales, +the shields being divisible into three layers. The anterior part of +the body is covered dorsally by a single large shield which differs +from those of the Heterostraci in having the inner layer ossified. The +middle layer contains canals for the passage of blood vessels, but the +exoskeleton shows no impressions of dermal sense organs. The posterior +part of the body is covered by large quadrangular scales. Paired +appendages are absent, but median dorsal and caudal fins occur +supported by scales, not fin-rays. _Cephalaspis_, the best known of +these animals, occurs in beds of Lower Devonian age. + + +_Suborder_ (3). ANTIARCHA. + +The exoskeleton is formed of bony plates, the dorsal and ventral +shields each consisting of several symmetrically arranged pieces. The +tail may be covered with small scales or may be naked. The head is +articulated with the trunk, and its angles are drawn out into a pair +of segmented paddle-like appendages, covered with dermal plates. The +orbits are close together. A dorsal fin and traces of mouth parts +occur in _Pterichthys_, but the endoskeleton is unknown. The best +known forms _Pterichthys_[26] and _Asterolepis_ occur in beds of Lower +Devonian age. + + +GENERAL ACCOUNT OF THE SKELETON OF MARSIPOBRANCHII. + +The Marsipobranchii are worm-like animals. The living forms include +two families, the Myxinoidei (Hags)--genera _Myxine_ and +_Bdellostoma_--and the Petromyzontidae (Lampreys). + +Three species of _Petromyzon_ are known, _P. fluviatilis_, _P. +marinus_ and _P. planeri_. The larval forms were for a long time +thought to belong to a separate genus and were called _Ammocoetes_. + +The Myxinoids, although very highly specialised in their own way, are +at distinctly a lower stage of development than the adult Lamprey, and +come nearer to the larval Lamprey or Ammocoete. + +SPINAL COLUMN. + +[Illustration FIG. 4. A, DORSAL; B, LATERAL AND C, VENTRAL VIEW OF THE +SKULL OF _Petromyzon marinus_ × 1 (after PARKER). + + 1. horny teeth. + 2. labial cartilage. + 3. anterior dorsal cartilage. + 4. posterior dorsal cartilage. + 5. nasal capsule. + 6. auditory capsule. + 7. dorsal portion of trabeculae. + 8. lateral distal mandibular. + 9. lingual cartilage. + 10. branchial basket. + 11. cartilaginous cup supporting pericardium. + 12. sheath of notochord. + 13. neural plate.] + +In Myxinoids and larval lampreys, the notochord is enclosed in a thick +chordal sheath, in connection with which in the tail region there +occur cartilaginous pieces forming neural arch elements. In the trunk +region, however, no cartilage occurs in connection with the spinal +column, the only cartilage present being that forming the radiale of +the dorsal fin. On the other hand in most species of lamprey +(_Petromyzon_) cartilaginous pieces forming imperfect neural arches +(fig. 4, B, 13) are found lying in the tough skeletogenous layer +dorsal to the notochord, and extending throughout the whole length of +the trunk and tail. Two of these pieces, which are probably homologous +with the neural plates (see p. 72) of Elasmobranchs, occur to each +_neuromere_, or segment as determined by the spinal nerves. The +dorsal and caudal fins are supported by paired cartilaginous radiale +which are connected proximally with the skeletogenous layer. + +THE SKULL. + +In Myxinoids the cranium is a mere cartilaginous floor without side +walls or roof, and the trabeculae[27] end without growing forwards +into cornua. In Lampreys the trabeculae grow forwards and send up +plates of cartilage which meet above (fig. 4, 7) and form side walls +and a roof for part of the brain case. In Lampreys a labial suctorial +apparatus is well developed, including a large ring-like piece of +cartilage (fig. 4, 2) which supports the oral funnel and bears a large +armament of horny teeth. In Myxinoids on the other hand the labial +skeleton is small and consists merely of barbels round the mouth. + +The olfactory organ of Myxinoids has a very curious skeleton. It is +covered with a kind of grating of cartilage which is prolonged in +front into a tube composed of a series of imperfect cartilaginous +rings. In Lampreys the olfactory organ opens merely by a short +membranous passage. In correlation with the small development of the +labial suctorial apparatus in Myxinoids the lingual apparatus is very +greatly developed. The tongue in _Myxine_ has been said to 'dominate +the whole body' (Parker). It is supported by a great median +cartilaginous bar which when followed forwards first becomes bifid and +still further forwards becomes four-cleft. + +The horny teeth in Myxinoids are chiefly borne on the very large +supralingual apparatus. They form a double series arranged in the form +of an arch. In _Myxine_ there are seven large teeth and nine small +ones on each side. In _Bdellostoma_ the teeth of the two rows are more +equal in size. In _Bdellostoma_ and _Myxine_ it has been shown that +imperfect calcified teeth occur below the horny teeth. + +In Lampreys the lingual apparatus (fig. 4, C, 9) is well developed, +but not excessively so. It consists of a long median cartilaginous bar +which ends in front with a semicircular piece of cartilage supporting +the median part of the tongue. + +In both Myxinoids and Lampreys there is a complicated branchial basket +apparatus, but while in Myxinoids the basket apparatus is +interbranchial, formed deep within the head near the hypoblastic +lining of the throat, in Lampreys it is extra-branchial and formed +outside the head cavities (fig. 4, 10). The two sides of the basket +apparatus in _Myxine_ are not symmetrical. In the interbranchial +basket apparatus of Myxinoids the hyoid and first and second branchial +arches can be recognised. Traces of the interbranchial skeleton of +Myxinoids can be detected in Lampreys, and similarly in Myxinoids, +there are indications of the extra-branchial skeleton of _Petromyzon_. +The branchial basket in Lampreys forms at its posterior end a kind of +cup which supports the pericardium (fig. 4, 11). + +A remarkable Cyclostome named _Palaeospondylus_[28] has recently been +described from the Scottish Old Red Sandstone. It differs however from +all living Cyclostomes, in having a spinal column formed of distinct +vertebrae with well-developed neural arches. The caudal fin is well +developed and the dorsal radiale are forked as in lampreys. The skull +is well calcified and the auditory capsules are specially large. The +mouth is very similar to that of lampreys, being circular and without +jaws; it is provided with barbels or cirri. There is no trace of limbs +and the average length is only about 1-1-1/2 inches. + + +FOOTNOTES: + +[24] See W.K. Parker On the skeleton of the Marsipobranch fishes, +_Phil. Trans._ 1883, London. + +[25] See A. Smith Woodward, _Catalogue of Fossil Fish in the British +Museum_, Part II., 1891. A. Smith Woodward, _Nat. Sci._ vol. I. 1892, +p. 596. + +[26] See R.H. Traquair, _Ann. Nat. Hist._, ser. 6, vol. II. 1888, p. +485. + +[27] See p. 17. + +[28] R.H. Traquair, _Ann. Nat. Hist._ vol. VI. 1890, p. 485; _P. Phys. +Soc. Edinb._ vol. XII. 1892-93, pp. 87-94, and 312-320. A. Smith +Woodward, _Nat. Sci._ vol. III. p. 128, 1893. + + + + +CHAPTER V. + +(II.) GNATHOSTOMATA. + + +THE mouth is supported by definite jaws. + + +ICHTHYOPSIDA. + +The epiblastic exoskeleton is generally unimportant, the mesoblastic +exoskeleton is usually well developed. + +The notochord with its membranous sheath (1) may remain unmodified, or +(2) may be replaced by bone or cartilage derived from the +skeletogenous layer, or (3) may be calcified to a varying extent. + +The first vertebra is not homologous throughout the whole series and +so is not strictly comparable to the atlas of Sauropsids and Mammals. + +The centra of the vertebrae have no epiphyses. The skull may be (_a_) +incomplete and membranous, or (_b_) more or less cartilaginous, or +(_c_) bony. Membrane bones are not included in the cranial walls, and +there are large unossified tracts in the skull. When membrane bones +are developed in connection with the skull, a large parasphenoid +occurs. The basisphenoid is always small or absent. The skull may be +immovably fixed to the vertebral column, or may articulate with it by +a single or double occipital condyle. When the occipital condyle is +double, it is formed by the exoccipitals, and the basi-occipital is +small or unossified. The mandible may be (_a_) cartilaginous, (_b_) +partially ossified, or (_c_) membrane bones may be developed in +connection with it,--if so, there is usually more than one membrane +bone developed in connection with each half. + +There are at least four pairs of branchial arches present during +development. The sternum, if present, is not costal in origin. + + +CLASS I. PISCES. + +The exoskeleton is in the form of scales, which may be entirely +mesoblastic or dermal in origin (e.g. _cycloid_ and _ctenoid_ scales), +or may be formed of both mesoblast and epiblast (e.g. _placoid_ and +_ganoid_ scales). Large bony plates may be derived from both these +types of scale. In general fish with a greatly developed dermal armour +have the endoskeleton poorly developed; and the converse also holds +good. + +The integument of the dorsal and ventral surfaces is commonly +prolonged into longitudinal unpaired fins, supported by an internal +skeleton. These fins are distinguished according to their position as +dorsal, caudal and anal fins. The dorsal and anal fins are used +chiefly as directing organs, the caudal fin is however a most +important organ of propulsion. + +Three types of tail are found in fishes, viz.:-- + +1. The =diphycercal=, in which the axis is straight and the tail is +one-bladed and symmetrical, an equal proportion of radiale[29] being +attached to the upper and lower surfaces of the axis. + +2. The =heterocercal=, in which the tail is asymmetrical and the axis +is bent upwards, the proportion of radiale or of fin-rays attached to +its upper surface being much smaller than that attached to its lower +surface. + +3. The =homocercal=, in which the tail though externally symmetrical, +so far resembling the diphycercal type, is internally really +heterocercal, the great majority of the radiale or of the fin-rays +being attached to the lower surface of the axis. + +The cranium in the simplest cases (e.g. Selachii) forms a +cartilaginous box enclosing the brain and sense organs; in bony fishes +it is greatly complicated. When palatine or pterygoid bones are +present they are formed by the ossification of cartilage; in +Sauropsida and Mammalia they are laid down as membrane bones. There is +no tympanic cavity or auditory ossicle in relation to the ear. + +There are two principal types of suspensorium by means of which the +jaws are attached to the cranium:-- + +(1) The =Autostylic=. This is the primitive condition in which the +mandibular arch articulates with the base of the cranium in front of +the hyoid and in a similar manner. + +(2) The =Hyostylic=. In this case the mandibular arch becomes +connected with the hyomandibular and supported by the hyoid arch. +These terms are more fully discussed in Chapter VIII. + +There is always an internal framework supporting the gills; it usually +consists of the hyoid arch and five, rarely six or seven, pairs of +branchial arches. The limbs are represented by two pairs of fins, the +pectoral and the pelvic; they are not divided into proximal, middle +and distal portions. The ribs do not unite with a median ventral +sternum, or meet in the mid-ventral line in any other way in the trunk +region. + + +_Order I._ ELASMOBRANCHII. + +The exoskeleton is in the form of placoid scales which are sometimes +so numerous as to give the whole skin a rough surface forming +shagreen. In some cases the placoid scales are enlarged to form plates +or spines capped or coated with enamel. These spines may be imbedded +in the flesh in front of the paired or unpaired fins, or may be +attached to the tail. They are specially characteristic of the +suborder Acanthodii. The endoskeleton is cartilaginous and true bone +is never found. Much of the skeleton, especially of the vertebral +column, is however often calcified, this being especially well seen in +the anterior part of the vertebral column of Rays (Raiidae). In living +forms cartilaginous biconcave vertebrae are always well developed, but +in some extinct forms the notochord persists unconstricted. Neural and +haemal arches are however always developed; they sometimes remain +separate, sometimes fuse with the centra. Ribs are often wanting and +when present are often not separated off from the vertebrae. The +cranium is a simple cartilaginous box whose most prominent parts are +the capsules which enclose the sense organs. The skull is sometimes +immovably fixed to the vertebral column, sometimes articulates with it +by means of two condyles. There is no operculum and no representative +of the maxilla or premaxillae. The teeth are very variable. Large +pectoral and pelvic fins always occur. + +The Elasmobranchii may be divided into four suborders:-- + + (1) Ichthyotomi. + (2) Pleuropterygii. + (3) Selachii. + (4) Acanthodii. + + +_Suborder_ (1). ICHTHYOTOMI[30]. + +The members of this suborder range from the Devonian to the Permian +and so have long been extinct. + +The endoskeletal cartilage has granular calcifications evenly +distributed throughout it. The notochord is unconstricted, but the +neural and haemal arches are well-developed, and the neural spines are +long and slender. There is a continuous dorsal fin with separate +basalia and radiale. The tail is diphycercal, and the pectoral fins +are typical archipterygia[31]. The pelvic fins of the male are +prolonged to form claspers. + +The best known of these primitive Elasmobranchs are the +Pleuracanthidae. + + +_Suborder_ (2). PLEUROPTERYGII. + +This suborder was formed for the reception of _Cladoselache_, an +Elasmobranch found in the Lower Carboniferous of Ohio[32]. + +The exoskeleton is in the form of small, thickly-studded dermal +denticles. The vertebral centra are unossified, and the tail is +strongly heterocercal. There were certainly five, perhaps seven gill +slits, and the suspensorium is apparently hyostylic. The paired fins +are, according to the view which derives them by concentration from +continuous lateral folds, the most primitive known (see p. 129) and +claspers are absent. + + +_Suborder_ (3). SELACHII. + +Cartilaginous or partially calcified biconcave vertebrae are always +well developed; they constrict the notochord intervertebrally. The +neural and haemal arches and spines are stout and intercalary +cartilages (interdorsalia) are present. The tail is heterocercal, but +in some cases (_Squatina_) approaches the diphycercal condition. In +most cases the suspensorium is hyostylic, the jaws being attached to +the cranium by means of the hyomandibular, and the palato-pterygo +quadrate bar not being fused to the cranium. There are generally five +pairs of branchial arches, and gill rays are borne on the posterior +surface of the hyoid arch, and on both the anterior and posterior +surfaces of the first four branchial arches. The Notidanidae differ +from most Selachians in two respects, first as regards the +suspensorium,--Meckel's cartilage articulating directly with the +palato-pterygo-quadrate bar, and not being connected with the hyoid +arch; and secondly as regards the number of branchial arches,--six +pairs occurring in _Hexanchus_ and seven in _Heptanchus_. + +The pectoral fins are without the segmented axis of the +archipterygium. In most cases they are sharply marked off from the +body and lie almost at right angles to it; but in the Rays they have +the form of lateral expansions in the same plane as the body, from +which they are not sharply marked off. The pelvic fins in the male +bear long grooved cartilaginous rods which are accessory copulatory +organs or claspers. + +There are two principal groups of Selachii, the Squalidae or Sharks +and Dogfish, and the Batoidei or Skates and Rays. The Squalidae have +the shape of ordinary fish, the pectoral fins are vertically placed +and the body ends in a powerful heterocercal tail. The Batoidei have +flattened bodies owing to the great size and horizontal position of +the pectoral fins. The tail is long and thin and is often armed with +spines. The teeth in Selachii differ much in character in the +different forms, and are always arranged in numerous rows. They are +generally pointed and triangular or conical in the Squalidae, while in +the Batoidei they are often broad and flattened. + + +_Suborder_ (4). ACANTHODII. + +The fishes included in this group are all extinct and in some respects +are intermediate between Elasmobranchii and Ganoidei. The body is +elongated and closely covered with small scales consisting of dentine +enamelled at the surface. The notochord is persistent and the +calcification of the endoskeletal cartilage is only superficial. The +tail is heterocercal. The jaws bear small conical teeth, or in some +cases are toothless. The skeleton of all the fins differs from that of +modern Elasmobranchs in having the cartilaginous radiale much reduced, +and the fins are nearly always each provided with an anterior spine, +which except in the case of the pectoral fins is merely inserted +between the muscles. These spines are really enormous dermal fin-rays; +the pectoral fin-spine is articulated to the pectoral girdle. + +The suborder includes many well-known extinct forms like _Acanthodes_ +and _Diplacanthus_; it ranges from the Devonian to the Permian. + + +PISCES, HOLOCEPHALI. + +_Order II._ HOLOCEPHALI. + +This order includes a single suborder only. + + +_Suborder._ CHIMAEROIDEI. + +[Illustration FIG. 5. SKULL OF A MALE _Chimaera monstrosa_ (after +HUBRECHT). + + 1. nasal capsule. + 2. cartilaginous appendage to the fronto-nasal region. + 3. erectile appendage. + 4. foramen by which the ophthalmic nerves leave the orbit. + 5. foramen by which the ophthalmic branch of the Vth nerve enters + the orbit. + 6. auditory capsule. + 7. interorbital septum. + 8. mandible articulating with an outgrowth from the posterior part + of the palato-pterygo-quadrate. + 9. teeth. + 10. labial cartilage. + II. III. V. VII. IX. X. foramina for the passage of cranial nerves.] + +These singular fish have the skin smooth and in living forms almost or +quite scaleless. The palato-pterygo-quadrate bar and hyomandibular are +fused to the cranium, and Meckel's cartilage articulates directly with +the part corresponding to the quadrate. The skull is distinctly +articulated with the spinal column, the notochord is persistent and +unconstricted, and the skeletogenous layer shows no trace of metameric +segmentation, though in the neural arches this segmentation is +readily traceable. The neural arches of the first few vertebrae are +fused together and completely surround the notochord, while they do +not in other parts of the body. The tail is diphycercal. Of the living +genera, in _Callorhynchus_ there is no trace of calcification in the +skeletogenous layer, while in _Chimaera_ rings of calcification are +found, there being three to five for each vertebra as indicated by the +foramina for the exit of the spinal nerves. The pelvic fins are +produced into claspers. Besides the living genera _Chimaera_, +_Harriotta_ and _Callorhynchus_ a fair number of fossil forms are +known, e.g. _Ischyodus_. + + +_Order_ III. GANOIDEI. + +The fishes included under the term Ganoidei form a very heterogeneous +group, some of which closely approach the Dipnoi, others the +Elasmobranchii, others the Teleostei. The great majority of them are +extinct, only eight living genera being known; these are all +inhabitants of the northern hemisphere, and with the exception of +_Acipenser_, which is both fluviatile and marine, are entirely +confined to fresh water. + +The following is a list of the living genera of Ganoids with their +respective habitats:-- + + _Acipenser._ Rivers and seas of the northern hemisphere. + + _Scaphirhynchus._ Mississippi and rivers of Central Asia. + + _Polyodon_ (_Spatularia_). Mississippi. + + _Psephurus._ Yan-tse-kiang, and Hoangho. + + _Polypterus._ Rivers of tropical Africa. + + _Calamoichthys._ Some rivers of West Africa. + + _Lepidosteus._ Freshwaters of Central and North America and + Cuba. + + _Amia._ Rivers of Carolina. + +The exoskeleton is very variable, thus the body may be:-- + +(_a_) Naked or with minute stellate ossifications as in the +Polyodontidae. (_b_) Partially covered with large detached bony +plates as in _Scaphirhynchus_ and _Acipenser_. (_c_) Entirely covered +with rhomboidal ganoid scales as in _Lepidosteus_, _Polypterus_, +_Palaeoniscus_ and many extinct forms. (_d_) Covered with rounded +scales shaped like the cycloid scales of Teleosteans as in _Amia_. +(_e_) Having the trunk and part of the tail covered with rhomboidal +scales, and the remainder of the tail with rounded scales as in +_Trissolepis_. + +The teeth also are very variable. The endoskeleton shows every stage +of transition from an almost entirely cartilaginous state as in +_Acipenser_ to a purely bony state as in _Lepidosteus_. Sometimes, as +in _Acipenser_, the notochord persists, and its sheath is unsegmented; +sometimes, as in _Lepidosteus_, there are fully formed vertebrae. The +tail may be heterocercal, as in _Acipenser_, or diphycercal as in +_Polypterus_. The cartilaginous cranium is always covered with +external membrane bone to a greater or less extent, and the +suspensorium is markedly hyostylic. The pectoral girdle is formed of +two parts, one endoskeletal and cartilaginous, corresponding with the +pectoral girdle of Elasmobranchs, and one exoskeletal and formed of +membrane bones, corresponding with the clavicular bones of +Teleosteans. The pelvic fins are always abdominal. The fins often, as +in _Polypterus_, have spines (fulcra) attached to their anterior +borders. + +The order Ganoidei may be divided into three suborders: + + (1) CHONDROSTEI. Living genera _Acipenser_, _Scaphirhynchus_, + _Polyodon_ and _Psephurus_. + + (2) CROSSOPTERYGII. Living genera _Polypterus_ and + _Calamoichthys_. + + (3) HOLOSTEI. Living genera _Lepidosteus_ and _Amia_. + + +_Suborder_ (1). CHONDROSTEI. + +The skull is immovably fixed to the vertebral column. By far the +greater part of the skeleton is cartilaginous. The notochord is +persistent and unconstricted, its sheath is membranous, but +cartilaginous neural and haemal arches are developed. Intercalary +pieces (interdorsalia) occur between the neural arches, and similar +pieces (interventralia) between the haemal arches. The cranium is +covered with membrane bone, and teeth are but slightly developed. The +tail is heterocercal. Gill rays occur on the hyoid arch, and the gills +are protected by a bony operculum attached to the hyomandibular. The +skin (1) may be almost or quite naked, (2) may carry bony plates +arranged in rows, or may be covered (3) with rhomboidal scales, or (4) +partly with rhomboidal, partly with cycloidal scales. + + +_Suborder_ (2). CROSSOPTERYGII. + +The exoskeleton has the form of cycloidal or rhomboidal scales. The +condition of the vertebral column differs in the different genera. +Sometimes, as in _Polypterus_, there are well-developed ossified +vertebrae; sometimes, as in many extinct forms, the notochord persists +and is unconstricted. The tail may be diphycercal or heterocercal. The +pectoral and sometimes the pelvic fins consist of an endoskeletal axis +bearing a fringe of dermal rays. + + +_Suborder_ (3). HOLOSTEI. + +The exoskeleton has the form of cycloidal or rhomboidal scales. The +notochord is constricted and its sheath is segmented and ossified, +forming distinct vertebrae, which are generally biconcave, sometimes +opisthocoelous (_Lepidosteus_). The cartilaginous cranium is largely +replaced by bone, and in connection with it we find not only membrane +bone, but cartilage bone, as the basi-occipital, exoccipitals, and +pro-otic are ossified. The tail is heterocercal. The suspensorium +resembles that of Teleosteans, consisting of a proximal ossification, +the hyomandibular, which is movably articulated to the skull and a +distal ossification, the symplectic. The two are separated by some +unossified cartilage. The cartilaginous upper and lower jaws are to a +great extent surrounded and replaced by a series of membrane bones. + + +_Order_ IV. TELEOSTEI. + +The exoskeleton is sometimes absent but generally consists of +overlapping cycloid or ctenoid scales. Bony plates are sometimes +present, as in the Siluridae, or the body may be encased in a complete +armour of calcified plates, as in _Ostracion_. Enamel is however never +present, and the plates are entirely mesodermal. The skeleton is bony, +but in the skull much cartilage generally remains. The vertebral +centra are usually deeply biconcave, and the tail is of the masked +heterocercal type distinguished as _homocercal_. In the skull the +occipital region is always completely ossified, while the sphenoidal +region is generally less ossified. The skull has usually a very large +number of membrane bones developed in connection with it. The teeth +vary much in character in the different members of the order, but are +as a rule numerous and pointed, and are ankylosed to the bone. The +suspensorium is hyostylic and the jaws have much the same arrangement +as in the Holostei. There are five pairs of branchial arches, of which +all except the last bear gill rays. A series of dermal opercular bones +is developed in connection with these arches. The pectoral girdle +consists almost entirely of dermal clavicular bones. The pelvic girdle +has disappeared, its place being taken by the enlarged and ossified +dermal fin-rays of the pelvic fins. + +The group includes the vast majority of living fish (see p. 33). + + +_Order_ V. DIPNOI. + +The exoskeleton is of two types; dermal bones are largely developed in +the head region, while the tail and posterior part of the body may be +naked or may be covered with overlapping scales. The cranium remains +chiefly cartilaginous, the palato-pterygo-quadrate bar is fused with +the cranium, and the suspensorium is autostylic. The gill clefts are +feebly developed and open into a cavity covered by an operculum. The +notochord is persistent and unconstricted, and the limbs are +archipterygia. The pelvic fins are without claspers. + + +_Suborder_ (1). SIRENOIDEI[33]. + +The head has well developed membrane bones. The trunk is covered with +overlapping scales and bears no bony plates. Three pairs of teeth are +present, two in the upper and one in the lower jaw, the two principal +pairs of teeth are borne on the palato-pterygoids and splenials, while +the third pair are found in the vomerine region. The tail is +diphycercal in living forms. In the extinct Dipteridae it is +heterocercal. The pectoral girdle includes both membrane and cartilage +bones. The pelvic girdle consists of a single bilaterally symmetrical +piece of cartilage. + +This suborder is represented by the living genera _Ceratodus_, +_Protopterus_ and _Lepidosiren_, and among extinct forms by the +Dipteridae and others. + + +_Suborder_ (2). ARTHRODIRA. + +Bony plates are developed not only on the head but also on the +anterior part of the trunk, where they consist of a dorsal, a ventral, +and a pair of lateral plates which articulate with the cranial shield. +The posterior part of the trunk is naked. The tail is diphycercal. The +jaws are shear-like, and their margins are usually provided with +pointed teeth whose bases fuse with the tissue of the jaw and +constitute dental plates. There seem to have been three pairs of these +plates, arranged as in the Sirenoidei, the principal ones in the upper +jaw being borne on the palato-pterygoids. Small pelvic fins are +present, but pectoral fins are unknown. + +The Arthrodira occur chiefly in beds of Devonian and Carboniferous +age. Two of the best known genera are _Coccosteus_ from the European +Devonian and _Dinichthys_, a large predatory form from the lower +Carboniferous of Ohio. + + +FOOTNOTES: + +[29] See p. 79. + +[30] For this and other groups of extinct fish see A. Smith Woodward, +_Catalogue of Fossil Fish in the British Museum_, Parts I.-III. +London, 1889-95. + +[31] See p. 127. + +[32] See B. Dean, _J. Morphol._ vol. IX. pp. 87-114, 1894, and _Nat. +Sci._ vol. VIII. p. 245, 1896. + +[33] A. Günther, _Phil. Trans._ vol. 161, Part II. 1871, p. 511. T.H. +Huxley, "On Ceratodus and the classification of fishes," _P.Z.S._ +1876, p. 24. + + + + +CHAPTER VI. + +THE SKELETON OF THE DOGFISH[34]. + +_Scyllium canicula._ + + +I. EXOSKELETON. + +The exoskeleton of the dogfish is mainly composed of placoid scales, +each of which consists of a little bony base imbedded in the skin, +bearing a small backwardly-directed spine formed of dentine capped +with enamel. The scales are larger on the dorsal than on the ventral +surface, and on the jaws they are specially large and regularly +arranged in rows, there forming the teeth. The margins of the jaws or +lips are without scales. + +A second exoskeletal structure is found in the fins, all of which, +both paired and unpaired, have, in addition to their cartilaginous +endoskeleton, large numbers of long slender horny fibres, the +fin-rays, which are of exoskeletal origin. + + +II. ENDOSKELETON. + +The endoskeleton of the dogfish consists almost entirely of cartilage, +which however may become calcified in places, e.g. the centrum of each +vertebra is lined by a layer of calcified tissue. + +The endoskeleton is divisible into an =axial= portion consisting of +the vertebral column, skull, and skeleton of the median fins, and an +=appendicular= portion consisting of the skeleton of the paired fins +and their girdles. + + +1. THE AXIAL SKELETON. + +A. THE VERTEBRAL COLUMN AND RIBS. + +The vertebral column consists of a series of some hundred and thirty +vertebrae, each of which is united with its predecessor and successor +in such a way as to allow a large amount of flexibility. + +These vertebrae are developed round an unsegmented rod, the +=notochord=, which forms the axial support of the embryo. The +notochord remains continuous throughout the whole vertebral column, +but is greatly constricted opposite the middle of each vertebra, and +thus rendered moniliform. The vertebrae are divided into two groups, +an anterior group of trunk vertebrae, and a posterior group of caudal +or tail vertebrae. + +A typical vertebra consists of a middle portion, the =centrum=, a +dorsal portion, the =dorsal= or =neural arch=, which surrounds the +spinal cord, and a ventral portion, the =ventral= or =haemal arch=, +which similarly encloses a space. + +The tail vertebrae of the dogfish have this typical arrangement, the +trunk vertebrae have the haemal arches modified. + +Each =centrum= is a short cylinder of cartilage surrounding an +hourglass-shaped cavity occupied by the notochord. The =neural arches= +are composed of three separate elements, the =vertebral neural plates= +(basidorsalia), =intervertebral neural plates= (interdorsalia), and +=neural spines= (supradorsalia). + +The =vertebral neural plates= are in the adult fused with their +respective centra, and are notched behind for the exit of the ventral +(motor) roots of the spinal nerves. The =intervertebral neural plates= +are polygonal pieces alternating with the vertebral neural plates; +they are notched behind, but at a more dorsal level than are the +vertebral neural plates, for the exit of the dorsal or sensory roots +of the spinal nerves. + +The =neural spines= are small patches of cartilage filling up the gaps +between the dorsal ends of the neural plates. + +The =haemal arches= (basiventralia) differ much in the trunk and tail +portions of the vertebral column. In the trunk portion the centra are +flattened below, and the two halves of the haemal arch diverge from +one another as blunt =ventri-lateral processes= to +which short cartilaginous rods, the =ribs=, are attached. Further back +at about vertebra 37, the two halves of the haemal arch project +downwards and meet forming a complete arch. Further back still, +towards the hind end of the tail, the haemal arches bear median +=haemal spines= (ventrispinalia). + + +B. THE SKULL. + +The skull of the dogfish remains cartilaginous throughout the life of +the animal, and has consequently a far more simple structure than have +the skulls of higher animals, in which complication has been produced +by the development of bone. + +The skull consists of the following parts:-- + +(1) a dorsal portion, the =cranium=, which lodges the brain, and to +the sides of which the capsules of the auditory and olfactory sense +organs are united. The cranium may be compared to an unsegmented +continuation of the vertebral column; + +(2) a number of ventral structures, disconnected or only loosely +connected with the cranium. These together constitute the =visceral +skeleton= forming the jaws and supporting the gills. + +(1) THE CRANIUM. + +The =Cranium= is an oblong box, with a flattened floor and a more +irregular roof. Its sides are expanded in front owing to the +olfactory capsules, and behind owing to the auditory capsules, while +in the middle they are deeply hollowed to form the orbits. + +(_a_) On the dorsal surface of the cranium the following points should +be noticed. First at the anterior end, the large thin-walled =nasal= +or =olfactory capsules= (fig. 6, 1), each of which is drawn out into a +narrow cartilaginous process. + +The olfactory capsules have no ventral walls, and are separated from +one another by the =internasal septum=, which is drawn out into a +third slender process. These three processes together constitute the +=rostrum= (fig. 6, 2). + +Behind the olfactory capsules comes a large, nearly circular, hole, +the =anterior fontanelle=, slightly behind which are the two +=ophthalmic foramina=. The dorsal and ventral boundaries of the orbits +are respectively formed by the prominent =supra-orbital= and +=suborbital ridges=. Behind are the =auditory capsules= (fig. 6, 8), +each of which is marked by a pair of prominent ridges, converging +towards the middle line to a pair of apertures. These apertures +communicate with two canals, the =aqueductus vestibuli=, which lead +into the internal ear. The two ridges lodge respectively the =anterior +and posterior vertical semicircular canals= of the ear. + +(_b_) The principal structures to be noted in a side view of the +cranium are contained in the =orbit= or eye-cavity. Near the base of +the orbit at its anterior end is seen the small =orbitonasal foramen= +(fig. 6, 7), for the passage of blood-vessels, not nerves. Above it is +the large =ophthalmic foramen= (fig. 6, 5) so prominent in a dorsal +view of the skull; through it the ophthalmic branches of the fifth and +seventh nerves pass. Slightly further back near the ventral surface is +the large =optic foramen= (fig. 6, II.) for the passage of the second +nerve. Vertically above the optic foramen, near the dorsal surface, is +the very small =foramen for the fourth nerve= (fig. 6, IV.). Behind +and a little above the optic foramen is another small aperture, the +=foramen for the third nerve=. Behind and slightly below this is the +large =foramen for the sixth and main branches of the fifth and +seventh nerves= (fig. 6, V.). In front of and slightly below this +foramen are seen two other small apertures; the more anterior and +ventral of these (fig. 6, 4) is for the passage of a vessel connecting +the efferent artery of the hyoid gill with the internal carotid artery +inside the skull, the more posterior and dorsal is for the +=interorbital canal= (fig. 6, 3) which unites the two orbital sinuses. +Above and very slightly in front of the large foramen for the sixth +and main parts of the fifth and seventh nerves, are two small foramina +(fig. 6, Va., and VIIa.), through which the =ophthalmic branches of +the fifth and seventh nerves= enter the orbit. Behind and slightly +below the large foramen just mentioned is a small hole through which +the external carotid enters the orbit (fig. 6, 9). + +[Illustration Fig. 6. LATERAL VIEW OF THE SKULL OF A DOGFISH +(_Scyllium canicula_) × 2/3.] + + 1. nasal capsule. + 2. rostrum. + 3. interorbital canal. + 4. foramen for hyoidean artery. + 5. foramen for the exit of the + ophthalmic branches of + Vth and VIIth nerves. + 6. foramen through which the + external carotid leaves the + orbit. + 7. orbitonasal foramen. + 8. auditory capsule. + 9. foramen through which the + external carotid enters the + orbit. + 10. ethmo-palatine ligament. + 11. palato-pterygo-quadrate bar. + 12. Meckel's cartilage. + 13. hyomandibular. + 14. cerato-hyal. + 15. pharyngo-branchial. + 16. epi-branchial. + 17. cerato-branchial. + 18. gill filaments, nearly all have + been cut off short for the + sake of clearness. + 19. extra-branchial. + 20. pre-spiracular ligament. + II. III. IV. V. Va. VIIa. foramina + for passage of cranial nerves. + +Behind the orbit is the =auditory capsule=. This is marked below by a +prominent =surface for the articulation of the hyomandibular=, above +which is the deep =postorbital groove= for the passage of a +blood-vessel, connecting the orbital and anterior cardinal sinuses. + +(_c_) Passing to the posterior end of the cranium: in the centre is +seen the large =foramen magnum= through which the brain and spinal +cord communicate. The =notochord= enters the skull just below this +foramen, and on each side of the notochord is a projection, the +=occipital condyle=, by which the first vertebra articulates with the +skull. + +External to the condyles are the prominent =pneumogastric foramina= +for the passage of the tenth nerves, and further to the sides, just +beyond the posterior vertical semicircular canals, are a pair of deep +pits in which lie the =foramina for the ninth nerves= (fig. 6, IX). + +(_d_) The broad and flat ventral surface of the cranium is continued +in front as the =internasal septum= and terminated laterally by the +=suborbital ridges=. At a little behind the middle it is traversed by +two shallow grooves along which the internal carotid arteries run. At +the divergent ends of these grooves are seen two small apertures +through which the external carotids enter the orbit (fig. 6, 9), and +at the point where they meet is a single small aperture through which +the internal carotid enters the cranium. + +(2) THE VISCERAL SKELETON. + +The =Visceral skeleton= forms a series of seven cartilaginous arches +or hoops, surrounding the anterior part of the alimentary canal, and +enclosing a wide but rather shallow space. + +(_a_) The first or =mandibular arch= is the largest of the series, and +forms the upper and lower jaws. Each half of the upper jaw or +=palato-pterygo-quadrate= bar is formed by a thick cartilaginous rod +which meets its fellow in the middle line in front, the two being +united by ligament. Each half is connected to the cranium just in +front of the orbit by the =ethmo-palatine ligament= (fig. 6, 10), and +at its hind end articulates with one of the halves of the lower jaw. +Each half of the lower jaw or =Meckel's cartilage= (fig. 6, 12) is a +cartilaginous bar, wide behind but narrow in front, where it is united +to its fellow by a median ligament. Imbedded in the tissue external to +the upper jaw are a pair of =labial cartilages=, and a similar but +smaller pair are imbedded in the tissue external to the lower jaw. + +The jaws are developed from a structure whose dorsal and ventral +portions subsequently become of very different importance. The ventral +portion forms both upper and lower jaws, the former being developed as +an outgrowth from the latter. The dorsal portion forms only the +=pre-spiracular ligament= (fig. 6, 20), a strong fibrous band +containing a nodule of cartilage, and running from the anterior part +of the auditory capsule to the point where the jaws are connected with +the hyomandibular. + +(_b_) The =hyoid arch= consists of a pair of cartilaginous rods which +are attached at their dorsal ends to the cranium, and are united +ventrally by a broad median plate of cartilage, the =basi-hyal=. Each +rod is divided into a dorsal portion, the =hyomandibular= and a +ventral portion, the =cerato-hyal=. The =hyomandibular= (fig. 6, 13) +is a short stout rod of cartilage projecting outwards, and somewhat +backwards and downwards from the cranium, with which it articulates +behind the orbit and below the postorbital groove. Its distal end +articulates with a rather long slender bar, the =cerato-hyal= (fig. 6, +14), which is in its turn attached to the side of the =basi-hyal=. The +=basi-hyal= is a broad plate, rounded in front and drawn out behind +into two processes to which the two halves of the first branchial arch +are attached. The posterior surfaces of both hyomandibular and +cerato-hyal bear slender cartilaginous processes, the =gill rays=. The +hyoid arch forms the main =suspensorium= or means by which the jaws +are attached to the cranium. This attachment is chiefly brought about +by a series of short ligaments which connect the posterior ends of +both upper and lower jaws with the hyomandibular, but there is also a +ligament connecting the lower jaw with the cerato-hyal. The attachment +of the jaws to the cranium is also partially effected by the +pre-spiracular and ethmo-palatine ligaments. + +(_c_) Each of the five =branchial arches= is a hoop, incomplete above +and formed of four or more pieces of cartilage. The most dorsal +elements, the =pharyngo-branchials=, are flattened, pointed plates +whose free inner ends run obliquely backwards, and terminate below the +vertebral column. They are connected at their outer ends with the +short broad =epi-branchials= (fig. 6, 16) which lie at the sides of +the pharynx. From the epi-branchials arise the long =cerato +branchials= (fig. 6, 17) which run forwards and inwards along the +ventral wall of the pharynx. The first four cerato-branchials are +connected with small rods, the =hypo-branchials=, which run backwards +to meet one another in the middle line. The last two pairs of +hypo-branchials and the fifth cerato-branchials are connected with a +broad median plate, the =basibranchial=. Along the outer sides of the +second, third and fourth cerato-branchials are found elongated curved +rods, the =extra-branchials= (fig. 6, 19). The epi-branchials and +cerato-branchials bear gill rays along their posterior borders. + + +C. THE SKELETON OF THE MEDIAN FINS. + +The =dorsal fins= have a skeleton consisting of a series of short +cartilaginous rods, the =basals= or basalia, which slope obliquely +backwards. Their bases are imbedded in the muscles of the back, while +their free ends bear a number of small polygonal cartilaginous plates, +the =radials= or radiale. Associated with this cartilaginous skeleton +are a number of long slender horny fibres, the fin-rays, which have +been already referred to in connection with the exoskeleton. The +skeleton of the other median fins mainly consists of these fibres, the +cartilaginous portion being reduced or absent. + + +2. THE APPENDICULAR SKELETON. + +This includes the skeleton of the two pairs of limbs and of their +respective girdles. + +THE PECTORAL GIRDLE forms a crescent-shaped hoop of cartilage, +incomplete above and lying just behind the visceral skeleton. The +mid-ventral part of the hoop is the thinnest portion, and is drawn out +in front into a short rounded process which is cupped dorsally and +supports part of the floor of the pericardium (fig. 7, 1). On each +side of this flattened mid-ventral portion the arch becomes very thick +and bears on its outer border a surface with which the three basal +cartilages of the fin articulate. The dorsal ends or scapular portions +of the girdle form a pair of gradually tapering horns. + +THE PECTORAL FIN articulates with the pectoral girdle by means of +three basalia or basal cartilages, the =propterygium=, =meso +pterygium= and =meta-pterygium=. The most anterior and the smallest of +these is the =propterygium= (fig. 7, 5), while the most posterior +one, the =meta-pterygium= (fig. 7, 3), is much the largest. Along the +outer borders of the three basalia are arranged a series of close set +cartilaginous pieces, the =radiale=. The propterygium supports only a +single radial, which is however much larger than any of the others. +The meso-pterygium also supports only a single radial which divides +distally. + +[Illustration FIG. 7. SEMIDORSAL VIEW OF THE PECTORAL GIRDLE AND FINS +OF A DOGFISH (_Scyllium canicula_) × 2/3. + +The gaps between the radiale are blackened. + + 1. hollow in the mid-ventral part + of the pectoral girdle which + supports the pericardium. + 2. dorsal (scapular portion) of + pectoral girdle. + 3. meta-pterygium. + 4. meso-pterygium. + 5. propterygium. + 6. propterygial radial. + 7. meso-pterygial radial. + 8. meta-pterygial radial. + 9. outline of the distal part of + the fin which is supported + by horny fin-rays.] + +The meta-pterygium bears about twelve long narrow radials, the first +nine of which are traversed by a transverse joint at about two-thirds +of the way from their origin. Succeeding the radials are a series of +small polygonal pieces of cartilage arranged in one or more rows and +attached to the ends of the radials, and finally the fin is completed +by the dermal fin-rays. + +[Illustration FIG. 8. DORSAL VIEW OF THE PELVIC GIRDLE AND FINS OF A +MALE DOGFISH (_Scyllium canicula_). + + 1. pelvic girdle. + 2. basi-pterygium. + 3. clasper. + 4. radiale.] + +THE PELVIC GIRDLE is much smaller than the pectoral. It is formed of a +stout nearly straight bar of cartilage placed transversely across the +ventral region of the body. The bar has no dorsal or lateral +extensions, and is terminated by short blunt processes. It bears on +its posterior surface a pair of facets with which the pelvic fins +articulate. + +THE PELVIC FIN is smaller and more simply constructed than is the +pectoral. It consists of a long, somewhat curved rod, the +=basi-pterygium= (fig. 8, 2), running directly backwards on the inner +side of the fin, and articulating in front with the pelvic girdle. +From its outer side arise a series of about fourteen parallel +cartilaginous radials which bear smaller polygonal pieces. The +anterior one or two of these radials may articulate independently with +the pelvic girdle. In the adult male dogfish the distal end of the +basi-pterygium bears a stout rod nearly as long as itself, and grooved +on the dorsal surface. This is the skeleton of the =clasper= (fig. 8, +3). + +FOOTNOTE: + +[34] See Marshall and Hurst's _Practical Zoology_, 4th ed. London, +1895, p. 214. + + + + +CHAPTER VII. + +THE SKELETON OF THE CODFISH. (_Gadus morrhua._) + + +I. EXOSKELETON. + +The exoskeleton includes + +(1) =Scales.= These are of the type known as =cycloid= and consist of +flat rounded plates composed of concentrically arranged laminae of +calcified matter, with the posterior margin entire. The anterior end +of each scale is imbedded in the skin and is overlapped by the +preceding scales. + +(2) The =teeth=. These are small, pointed, calcified structures +arranged in large groups on the premaxillae, mandible, vomer, and +superior and inferior pharyngeal bones. + +(3) The =fin-rays=. These are delicate, nearly straight bony rods +which support the fins. + + +II. ENDOSKELETON. + +The endoskeleton of the Codfish, though partially cartilaginous, is +mainly ossified. + +It is divisible into an =axial portion=, including the skull, +vertebral column, ribs, and skeleton of the median fins, and an +=appendicular portion=, including the skeleton of the paired fins and +their girdles. + +1. THE AXIAL SKELETON. + +A. THE VERTEBRAL COLUMN. + +This consists of a series of some fifty-two vertebrae, all completely +ossified. + + +It is divisible into two regions only, viz. the =trunk= region, the +vertebrae of which bear movable ribs, and the =caudal= or =tail= +region, the vertebrae of which do not bear movable ribs. + +=Trunk vertebrae.= + +These are seventeen in number; the ninth may be described as typical +of them all. It consists of a short deeply biconcave =centrum= whose +two cavities communicate by a narrow central canal. From the dorsal +surface of the anterior half of the centrum arise two strong plates, +the dorsal or =neural processes=, which are directed obliquely +backwards and meet forming the dorsal or =neural arch=. This is +produced into a long backwardly-directed dorsal or =neural spine=. + +From the lower part of the anterior edge of each neural arch arise a +pair of blunt triangular projections which overhang the posterior half +of the preceding centrum, and bear a pair of flattened surfaces which +correspond to the anterior or =prezygapophyses= of most vertebrae, +they differ however from ordinary prezygapophyses in the fact that +they look downwards and outwards. From the posterior end of the +centrum arise a pair of short blunt processes each of which bears an +upwardly- and inwardly-directed articulating surface corresponding to +a =postzygapophysis=. + +The two halves of the ventral arch form a pair of large +=ventri-lateral processes= which arise from the anterior half of the +centrum and pass outwards and slightly backwards and downwards. + +Behind these there arises on each vertebra a second outgrowth which is +small and flattened, and like the ventri-lateral process serves to +protect the air-bladder. The surface of the centrum is marked by more +or less wedge-shaped depressions, one in the mid-dorsal line, and two +on the ventral surface immediately mesiad to the bases of the +ventri-lateral process. There are also a number of smaller +depressions. + +The space between one centrum and the next is in the fresh skeleton +filled up by the gelatinous remains of the =notochord=. + +The first few vertebrae differ from the others in having very short +centra and no ventri-lateral processes. + +The first vertebra comes into very close relation to the posterior +part of the skull, articulating with the exoccipitals. In the next few +vertebrae the centra gradually lengthen, and at the fourth or fifth +vertebra the ventri-lateral processes appear and gradually increase in +size as followed back. They likewise gradually come to arise at a +lower level on the centrum, and also become more and more downwardly +directed, till at the last trunk vertebra they nearly meet. + +The =neural spines= of the anterior trunk vertebrae are much longer +than those of the posterior ones, that of the first vertebra being the +largest and longest of all, and articulating with the skull. The +spinal nerves pass out through wide notches or spaces between the +successive neural arches. + +=Caudal vertebrae.= + +The caudal vertebrae are about thirty-five in number, each consists of +a centrum with a slender backwardly-directed dorsal or neural arch, +similar to those of the posterior trunk vertebrae. The two halves of +the ventral or haemal arch however do not form outwardly-directed +ventri-lateral processes, but arise on the ventral surface of the +centrum, and passing downwards meet and enclose a space; they thus +form a complete canal, and are prolonged into a backwardly-directed +ventral or =haemal spine=. The anterior haemal arches are much larger +than the corresponding neural arches, but when followed back they +gradually decrease in size, till at about the twenty-fourth caudal +vertebra they are nearly as small as the neural arches. The last +caudal vertebra is succeeded by a much flattened =hypural= bone or +=urostyle=, which together with the posterior neural and haemal spines +supports the tail-fin. + +B. THE RIBS. + +The =ribs= are slender, more or less cylindrical bones attached to the +poster-dorsal faces of the ventri-lateral processes of all the trunk +vertebrae except the first and second. The earlier ones are thicker +and more curved; the later ones thinner and more nearly straight. The +ribs are homologous with the distal parts of the haemal arches of the +caudal vertebrae. + +Associated with the ribs are a second series of rib-like bones, the +=intermuscular bones=. These are slender, curved bones which arise +from the ribs or from the ventri-lateral processes at a distance of +about an inch from the centra, and curve upwards, outwards and +backwards. In the anterior region where the ventri-lateral processes +are short they arise from the ribs, further back they arise from the +ventri-lateral processes. + +C. THE UNPAIRED OR MEDIAN FINS. + +These are six in number, three being =dorsal=, one =caudal= and two +=anal=. + +The =dorsal= and =anal= fins each consist of two sets of structures, +the =fin-rays= and the =interspinous bones=. Each fin-ray forms a +delicate, nearly straight, bony rod which becomes thickened and +bifurcated at its proximal or vertebral end, while distally it is +transversely jointed and flexible, frequently also becoming more or +less flattened. + +The first dorsal fin has thirteen rays, the second, sixteen to +nineteen, the third, seventeen to nineteen. The first anal fin has +about twenty-two, the second anal fourteen. In each fin the posterior +rays rapidly decrease in size when followed back. + +The =interspinous bones= of the dorsal and anal fins alternate with +the neural and haemal spines respectively, and form short, +forwardly-projecting bones, each attached proximally to the base of +the corresponding fin-ray. + +The =caudal fin= consists of a series of about forty-three rays which +radiate from the posterior end of the vertebral column, being +connected with the urostyle or hypural bone, and with the posterior +neural and haemal spines without the intervention of interspinous +bones. Like the other fin-rays those forming the caudal fin are +transversely jointed, and are widened and frayed out distally. + +The tail-fin in the Cod is =homocercal=, i.e. it appears to be +symmetrically developed round the posterior end of the vertebral +column, though in reality a much greater proportion is attached below +the end of the vertebral column than above it. It is a masked +heterocercal tail. + +THE SKULL. + +Owing to the fact that very little cartilage remains in the skull of +the adult Codfish, its relation to the completely cartilaginous skull +of the Dogfish is not easily seen. Before describing it therefore, the +skull of the Salmon will be described, as it forms an intermediate +type. + +THE SKULL OF THE SALMON[36]. + +The Salmon's skull consists of (1) the =chondrocranium=, which remains +partly cartilaginous and is partly converted into cartilage bone, +especially in the occipital region, (2) a large series of plate-like +membrane bones. + +THE CHONDROCRANIUM. + +This is an elongated structure, wide behind owing to the fusion of the +large auditory capsules with the cranium, and elongated and tapering +considerably in front; in the middle it is much contracted by the +large orbital cavities. + +DORSAL SURFACE OF THE CRANIUM. + +In the centre of the posterior end of the dorsal surface is the +=supra-occipital= (fig. 9, A, 1) with a prominent posterior ridge. +It is separated by two tracts of unossified cartilage from the large +series of bones connected with the =auditory organ=. The first of +these is the =epi-otic= (fig. 9, 2), which is separated by only a +narrow tract of cartilage from the supra-occipital, and is continuous +laterally with the large =pterotic= (fig. 9, A, 3) which overlaps in +front a smaller bone, the =sphenotic= (fig. 9, 4). Both epi-otic and +pterotic are drawn out into rather prominent backwardly-projecting +processes. + +[Illustration FIG. 9. A. DORSAL AND B. VENTRAL VIEW OF THE CRANIUM OF +A SALMON (_Salmo salar_) from which most of the membrane bones have +been removed (after PARKER). Cartilage is dotted. + + 1. supra-occipital. + 2. epi-otic. + 3. pterotic. + 4. sphenotic. + 5. frontal. + 6. median ethmoid. + 7. parietal. + 8. lateral ethmoid. + 9. parasphenoid. + 10. vomer. + 11. exoccipital. + 12. opisthotic. + 13. alisphenoid. + 14. orbitosphenoid. + 16. foramen for passage of an artery. + 17. pro-otic. + 18. articular surface for hyomandibular. + +II. VII. IX. X. foramina for the passage of cranial nerves.] + +The greater part of the remainder of the dorsal surface is formed of +unossified cartilage which is pierced by three large vacuities or +=fontanelles=. The anterior fontanelle is unpaired, and lies far +forward near the anterior end of the long cartilaginous snout, the two +larger posterior ones lie just in front of the supra-occipital and +lead into the cranial cavity. In front of the orbit the skull widens +again, and is marked by two considerable =lateral ethmoid= (fig. 9, 8) +ossifications. In front of these are a pair of deep pits, the =nasal +fossae=, at the base of which are a pair of foramina through which the +olfactory nerves pass out; they communicate with a space, the =middle +narial cavity=, seen in a longitudinal section of the skull. + +The long cartilaginous snout is more or less bifid in front, +especially in the male (fig. 9). + +POSTERIOR END OF THE CRANIUM. + +The =foramen magnum= forms a large round hole leading into the cranial +cavity, and is bounded laterally by the two =exoccipitals= and below +by them, and to a very slight extent by the =basi-occipital=, the +three bones together forming a concave =occipital condyle= by which +the vertebral column articulates with the skull. + +The exoccipitals are connected laterally with a fourth pair of +auditory bones, the =opisthotics=, and just meet the epi-otics +dorsolaterally, while dorsally they are separated by a wide tract of +unossified cartilage from the supra-occipital. + +The opisthotics are connected laterally with the pterotics. + +SIDE OF THE CRANIUM. + +At the posterior end is seen the =basi-occipital= in contact above +with the =exoccipital=, which is pierced by a prominent foramen for +the exit of the tenth nerve. In front of this lies a small foramen, +sometimes double, for the ninth nerve. + +[Illustration FIG. 10. LATERAL VIEW OF THE CHONDROCRANIUM OF A SALMON +(_Salmo salar_) (after PARKER). A few membrane bones are also shown. +Cartilage is dotted. + + 1. supra-occipital. + 2. epi-otic. + 3. pterotic. + 4. opisthotic. + 5. exoccipital. + 6. basi-occipital. + 7. parasphenoid. + 8. sphenotic. + 9. alisphenoid. + 10. orbitosphenoid. + 11. lateral- or ectethmoid. + 12. olfactory pit; the vomerine teeth are seen just below. + 14. pro-otic. + 15. basisphenoid. + 16. foramen for the passage of an artery. + 17. anterior fontanelle. + 18. posterior fontanelle. + +I. II. V. VII. IX. X. foramina for the passage of cranial nerves.] + +In front of the exoccipital is the large =pro-otic= pierced by two +prominent foramina. Through the more dorsal of these (fig. 10, VII.) +the facial nerve passes out, while the more ventral (fig. 10, 16) is +for the passage of an artery. Dorsal to the exoccipital are the +=opisthotic= and =pterotic=, and dorsal to the pro-otic is the +=sphenotic=. The =pterotic= is marked by a prominent groove often +lined by cartilage, which is continued forwards along a tract of +cartilage between the pro-otic and sphenotic. With this groove the +hyomandibular articulates. + +There are considerable ossifications in the sphenoidal region of the +side of the cranium. The anterior boundary of the posterior fontanelle +is formed by the large =alisphenoid=, which is continuous behind with +the pro-otic and sphenotic, and below with a slender =basisphenoid=. +Both in front of and behind the basisphenoid there are considerable +vacuities in the walls of the cranium; through the posterior of these +openings (fig. 10, V.) the main part of the trigeminal nerve passes +out, and through the anterior one, the optic (fig. 10, II.). The +alisphenoid is continuous in front with the =orbitosphenoid= (fig. 10, +10), which is pierced by the foramen for the exit of the first nerve +(fig. 10, I.), and in front of the orbitosphenoid there is a large +vacuity. The =lateral ethmoid= is seen in the side view as well as in +the dorsal view. Further forwards are seen the olfactory pits, and the +long cartilaginous snout. + +A =ventral view= of the cartilaginous cranium shows much the same +points as the side view. The basisphenoid appears on the surface +immediately in front of the basi-occipital. + +THE SKULL WITH MEMBRANE BONES. + +The =dorsal surface=. The greater part of the dorsal surface in front +of the supra-occipital is overlaid by a pair of large rough _frontals_ +(figs. 9, A, 5, and 10, 5). They cover the posterior fontanelles and +stretch over from the sphenotic to the lateral ethmoid, forming a roof +for the orbit. They meet in the middle line behind, but in front are +separated by a narrow tract of unossified cartilage, and are +overlapped by the _median ethmoid_ (figs. 9, A, 6, and 11, 6). At the +sides of the supra-occipital behind the frontals are a pair of small +_parietals_ (figs. 9, A, 7, and 11, 7). + +[Illustration FIG. 11. LATERAL VIEW OF THE SKULL OF A SALMON (_Salmo +salar_) (after PARKER). Cartilage is dotted. + + 1. supra-occipital. + 2. epi-otic. + 3. pterotic. + 4. sphenotic. + 5. frontal. + 6. median ethmoid. + 7. parietal. + 8. nasal. + 9. lachrymal. + 10. suborbital. + 11. supra-orbital. + 12. cartilaginous sclerotic. + 13. ossification in sclerotic. + 14. meso-pterygoid. + 15. meta-pterygoid. + 16. palatine. + 17. jugal. + 18. quadrate. + 19. maxillae + 20. premaxillae. + 21. articular. + 22. angular. + 23. dentary. + 24. hyomandibular. + 25. symplectic. + 26. epi-hyal. + 27. cerato-hyal. + 28. hypo-hyal. + 29. glosso-hyal. + 30. opercular. + 31. sub-opercular. + 32. infra-opercular. + 33. pre-opercular. + 34. supratemporal. + 35. branchiostegal rays. + 36. basi-branchiostegal.] + +In a =ventral view= the cranium is seen to be chiefly covered by two +large membrane bones, the _parasphenoid_ (fig. 9, B, 9) behind, the +_vomer_ in front. A view of the =posterior end= differs from that of +the cartilaginous cranium only in the fact that the end of the +_parasphenoid_ appears lying ventral to the basi-occipital. + +The =lateral view= differs very markedly from that of the +cartilaginous cranium, there being a great development of membrane +bone in connection with the jaws and branchial apparatus. Lying +dorsally are seen the _median ethmoid_, _frontal_, _parietal_, and +=supra-occipital= as before. Lying external to the middle of the +_median ethmoid_ is seen the small _nasal_ (fig. 11, 8), and below the +hinder part is the _lachrymal_. The _lachrymal_ (fig. 11, 9) forms the +first of a series of seven small bones which surround the orbit +forming the =orbital ring=. Of these the one lying immediately in the +mid-ventral line of the orbit is the _suborbital_, while the one lying +in the mid-dorsal line and attached to the frontal is the +_supra-orbital_ (fig. 11, 11). The orbit has a cartilaginous +_sclerotic_ in which are two small ossifications (fig. 11, 13) +laterally placed. + +BONES OF THE UPPER JAW. + +The =palato-pterygo-quadrate bar= is in a very different condition +from that of the dogfish, it is partially cartilaginous, partially +converted into cartilage bone, partially overlapped by membrane bone. +It is narrow in front but becomes much broader and deeper when +followed back. Its anterior end forms the =palatine= which bears +teeth, and in front is completely ossified, while behind the cartilage +is only sheathed by bone. + +Just behind the palatine the outer part of the cartilage is ossified, +forming two small bones, the =pterygoid= and =meso-pterygoid=, while +behind them is a larger, somewhat square bone, the =meta-pterygoid= +(fig. 11, 15). + +Below the meta-pterygoid is a tract of unossified cartilage, and then +comes the =quadrate= (fig. 11, 18). + +The lower angle of the quadrate bears a cartilaginous =condyle= with +which the mandible articulates. In front of the palatine the +cartilaginous snout is overlapped by three membrane bones, the +_jugal_, _maxilla_ and _premaxillae_. + +The _premaxillae_ (fig. 11, 20), the largest of these, overlaps the +maxilla behind; both bones bear teeth. The _jugal_ (fig. 11, 17) lies +above the maxilla and overlaps it in front. + +THE LOWER JAW. + +The =lower jaw= is a strong bar and is like the upper jaw, partly +cartilaginous, forming =Meckel's cartilage=, partly ossified, and +sheathed to a considerable extent in membrane bone. + +The outer side and posterior end is ossified, forming the large +=articular= (fig. 11, 21), but the condyle is cartilaginous and the +anterior part of the articular forms merely a splint on the outer side +of Meckel's cartilage, which extends beyond it for a considerable +distance. The angle of the jaw just below the condyle is formed by a +small _angular_ (fig. 11, 22), and the anterior two-thirds of the jaw +is sheathed in the large tooth-bearing _dentary_ (fig. 11, 23). + +THE HYOID ARCH. + +The =hyoid arch= has a number of ossifications in it and is closely +connected with the mandibular arch. + +The =hyomandibular= (fig. 11, 24) is a large bone which articulates +with a shallow groove lined by cartilage and formed partly in the +pterotic, partly in front of it. The hyomandibular is overlapped in +front by the meta-pterygoid, while below it tapers and is succeeded by +a small area of unossified cartilage followed by the forwardly +directed =symplectic= which fits into a groove in the quadrate. + +The unossified tract between the hyomandibular and symplectic is +continuous in front with a strong bar, which remains partly +cartilaginous and is partly converted into cartilage bone. The +proximal part is ossified, forming the =epi-hyal=, the middle part +forms the =cerato-hyal= (fig. 11, 27), in front of which is the small +=hypo-hyal=. The hyoid arches of the two sides are united by the large +tooth-bearing =glosso-hyal= (fig. 11, 29). Attached to the lower +surface of the hyoid arch are a series of twelve flat _branchiostegal +rays_ (fig. 11, 35). Each overlaps the one in front of it, the +posterior one being the largest. The branchiostegal rays of the two +sides are united in front by an unpaired membrane bone, the +_basi-branchiostegal_ (fig. 11, 36). + +=Opercular bones.= Behind the hyomandibular there is a large bony +plate, the =operculum=, formed of four large membrane bones. The +anterior of these, the _pre-opercular_ (fig. 11, 33), is crescentic in +shape, and with its upper end a small _supratemporal_ (fig. 11, 34) is +connected. + +Behind the upper part of the pre-opercular is the largest of the +opercular bones, the _opercular proper_. Its lower edge overlaps the +sub-opercular, and both opercular and sub-opercular are overlapped by +the _infra-opercular_ (fig. 11, 32) in front. The infra-opercular is +in its turn overlapped by the _pre-opercular_. + +BRANCHIAL ARCHES. + +There are five branchial arches, the first four of which bear gill +rays. Each of the first three consists of a shorter upper portion +directed obliquely backwards and outwards, and a longer lower portion +forming a right angle with the upper and directed obliquely forwards +and inwards. The greater part of each arch is ossified. + +The upper part of either of the first two consists of a short tapering +=pharyngo-branchial= directed inwards, and of a long =epi-branchial= +tipped with cartilage at both ends. The junction of the upper and +lower parts is formed by a cartilaginous hinge-joint between the +epi-branchial and cerato-branchial. The =cerato-branchial= is a long +bony rod separated by a short area of cartilage from the +=hypo-branchial=, which is succeeded by the =basibranchial= meeting +its fellow in the middle line. The =fourth arch= has a short +epi-branchial and no ossified pharyngo-branchial, while the fifth is +reduced to little more than the cerato-branchial, which bears a few +teeth on its inner edge. All the branchial arches have projecting from +their surfaces a number of little processes which act as strainers. +The first and fourth arches have one series of these, the second and +third have two. + + +THE SKULL OF THE CODFISH[37]. + +A full description having been already given of the Salmon's skull, +that of the Codfish will be described in a briefer manner. The skull +is very fully ossified, and the great number of plate-like bones +render it a very complicated structure. + +THE CRANIUM. + +At the posterior end of the dorsal surface is the large +=supra-occipital=, which is drawn out behind into the large blade-like +=occipital spine=. On each side of the supra-occipital are the small +irregular _parietals_, while in front of it the roof of the skull is +mainly formed by the very large unpaired _frontal_. + +A complicated series of bones are developed in connection with the +=auditory capsule=, which forms a large projecting mass united with +the side of the cranium and drawn out behind into a pair of strong +processes, the =epi-otic= and =parotic= processes. Both these +processes are connected behind with a large V-shaped bone, the +_post-temporal_ (fig. 13, 1), which will be described when dealing +with the pectoral girdle. The =epi-otic process= is formed by the +=epi-otic=, which is continuous in front with the parietal. The +=parotic process= is formed by two larger bones, a more dorsal one, +the =pterotic=, and a more ventral and internal one, the =opisthotic=, +which is continuous in front with the large =pro-otic=. Intervening +between the pterotic and frontal is another rather large bone, the +=sphenotic=, this articulates below with the pro-otic. The pterotic +and sphenotic together give rise to a large concave surface by which +the hyomandibular articulates with the cranium. Several of the cranial +nerves pass out through the bones of the auditory capsule. The ninth +leaves by a foramen near the posterior border of the opisthotic, the +fifth and seventh by a notch in the anterior border of the pro-otic. + +A number of bones are likewise developed in connection with the orbit +forming the =orbital ring=. Of these the most anterior, the +_lachrymal_, is much the largest, the others are five to seven in +number, the most ventral being the _suborbital_. The sclerotic coat of +the eye is cartilaginous. + +Two pairs of bones and one unpaired bone are developed in connection +with the =olfactory capsules=, of these, the _nasals_ are narrow bones +lying next the lachrymals, but nearer the middle line; they overlap +the second pair of bones, the irregular =lateral ethmoids=. These meet +one another in the middle line, and are overlapped behind by the +frontal. They articulate laterally with the lachrymal and palatine, +and ventrally with the parasphenoid. + +In a =posterior view= the foramen magnum and the four bones which +surround it and together form the occipital segment are well seen. On +the ventral side is the =basi-occipital=, terminated posteriorly by a +slightly concave surface which articulates with the centrum of the +first vertebra. The sides of the foramen magnum are formed by the +=exoccipitals=, a pair of very irregular bones, pierced by a pair of +prominent foramina for the exit of the tenth nerves. The exoccipitals +also bear a pair of surfaces for articulation with corresponding ones +on the neural arch of the first vertebra. The most dorsal of the four +bones is the supra-occipital. + +On the ventral surface of the cranium in front of the basi-occipital +is seen the _parasphenoid_, a very long narrow bone which underlies +the greater part of the cranium. Behind, it articulates dorsally with +the basi-occipital and dorsolaterally with the pro-otics and +opisthotics, in front it articulates dorsally with the lateral ethmoid +and ventrally with the vomer. At the sides of the parasphenoid are the +small =alisphenoids= articulating above with the postfrontals, in +front with the frontals, and behind with the pro-otics. + +The _vomer_ is an unpaired bone lying immediately in front of the +parasphenoid. In front it terminates with a thickened curved margin +bearing several rows of small teeth; behind it tapers out into a long +process which underlies the anterior part of the parasphenoid. +Immediately dorsal to the vomer is another median bone, the _median +ethmoid_; this is truncated in front and tapers out behind into a +process which fits into a groove on the ventral side of the frontal. + +BONES IN CONNECTION WITH THE UPPER JAW. + +These bear a close resemblance to those of the Salmon. The most +anterior bone is the _premaxillae_, a thick curved bone meeting its +fellow in the middle line. The point of junction of the two is drawn +out into a short process, and the oral surface is thickly covered with +small teeth. The dorsal ends of the premaxillae are seen in the fresh +skull to meet a large patch of cartilage. Behind the premaxillae is the +_maxilla_, a long rod-like toothless bone, somewhat expanded at the +upper end where it articulates with the premaxillae and vomer. + +Articulating in front with the anterior end of the maxilla and with +the =lateral ethmoid= is a very irregular bone, the =palatine= (fig. +12, 1); it articulates behind with two flat bones, the =pterygoid= and +=meso-pterygoid=. The pterygoid is united behind with two more bones, +the =quadrate= (fig. 12, 4) and =meta-pterygoid=. The =quadrate= is a +rather stout irregular bone, bearing on its lower surface a prominent +saddle-shaped articulating surface for the mandible. The palatine, +pterygoid and quadrate bones are the ossified representatives of the +palato-pterygo-quadrate bar of the Dogfish. + +[Illustration FIG. 12. MANDIBULAR AND HYOID ARCHES OF A COD (_Gadus +morrhua_) × 1/2 (Brit. Mus.). + + 1. palatine. + 2. meso-pterygoid. + 3. pterygoid. + 4. quadrate. + 5. symplectic. + 6. meta-pterygoid. + 7. hyomandibular. + 8. angular. + 9. articular. + 10. dentary. + 11. inter-hyal. + 12. epi-hyal. + 13. cerato-hyal. + 14. hypo-hyal. + 15. uro-hyal. + 16. branchiostegal rays.] + +The quadrate is united behind with the =symplectic= (fig. 12, 5), and +the meta-pterygoid with the symplectic and =hyomandibular=, both of +which bones will be described immediately in connection with the hyoid +arch. + + +THE LOWER JAW. + +The =lower jaw or mandible= like that of the Salmon is partly +cartilaginous, forming =Meckel's cartilage=, partly formed of +cartilage bone, partly of membrane bone. Meckel's cartilage is of +course not seen in the dried skull. + +The lower jaw includes one cartilage bone, the =articular= (fig. 12, +9), this is a large bone connected by a saddle-shaped surface with the +quadrate. Meckel's cartilage lies in a groove on its under surface, +and projects beyond it in front. The _angular_ is a small thick bone +united to the lower surface of the articular at its posterior end. The +_dentary_ (fig. 12, 10) is a large tooth-bearing bone meeting its +fellow in the middle line in front, while the articular fits into a +deep notch at its posterior end. + + +THE HYOID ARCH. + +The =hyomandibular= (fig. 12, 7) is a large irregular bone, +articulating by a prominent rounded head with the sphenotic and +pterotic. It is united in front with the meta-pterygoid and +symplectic, and sends off behind a strong process which articulates +with the opercular. The =symplectic= is a long somewhat triangular +bone drawn out in front into a process which fits into a groove on the +inner surface of the quadrate. The distal portion of the hyoid arch is +strongly developed and consists of first the =inter-hyal= (fig. 12, +11), a short bony rod, which articulates dorsally with a patch of +cartilage intervening between the posterior part of the hyomandibular +and the symplectic. Below it is united with the apex of the triangular +=epi-hyal=, a bone suturally connected with the large =cerato-hyal= +(fig. 12, 13) which unites distally with two small =hypo-hyals=. To +the cerato-hyal are attached a series of seven strong curved +cylindrical rods, the _branchiostegal rays_. The first of these is the +smallest and they increase in size up to the last. The four dorsal +ones are attached to the outer surface of the cerato-hyal, the three +ventral ones to its inner surface. Interposed between the hypo-hyals +of the two sides is an unpaired somewhat triangular plate, the +uro-hyal or _basi-branchiostegal_ (fig. 12, 15). + + +THE BRANCHIAL ARCHES. + +The =branchial arches= are five in number and consist of the following +parts on each side. The dorsal end is formed of the =supra-pharyngeal= +bone, a large irregular bone covered ventrally with teeth of a fair +size, and representing the fused =pharyngo-branchials= of the four +anterior arches. Its external surface is continuous with four small +=epi-branchials= which pass horizontally backwards and outwards. Their +distal ends meet four long =cerato-branchials= which are directed +forwards and inwards and form the principal part of the arches. + +Each of the first three cerato-branchials articulates ventrally with a +=hypo-branchial=, and the hypo-branchials of the two sides are united +in the middle line by an unpaired =basibranchial=. The third +hypo-branchial is much flattened. The fourth cerato-branchial is +united by cartilage with the posterior surface of the third +hypo-branchial, which it meets near the middle line. + +The fifth arch consists only of the cerato-branchial, a wide structure +covered with teeth and generally called the =inferior pharyngeal +bone=. + +The skeleton of the =operculum= consists of the same four bones as in +the Salmon, namely the _opercular_, the _infra-opercular_, the +_pre-opercular_ and the _sub-opercular_. Of these the anterior bone, +the _pre-opercular_, is the largest, while the _infra-opercular_ is +the smallest. The _opercular_ has a facet for articulation with the +hyomandibular. + + +2. THE APPENDICULAR SKELETON. + +THE PECTORAL GIRDLE. + +This is of a highly specialised type. Membrane bones are greatly +developed, and the cartilage bones, the =scapula= and =coracoid=, are +much reduced in size and importance. + +[Illustration FIG. 13. THE RIGHT HALF OF THE PECTORAL GIRDLE AND RIGHT +PECTORAL FIN OF A COD (_Gadus morrhua_) × 1/2 (Brit. Mus.). + + 1. post-temporal. + 2. supra-clavicle. + 3. clavicle. + 4. coracoid. + 5. scapula. + 6. post-clavicle. + 7. brachial ossicles. + 8. dermal fin-rays.] + +The largest bone in the shoulder girdle is the _clavicle_ (fig. 13, +3), which is irregularly crescent shaped, thick in front and tapering +off behind. To the outer side of its upper part is attached a thick +cylindrical bone, the _supra-clavicle_, which passes upwards and is +connected with a strong =V= shaped bone, the _post-temporal_. The apex +of the =V= meets the supra-clavicle, the inner limb articulates with +the epi-otic process, the outer with the parotic process. Projecting +downwards from the upper part of the clavicle is a long bony rod, +flattened proximally and cylindrical and pointed distally, this is +the _post-clavicle_ (fig. 13, 6). + +The =scapula= (fig. 13, 5) is a small irregular plate of bone attached +to the inner side of the middle of the _clavicle_. The =coracoid=[38] +is a larger plate of similar character, irregularly triangular in +shape, attached to the inner side of the clavicle immediately below +the scapula. The scapula and coracoid bear the pectoral fin. + +THE PECTORAL FINS. + +Each of these consists of four small irregular bones, the =brachial +ossicles= (fig. 13, 7), bearing a series of about nineteen dermal +_fin-rays_. The brachial ossicles represent the reduced and modified +radiale and basalia of cartilaginous fish such as the dogfish. The +fin-rays (fig. 13, 8) which form the whole external portion of the fin +are long slender rods having essentially the same character as those +of the unpaired fins. + +THE PELVIC GIRDLE. + +The =pelvic girdle= in the Cod as in other Teleosteans is entirely +absent, its place being taken by the enlarged basi-pterygia of the +fins. + +THE PELVIC FINS. + +These have a very anomalous position in the Cod, being attached to the +throat in front of the pectoral girdle. Each consists of a basal +portion, the =basi-pterygium=, and of a number of dermal rays. The +basi-pterygium consists of an expanded ventral portion which meets its +fellow below in the middle line, and to which the rays are attached, +and of an inwardly-directed dorsal portion which also meets its fellow +and is imbedded in the flesh. The rays are six in number and are long +slender structures similar to those of the other fins. + + +FOOTNOTES: + +[35] See T.J. Parker's _Zootomy_, London, 1884, p. 86. + +[36] See W.K. Parker and G.T. Bettany, _The Morphology of the Skull_, +London, 1877, chap. 3. + +[37] T.J. Parker, _Zootomy_, London, 1884, p. 91. + +[38] According to G. Swirski, _Schultergurtel des Hechtes_, Dorpat, +1880, the true coracoid is aborted, and the so-called coracoid of +Teleosteans is really the precoracoid. + + + + +CHAPTER VIII. + +GENERAL ACCOUNT OF THE SKELETON IN FISHES[39]. + + +EXOSKELETON. + +The most primitive type of exoskeleton is that found in Elasmobranchs +and formed of =placoid= scales; these are tooth-like structures +consisting of dentine and bone capped with enamel, and have been +already described (p. 4). In most Elasmobranchs they are small and +their distribution is fairly uniform, but in the Thornback skate, +_Raia clavata_, they have the form of larger, more scattered spines. +In adult Holocephali and in _Polyodon_ and _Torpedo_ there is no +exoskeleton, in young Holocephali, however, there are a few small +dorsal ossifications. + +The plates or scales of many Ganoids may have been formed by the +gradual fusion of elements similar to these placoid scales, and often +bear a number of little tooth-like processes. In _Lepidosteus_, +_Polypterus_, and many extinct species, these _ganoid_ scales, which +are rhomboidal in form and united to one another by a peg and socket +articulation, enclose the body in a complete armour. In _Trissolepis_ +part of the tail is covered by rhomboidal scales, while rounded scales +cover the trunk and remainder of the tail. _Acipenser_ and +_Scaphirhynchus_ have large dermal bony plates which are not +rhomboidal in shape and do not cover the whole body. In _Acipenser_ a +single row extends along the middle of the back and two along each +side. + +The majority of Teleosteans have thin flattened scales which differ +from those of Ganoids in being entirely mesodermal in origin, +containing no enamel. There are two principal types of Teleostean +scales, the cycloid and ctenoid. A =cycloid= scale is a flat thin +scale with concentric markings and an entire posterior margin. A +=ctenoid= scale differs in having its posterior margin pectinate. The +Dipnoi have overlapping cycloid scales. The rounded scales of _Amia_ +and of many fossil ganoids such as _Holoptychius_ are shaped like +cycloid scales, but differ from them in being more or less coated with +enamel. In Eels and some other Teleosteans the scales are completely +degenerate and have almost disappeared. Some Teleosteans, like _Diodon +hystrix_, have scales with triradiate roots from which arise long +sharp spines directed backwards. These scales, which resemble teeth, +contain no enamel; they become erect when the fish inflates its body +into a globular form. Many Siluroids have dermal armour in the form of +large bony plates which are confined to the anterior part of the body. +In _Ostracion_ the whole body is covered by hexagonal plates, closely +united together. + +The =fin-rays= are structures of dermal origin which entirely or +partially support the unpaired fins, and assist the bony or +cartilaginous endoskeleton in the support of the paired fins. + +In Elasmobranchs, Dipnoi, and Chondrosteous ganoids the skeletons of +the fins are, as a rule, about half of exoskeletal, half of +endoskeletal origin, the proximal and inner portion being +cartilaginous and endoskeletal, the distal and outer portion being +exoskeletal, and consisting of horny or of more or less calcified +fin-rays. In bony Ganoids and Teleosteans the endoskeletal parts are +greatly reduced and the fins come to consist mainly of the fin-rays, +which are ossified and frequently become flattened at their distal +ends. + +The fin-rays of the ventral part of the caudal fin are carried by the +haemal arches; those of the dorsal and anal fins and of the dorsal +part of the caudal fin generally by interspinous bones, which in adult +Teleosteans alternate with the neural and haemal spines. In Dipnoi +these interspinous bones articulate with the neural and haemal spines. +In many Siluroids the anterior rays of the dorsal and pectoral fins +are developed into large spines which often articulate with the +endoskeleton, or are sometimes fused with the dermal armour plates. +Similar spines may occur in Ganoids in front of both the dorsal and +anal fins. _Polypterus_ has a small spine or _fulcrum_ in front of +each segment of the dorsal fin. Such spines are often found +fossilised, and are known as _ichthyodorulites_. + +Similar spines are found in many Elasmobranchs, but they are simply +inserted in the flesh, not articulated to the endoskeleton. They also +differ from the spines of Teleosteans and Ganoids in the fact that +they are covered with enamel, and often have their edges serrated like +teeth. In the extinct Acanthodii they generally occur in front of all +the fins, paired and unpaired. + +In _Trygon_, the Sting-ray, the tail bears a serrated spine which is +used for purposes of offence and defence. Many ichthyodorulites may +have been spines of this nature fixed to the tail, rather than spines +situated in front of the fins. The spines, which are always found in +front of the dorsal fin in Holocephali, agree with those of +Elasmobranchs in containing enamel, and with those of Teleosteans in +being articulated to the endoskeleton. + +TEETH. + +The teeth of fish[40] are subject to a very large amount of variation, +perhaps to more variation than are those of any other class of +animals. Sometimes, as in adult Sturgeons, they are entirely absent, +sometimes they are found on all the bones of the mouth, and also on +the hyoid and branchial arches. The teeth are all originally developed +in the mucous membrane of the mouth, but they afterwards generally +become attached to firmer structures, especially to the jaws. In +Elasmobranchs, however, they are generally simply imbedded in the +tough fibrous integument of the mouth. Their attachment to the jaws +may take place in three different ways. + +[Illustration FIG. 14. DIAGRAM OF A SECTION THROUGH THE JAW OF A SHARK +(_Odontaspis americanus_) showing the succession of teeth (Brit. Mus. +from specimen and diagram). + + 1. teeth in use. + 2. teeth in reserve. + 3. skin. + 4. cartilage of the jaw. + 5. encrusting calcification of cartilage. + 6. connective tissue. + 7. mucous membrane of the mouth.] + +(1) By an elastic hinge-joint, as in the Angler (_Lophius_), and the +Pike (_Esox lucius_). In the Angler the tooth is held by a fibrous +band attaching its posterior end to the subjacent bone, in the Pike by +uncalcified elastic rods in the pulp cavity. + +(2) By ankylosis, i.e. by the complete union of the calcified tooth +substance with the subjacent bone. This is the commonest method among +fish. + +(3) By implantation in sockets. This method is not very common among +fish. The teeth are sometimes, as in _Lepidosteus_, ankylosed to the +base of the socket. In this genus there is along each ramus of the +mandible a median row of large teeth placed in perfect sockets, and +two irregular lateral rows of small teeth ankylosed to the jaw. + +Dentine, enamel and cement are all represented in the teeth of fishes, +but the enamel is generally very thin, and cement is but rarely +developed. Dentine forms the main bulk of the teeth; it is sometimes +of the normal type, but generally differs from that in higher +vertebrates in being vascular, and is known as _vasodentine_. A third +type occurs, known as _osteodentine_; it is traversed by canals +occupied by marrow, and is closely allied to bone. + +[Illustration FIG. 15. PART OF THE LOWER JAW OF A SHARK (_Galeus_) +(from OWEN after ANDRÉ). + +1. teeth in use. + +2. reserve teeth folded back. + +3. part of the caudal spine of a Sting-ray (_Trygon_) which has +pierced the jaw and affected the growth of the teeth.] + +The teeth are generally continually renewed throughout life, but +sometimes one set persists. + +The teeth of Selachii are fundamentally identical with placoid +scales. They are developed from a layer of dental germs which occurs +all over the surface of the skin, except in the region of the lips. At +this point the layer of tooth-producing germs extends back into the +mouth, being projected by a fold of the mucous membrane (fig. 14, 7). +Here new teeth are successively formed, and as they grow each is +gradually brought into a position to take the place of its predecessor +by the shifting outwards of the gum over the jaw. Owing to this +arrangement sharks have practically an unlimited supply of teeth +(figs. 14 and 15). + +Two principal types of teeth are found in ELASMOBRANCHS. In Sharks and +Dogfish, on the one hand, the teeth are very numerous, simple, and +sharp-pointed, and are with or without serrations and lateral cusps. +Many Rays and fossil Elasmobranchs, on the other hand, have broad +flattened teeth adapted for crushing shells. Intermediate conditions +occur between these two extremes. Thus in _Cestracion_ and many +extinct sharks, such as _Acrodus_, while the median teeth are sharp, +the lateral teeth are more or less flattened and adapted for crushing. +In various species belonging to the genus _Raia_ the teeth of the male +are sharp, while those of the female are blunt. A very specialised +dentition is met with in the Eagle-rays (Myliobatidae), in which the +jaws are armed with flattened angular tooth-plates, arranged in seven +rows, forming a compact pavement; the plates of the middle row are +very wide and rectangular, those of the other rows are much smaller +and hexagonal. Lastly, in _Cochliodus_ the individual crushing teeth +are fused, forming two pairs of spirally-coiled dental plates on each +side of each jaw. _Pristis_, the Saw-fish, has a long flat +cartilaginous snout, bearing a double row of persistently-growing +teeth planted in sockets along its sides. Each tooth consists of a +number of parallel dentinal columns, united at the base, but elsewhere +distinct. + +In the HOLOCEPHALI--_Chimaera_, _Hariotta_ and _Callorhynchus_--only +three pairs of teeth or dental plates occur, two pairs in the upper +jaw, one in the lower. These structures persist throughout life and +grow continuously. The upper tooth structures are attached +respectively to the ethmoid or vomerine region of the skull, and to +the palato-pterygoids. The vomerine teeth are small, while those +attached to the mandible and the palato-pterygoid region are large and +bear several roughened ridges adapted for grinding food. The teeth of +the two opposite sides of the jaw meet in a median symphysis. The +teeth of _Chimaera_ are more adapted for cutting, those of +_Callorhynchus_ for crushing. Many extinct forms are known, some of +whose teeth are intermediate in structure between those of _Chimaera_ +and _Callorhynchus_. + +The teeth of GANOIDS are also extremely variable. Among living forms, +the Holostei are more richly provided with teeth than are any other +fishes, as they may occur on the premaxillae, maxillae, palatines, +pterygoids, parasphenoid, vomers, dentaries, and splenials. Among the +Chondrostei, on the other hand, the adult Acipenseridae are toothless; +small teeth however occur in the larval sturgeon, and in _Polyodon_ +many small teeth are found attached merely to the mucous membrane of +the jaws. Many fossil Ganoids have numerous flattened or knob-like +teeth, borne on the maxillae, palatines, vomers and dentaries. Others +have a distinctly heterodont dentition. Thus in _Lepidotus_ the +premaxillae bear chisel-like teeth, while knob-like teeth occur on the +maxillae, palatines and vomers. In _Rhizodus_ all the teeth are +pointed, but while the majority are small a few very large ones are +interspersed. + +In TELEOSTEANS, too, the teeth are eminently variable both in form and +mode of arrangement. They may be simple and isolated, or compound, and +may be borne on almost any of the bones bounding the mouth cavity, and +also as in the Pike, on the hyoid and branchial arches. The splenial +however never bears teeth and the pterygoid and parasphenoid only +rarely, thus differing from the arrangement in the Holostei. + +The isolated teeth are generally conical in form and are ankylosed to +the bone that bears them. Such teeth are, with a few exceptions such +as _Balistes_, not imbedded in sockets nor replaced vertically. + +In some fish beak-like structures occur, formed partly of teeth, +partly of the underlying jaw bones. These beaks are of two kinds: (1) +In _Scarus_, the parrot fish, the premaxillae and dentaries bear +numerous small, separately developed teeth, which are closely packed +together and attached by their proximal ends to the bone, while their +distal ends form a mosaic. Not only the teeth but the jaws which bear +them are gradually worn away at the margins, while both grow +continuously along their attached edge. (2) In Gymnodonts, e.g. +_Diodon_, the beaks are formed by the coalescence of broad calcified +horizontal plates, which when young are free and separated from one +another by a considerable interval. + +In some Teleosteans the differentiation of the teeth into biting teeth +and crushing teeth is as complete as in _Lepidosteus_. Thus in the +Wrasse (_Labrus_), the jaws bear conical slightly recurved teeth +arranged in one or two rows, with some of the anterior ones much +larger than the rest. The bones of the palate are toothless, while +both upper and lower pharyngeal bones are paved with knob-like +crushing teeth; such pharyngeal teeth occur also in the Carp but are +attached only to the lower pharyngeal bone, the jaw bones proper being +toothless. + +In DIPNOI the arrangement of the teeth is very similar to that in +Holocephali. The mandible bears a single pair of grinding teeth +attached to the splenials, and a corresponding pair occur on the +palato-pterygoids. In front of these there are a pair of small conical +vomerine teeth loosely attached to the ethmoid cartilage. The +palato-pterygoid teeth of _Ceratodus_ are roughly semicircular in +shape with a smooth convex inner border, and an outer border bearing a +number of strongly marked ridges. The teeth of the extinct Dipteridae +resemble those of _Ceratodus_ but are more complicated. + + +ENDOSKELETON. + +SPINAL COLUMN[41]. + +The spinal column of fishes is divisible into only two regions, a +caudal region in which the haemal arches or ribs meet one another +ventrally, and a precaudal region in which they do not meet. + +The various modifications of the spinal column in fishes can be best +understood by comparing them with the arrangement in the simplest type +known, namely _Amphioxus_. In _Amphioxus_ the notochord is immediately +surrounded by a structureless cuticular layer, the _chordal sheath_. +Outside this is the _skeletogenous layer_, which in addition to +surrounding the notochord and chordal sheath embraces the nerve cord +dorsally, and laterally sends out septa forming the _myomeres_. + +The CARTILAGINOUS GANOIDS[42] _Acipenser_, _Polyodon_ and +_Scaphirhynchus_ are the simplest fishes as regards their spinal +column. The notochord remains permanently unconstricted and is +enclosed in a chordal sheath, external to which is the skeletogenous +layer. In this layer the development of cartilaginous elements has +taken place. In connection with each _neuromere_, or segment as +determined by the points of exit of the spinal nerves, there are +developed two pairs of ventral cartilages, the ventral arches +(basiventralia) and intercalary pieces (interventralia); and at least +two pairs of dorsal pieces, the neural arches (basidorsalia) and +intercalary pieces (interdorsalia). The lateral parts of the +skeletogenous layer do not become converted into cartilage, so there +are no traces of vertebral centra. The ventral or haemal arches meet +one another ventrally and send out processes to protect the ventral +vessels. The neural arches do not meet, but are united by a +longitudinal elastic band. + +In Cartilaginous ganoids the only indications of metameric +segmentation are found in the neural and haemal arches. The case is +somewhat similar with the Holocephali and Dipnoi. + +In the HOLOCEPHALI the notochord grows persistently throughout life, +and is of uniform diameter throughout the whole body except in the +cervical region and in the gradually tapering tail. The chordal sheath +is very thick and includes a well-marked zone of calcification which +separates an outer zone of hyaline cartilage from an inner zone. There +are also a number of cartilaginous pieces derived from the +skeletogenous layer which are arranged in two series, a dorsal series +forming the neural arches and a ventral series forming the haemal +arches. These do not, except in the cervical region, meet one another +laterally round the notochord and form centra. To each neuromere there +occur a pair of basidorsals, a pair of interdorsals, and one or two +supradorsals. In the tail the arrangement is irregular. + +In the DIPNOI as in the Holocephali the notochord grows persistently +and uniformly, and the chordal sheath is thick and cartilaginous +though there are no metamerically arranged centra. The neural and +haemal arches and spines are cartilaginous and interbasalia +(intercalary pieces) are present. The basidorsalia and basiventralia +do not in _Ceratodus_ meet round the notochord and enclose it except +in the anterior part of the cervical and posterior part of the caudal +region. + +In ELASMOBRANCHII the chordal sheath is weak and the skeletogenous +layer strong. Biconcave cartilaginous vertebrae are developed, and as +is the case in most fishes, constrict the notochord _vertebrally_. + +Two distinct types of vertebral column can be distinguished in +Elasmobranchs[43]: + +1. In many extinct forms and in the living Notidanidae, _Cestracion_, +and _Squatina_, the dorsal and ventral arches do not meet one another +laterally round the centrum, and consequently readily come away from +it. + +2. In most living Elasmobranchs the arches meet laterally round the +centrum. + +The vertebrae are never ossified but endochondral calcification nearly +always takes place, though it very rarely reaches the outer surface of +the vertebrae. Elasmobranchs are sometimes subdivided into three +groups according to the method in which this calcification takes +place: + +1. =Cyclospondyli= (_Scymnus_, _Acanthias_), in which the calcified +matter is deposited as one ring in each vertebra. + +2. =Tectospondyli= (_Squatina_, _Raia_, _Trygon_), in which there are +several concentric rings of calcification. + +3. =Asterospondyli= (Notidanidae, _Scyllium_, _Cestracion_), in which +the calcified material instead of forming one simple ring, extends out +in a more or less star-shaped manner. + +In _Heptanchus_ the length of the vertebral centra in the middle of +the trunk is double that in the anterior and posterior portions, and +as the length of the arches does not vary, the long centra carry more +of them than do the short centra. + +In many Rays the skull articulates with the vertebral column by +distinct occipital condyles. + +In BONY GANOIDS the skeletogenous layer becomes calcified +ectochondrally in such a way that the notochord is pinched in at +intervals, and distinct vertebrae are produced. Ossification of the +calcified cartilage rapidly follows. In _Amia_ the vertebrae are +biconcave, in _Lepidosteus_ they are opisthocoelous, cup and ball +joints being developed between the vertebrae in a manner unique among +fishes. The notochord entirely disappears in the adult _Lepidosteus_, +but at one stage in larval life it is expanded vertebrally and +constricted intervertebrally in the manner usual in the higher +vertebrata, but unknown elsewhere among fishes. + +The tail of _Amia_ is remarkable from the fact that as a rule to each +neuromere, as determined by the exit of the spinal nerves, there are +two centra, a posterior one which bears nothing, and an anterior one +which bears the neural and haemal arches, these being throughout the +vertebral column connected with the centra by cartilaginous discs. + +In most TELEOSTEANS but not in the Plectognathi the neural arches are +continuous with the centra, which are nearly always deeply biconcave. + +In some cases many of the anterior vertebrae are ankylosed together +and to the skull. The vertebrae often articulate with one another by +means of obliquely placed flattened surfaces, the zygapophyses. The +centrum in early stages of development is partially cartilaginous, but +the neural arches and spines in the trunk at any rate, pass directly +from the membranous to the osseous condition. + + +FINS. + +The most primitive fins are undoubtedly the unpaired ones, which +probably originally arose as ridges or folds of skin along the +mid-dorsal line of the body, and passed thence round the posterior end +on to the ventral surface, partially corresponding in position and +function to the keel of a ship. + +In long 'fish' which pass through the water with an undulating motion +such simple continuous fins may be the only ones found, as in +_Myxine_. To support these median fins skeletal structures came to be +developed; these show two very distinct forms, viz. cartilaginous +endoskeletal pieces, the _radiale_, and horny exoskeletal fibres, the +_fin-rays_. Mechanical reasons caused the fin to become concentrated +at certain points and reduced at intervening regions. Thus a terminal +caudal fin arose and became the chief organ of propulsion, and the +dorsal and ventral fins became specialised to act as balancing organs. + +In some of the earlier Elasmobranchs, the Pleuracanthidae, the +endoskeletal cartilaginous radiale are directly continuous with +outgrowths from the dorsal and ventral arches of the vertebrae, and +form the main part of the fin. In later types of Elasmobranchs the +horny exoskeletal fin-rays have comparatively greater prominence. In +bony fish, as has been already stated, the horny fibres are replaced +by bony rays of dermal origin, and at the same time complete reduction +and disappearance of the cartilaginous radiale takes place. + +THE CAUDAL FIN. + +The caudal region of the spinal column in fishes is of special +importance. It is distinctly marked off from the rest of the spinal +column by the fact that the ventral or haemal arches meet one another +and are commonly prolonged into spines, while in the trunk region they +do not meet but commonly diverge from one another. + +In some fish the terminal part of the caudal region of the spinal +column retains the same direction as the rest of the spinal column. +The blade of the caudal fin is then divided into two nearly equal +portions, and is said to be =diphycercal=. This condition is generally +regarded as the most primitive one; it occurs in the Ichthyotomi, +Holocephali, all living Dipnoi, _Polypterus_ and some extinct +Crossopterygii, and a few Selachii and Teleostei. It occurs also in +deep-sea fish belonging to almost every group, and under these +conditions obviously cannot be regarded as primitive, but must be +looked on as a feature induced by the peculiar conditions of life. + +In the great majority of fish the terminal part of the caudal region +of the spinal column is bent dorsalwards, and the part of the blade of +the caudal fin which arises on the dorsal surface is much smaller than +is that arising on the ventral surface. Such a fin is said to be +=heterocercal=. + +Strictly speaking all fish whose tails are not diphycercal have +heterocercal tails, but the term is commonly applied to two-bladed +tails in which the spinal column forms a definite axis running through +the dorsal blade, while the ventral blade is enlarged and generally +forms the functional part of the tail. Such heterocercal tails are +found in nearly all Elasmobranchii, together with the living +cartilaginous Ganoidei, and many extinct forms belonging to the same +order; _Lepidosteus_, _Amia_, and the Dipteridae among Dipnoi, have +tails which, though obviously heterocercal, are not two-bladed. + +The vast majority of the Teleostei and some extinct Ganoidei have +heterocercal tails of the modified type to which the term =homocercal= +is applied. The hypural bones which support the lower half of the tail +fin become much enlarged, and frequently unite to form a wedge-shaped +bone which becomes ankylosed to the last ossified vertebral centrum. +The fin-rays then become arranged in such a way as to produce a +secondary appearance of symmetry. Some homocercal fish such as the +Perch have the end of the notochord protected by a calcified or +completely ossified sheath, the =urostyle=, to which several neural +and haemal arches may be attached, and which becomes united with the +centrum of the last vertebra; in others such as the Salmon the end of +the notochord is protected only by laterally placed bony plates. + +THE SKULL. + +It is often impossible to draw a hard and fast line between the +cranium and the vertebral column. This is the case for instance in +_Acipenser_ (fig. 18, 16) among Chondrostei, in _Amia_ among Holostei, +and in _Ceratodus_ and _Protopterus_ among Dipnoi. The occipital +region of the skull in _Amia_ is clearly formed of three cervical +vertebrae whose centra have become absorbed into the cranium, while +the neural arches and spines are still distinguishable. + +The simplest type of cranium is that found in ELASMOBRANCHS: it +consists of a simple cartilaginous box, which is generally immovably +fixed to the vertebral column, though in some forms, like _Scymnus_ +and _Galeus_, a joint is indicated, and in others, such as the +Rays, one is fairly well developed. The cranium in Elasmobranchs +is never bony, though the cartilage is sometimes calcified. It is +drawn out laterally into an antorbital process in front of the eye, +and a postorbital process behind it. The nasal capsules are always +cartilaginous, and the eye, as a general rule, has a cartilaginous +sclerotic investment. The cranium is often prolonged in front into +a rostrum which is enormously developed in _Pristis_ and some Rays. +The cartilaginous roof of the cranium is rendered incomplete by the +presence of a large hole, the anterior fontanelle. + +[Illustration FIG. 16. A. SKULL OF _Notidanus_ × 1/2 (Brit. Mus.). B. +SKULL OF _Cestracion_ × 1/3 (after GEGENBAUR). In neither case are the +branchial arches shown. + + 1. rostrum. + 2. olfactory capsule. + 3. ethmo-palatine process. + 4. palatine portion of palato-pterygo-quadrate bar. + 5. quadrate portion of bar. + 6. Meckel's cartilage. + 7. teeth. + 8. labial cartilage. + 9. hyomandibular. + 10. postorbital process. + II. optic foramen.] + +Two pairs of labial cartilages (fig. 16, B, 8) are often present. They +lie imbedded in the cheeks outside the anterior region of the jaws, +and are specially large in _Squatina_. + +As regards the visceral arches[44] the simplest and most primitive +condition of the jaws is that of the Notidanidae, in which the +mandibular and hyoid arches are entirely separate. In these primitive +fishes the palato-pterygo-quadrate bar articulates with the +postorbital process (fig. 16, 10), while further forwards it is united +to the cranium by the ethmo-palatine ligament. The hyoid arch is small +and is broadly overlapped by the mandibular arch. The term +=autostylic= is used to describe this condition of the suspensorium. +From this condition we pass in the one direction to that of +_Cestracion_ (fig. 16, B), in which the whole of the palato-pterygo +quadrate bar has become bound to the cranium, and in the other to that +of _Scyllium_. In _Scyllium_ (fig. 6), while the ethmo-palatine +ligament is retained, the postorbital articulation of the palato +pterygo-quadrate has been given up, so that the palato-pterygo +quadrate comes to abut on the hyomandibular and is attached to it by +ligaments. The pre-spiracular ligament (fig. 16, 20) running from the +auditory capsule also assists in supporting the jaws. + +Lastly we come to the purely =hyostylic= condition met with in Rays, +in which the mandibular arch is entirely supported by the +hyomandibular. In some Rays the hyoid is attached to the posterior +face of the hyomandibular near its proximal end, and may even come to +articulate with the cranium. + +The =visceral arches of Elasmobranchs= may be summarised as follows:-- + +1. The =mandibular arch=, consisting of a much reduced dorsal portion, +the pre-spiracular ligament, and a greatly developed ventral portion +from which both upper and lower jaws are derived. The mandible +(Meckel's cartilage) is the original lower member of the mandibular +arch, and from it arises an outgrowth which forms the upper jaw or +palato-pterygo-quadrate bar. In _Scymnus_ this bears a few +branchiostegal rays. + +2. The =hyoid arch=, which consists of the hyomandibular and the +hyoid, and bears branchiostegal rays on its posterior face. + +[Illustration FIG. 17. DORSAL VIEW OF THE BRANCHIAL ARCHES OF +_Heptanchus_. (From GEGENBAUR). + + 1. basi-hyal. + 2. cerato-hyal. + 3. second hypo-branchial. + 4. first cerato-branchial. + 5. first epi-branchial. + 6. first pharyngo-branchial. + 7. pharyngo-branchial, common to the sixth and seventh arches. + 8. basibranchial of second arch. + 9. basibranchial, common to the sixth and seventh arches.] + +3. The =branchial arches=, generally five in number, all of which +except the last bear branchiostegal rays. In the Notidanidae the +number of branchial arches is increased beyond the normal series, +thus in _Hexanchus_ there are six, and in _Heptanchus_ seven. There +are six also in _Chlamydoselache_ and _Protopterus_. + +4. The so-called external branchial arches which are cartilaginous +rods attached to all the visceral arches. They are especially large in +_Cestracion_. + +The skull in HOLOCEPHALI is entirely cartilaginous. The +palato-pterygo-quadrate bar is fixed to the cranium, and to it the +mandible articulates. There is a well-marked joint between the skull +and the spinal column. + +In living Cartilaginous GANOIDS the primitive cartilaginous cranium is +very massive, and is greatly prolonged anteriorly, while posteriorly +it merges into the spinal column. Although it is mainly cartilaginous +a number of ossifications take place in the skull, and membrane bones +are now found definitely developed, especially in connection with the +roof of the cranium. In _Acipenser_ (fig. 18) the ossifications in the +cartilage include the pro-otic, which is pierced by the foramen for +the fifth nerve, the alisphenoid, orbitosphenoid, ectethmoid, +palatine, pterygoid, meso-pterygoid, hyomandibular (fig. 18, 11), +cerato-hyal, all the cerato-branchials, and the first two +epi-branchials. Most of these structures are, however, partly +cartilaginous, though they include an ossified area. The membrane +bones too of _Acipenser_ are very well developed, they include a bone +occupying the position of the supra-occipital, and form a complete +dorsal cephalic shield. Resting on the ventral surface are a vomer and +a very large parasphenoid (fig. 18, 3). There is a bony operculum +attached to the hyomandibular, and membrane bones representing +respectively the maxilla and dentary are attached to the jaws. The +suspensorium is most markedly hyostylic. The palato-pterygo-quadrate +bar has a very curious shape and is quite separate from the cranium. +It is connected to the hyomandibular by a thick symplectic ligament +containing a small bone homologous with the symplectic of +Teleosteans. + +_Polyodon_ differs much from _Acipenser_, the membrane bones not being +so well developed though they cover the great cartilaginous snout. + +[Illustration FIG. 18. LATERAL VIEW OF THE SKULL OF A STURGEON +(_Acipenser sturio_). Nearly all the membrane bones have been removed +(Brit. Mus.). + + 1. nasal cavity. + 2. orbit. + 3. parasphenoid. + 4. vomer. + 5. pterygoid. + 6. maxilla. (The dotted line running from 6 passes into the mouth + cavity.) + 7. dentary. + 8. symplectic. + 10. palatine. + 11. hyomandibular. + 12. pharyngo-branchial. + 13. epi-branchial. + 14. cerato-branchial. + 15. hypo-branchial. + 16. coalesced anterior vertebrae. + 17. inter-hyal. + 18. cerato-hyal. + 19. rib.] + +The skull in _Polypterus_ (Crossopterygii) shows a great advance +towards the condition met with in Teleostei. The cranium remains to a +great extent unossified, and large dorsal and ventral fontanelles +pierce its walls. It is covered by a great development of membrane +bones, paired nasals, frontals, parietals, supra- and post-temporals, +and dermo-supra-occipitals among others being present. The +palato-pterygo-quadrate bar is fused to the cranium, and in connection +with it the following paired membrane bones appear, palatine, ecto-, +meso- and meta-pterygoid, and further forwards jugal, vomer, maxilla +and premaxillae. The membrane bones developed in connection with each +ramus of the mandible are the dentary, angular, and splenial, in +addition to the cartilage bone the articular. Several large opercular +bones occur. There are also a pair of large jugular or gular plates, +and several large opercular bones. + +In Bony Ganoids both cartilage bone and membrane bone is well +developed. The pro-otics and exoccipitals are well ossified, but the +supra-occipital and pterotics are not. Lateral ethmoids are developed, +and there are ossifications in the sphenoidal region which vary in +different forms. The place of the cartilaginous palato-pterygo +quadrate is taken by a series of bones, the quadrate behind and the +palatine, ecto-, meso- and meta-pterygoids in front. In _Lepidosteus_, +however, the palatine and pterygoid are membrane bones, as they are in +_Polypterus_ and the Frog. Paired maxillae, premaxillae, vomers and a +parasphenoid occur forming the upper jaw and roof of the mouth, and a +series of membrane bones are found investing the mandible and forming +the operculum. + +In _Amia_[45] membrane bones are as freely developed as they are in +Teleosteans; they include on each side a squamosal, four opercular +bones, a lachrymal, a pre-orbital, one or two suborbitals, two large +postorbitals and a supratemporal; while investing the mandible, +besides the dentary, splenial, angular, and supra-angular, there is an +unpaired jugular. The articular too is double and a mento-meckelian +occurs. In _Amia_ teeth are borne on the premaxillae, maxillae, vomers, +palatines and pterygoids. + +Bony Ganoids are the lowest animals in which squamosal bones are +found, and they do not occur in Teleosteans. + +The suspensorium in bony Ganoids, as in the Chondrostei, is hyostylic, +and there are two ossifications in the hyomandibular cartilage, viz. +the hyomandibular, and the symplectic. + +The skull of TELEOSTEI is very similar to those of _Lepidosteus_ and +_Amia_. Although the bony skull is greatly developed and very +complicated, much of the original cartilaginous cranium often +persists. Membrane bones are specially developed on the roof of the +skull where they include the parietal, frontal, and nasal bones. The +same bones are developed in connection with the upper jaw and roof of +the mouth as in bony Ganoids, but only two membrane bones occur in the +lower jaw, viz. the angular and dentary. A number of large +ossifications take place in the cartilage of the auditory capsules. In +some forms parts of the last pair of branchial arches are broadened +out and form the pharyngeal bones which bear teeth. The opercular +bones and those of the upper and lower jaws are quite comparable to +those of bony Ganoids. + +A full account of the Teleostean skull has been given in the case of +the Salmon (pp. 87-96) and the Cod (pp. 96-101). + +In DIPNOI the skull is mainly cartilaginous, but both cartilage- and +membrane-bone occur also. Cartilage-bone is found in the ossified +exoccipitals, while of membrane-bones _Protopterus_ has among unpaired +bones a fronto-parietal, a median ethmoid, and a parasphenoid, and +among paired bones nasals and large supra-orbitals. The skull of +_Ceratodus_ (fig. 19) has an almost complete roof of membrane bones, +including some whose homology is doubtful. The ethmo-vomerine region +is always cartilaginous, but bears small teeth. The palato-pterygo +quadrate bar is ossified and firmly united to the cranium, and the +mandible articulates directly with it (autostylic). Membrane bones are +freely developed in connection with the mandible, dentary, splenial, +and angular bones being all present. There are two opercular bones. + +In the extinct Dipteridae the cranium is very completely covered with +plates of dermal bone, and the skeleton in general is more ossified +than is the case in recent Dipnoi. + +Six pairs of branchial arches occur in _Protopterus_; _Ceratodus_ and +_Lepidosiren_ have five, like most other fish. The branchial arches +bear gill rakers. + +[Illustration FIG. 19. DORSAL (TO THE LEFT) AND VENTRAL (TO THE RIGHT) +VIEWS OF THE CRANIUM OF _Ceratodus miolepis_ (after GÜNTHER). + + 1. cartilaginous part of the quadrate with which the mandible + articulates. + 2. scleroparietal. + 3. frontal. + 4. ethmoid. + 5. nares. + 6. orbit. + 7. pre-opercular (squamosal). + 8. second rib. + 9. first rib. + 10. vomerine tooth. + 11. palato-pterygoid tooth. + 12. palato-pterygoid. + 13. parasphenoid. + 14. interopercular.] + +RIBS. + +As has been already mentioned (p. 24), although ribs commonly appear +to be the cut-off ends of the transverse processes, they are really +elements derived from the ventral or haemal arch. + +In Elasmobranchii and other cartilaginous fish they have the form of +small cartilaginous structures imperfectly separated from the +diverging halves of the ventral arch, and are often absent. + +In Teleostei and bony Ganoids they often have different attachments in +different parts of the body. In the tail region they are not +differentiated from the two halves of the ventral arch, which meet in +the middle line, and are prolonged into a haemal spine. In the +posterior trunk region they sometimes form distinct processes +diverging from the two halves of the ventral arch; while further +forward they may shift their attachment so as to arise from the dorsal +side of the two halves of the ventral arch and at some distance from +their ends, which now diverge as ventri-lateral processes. + +APPENDICULAR SKELETON. + +PECTORAL GIRDLE. + +The simplest type of pectoral girdle is found in Elasmobranchs. It is +entirely cartilaginous and consists of a curved ventrally-placed rod, +ending dorsally in two horn-like scapular processes which are +sometimes attached to the cranium or vertebral column. In Rays the +shoulder girdle is very large, and has a distinct suprascapular +portion forming a broad plate attached to the neural spines of the +vertebrae. There is often a cup-like glenoid cavity for the +articulation of the limb; this cavity is specially large in Rays and +is much pierced by holes. In Dipnoi the cartilaginous girdle still +occurs, but on it there is a deposit of membrane bone forming the +clavicle, infraclavicle, and supra-clavicle. These bones, which with +the exception of the clavicle, are unknown in higher vertebrates, are +better developed in Ganoids, and best of all in Teleosteans. They are +connected by the supratemporal with the epi-otic and opisthotic +regions of the cranium. Owing to this development of dermal bone, the +original cartilaginous arch becomes much reduced, but ossifications +representing the scapula and coracoid occur in bony Ganoids and +Teleosteans. + +PELVIC GIRDLE. + +In Elasmobranchs the pelvic girdle consists of a short ventral rod of +cartilage representing the ischium and pubis, which does not send up +dorsal iliac processes. In _Chimaera_ the pelvic girdle has a +flattened pointed iliac portion, and ventrally an unpaired movable +cartilaginous plate which bears hooks and is supposed to be copulatory +in function. Claspers of the usual type are present as well. The +Dipnoi have a primitive kind of pelvis in the form of a cartilaginous +plate lying in the mid-ventral line and drawn out into three horns +anteriorly. In Ganoids the pelvis has almost entirely disappeared, +though small cartilaginous vestiges of it remain in _Polypterus_. In +Teleosteans even these vestiges are gone, and in these fish and +Ganoids the place of the pelvis is taken by the enlarged basi-pterygia +(meta-pterygia) of the fins. + +PAIRED FINS[46]. + +As regards the origin of the limbs or paired fins of fishes there are +two principal views. One view, that of Gegenbaur, considers that limbs +and their girdles are derived from visceral arches which have migrated +backwards. The other view, which probably now has the greater number +of supporters, considers that the paired fins of fishes are of +essentially the same nature as the median fins. + +According to Gegenbaur's view[47] the =archipterygium= of _Ceratodus_ +(fig. 20) represents the lowest type of fin; it consists of a central +cartilaginous axis bearing a large number of radiale. The dorsal or +pre-axial radiale are more numerous than the ventral or postaxial, and +at the margin of the fin[48] the cartilaginous endoskeletal radiale +are replaced by horny exoskeletal fin-rays. + +[Illustration FIG. 20. LATERAL VIEW OF THE SKELETON OF _Ceratodus +miolepis_ (after GÜNTHER). + + 1. ethmoid. + 2. scleroparietal. + 3. frontal. + 4. cartilaginous posterior part of cranium. + 5. pre-opercular (squamosal). + 6. opercular. + 7. suborbital. + 8. orbit. + 9. pectoral girdle. + 10. proximal cartilage of pectoral fin. + 11. pectoral fin. + 12. pelvic girdle. + 13. pelvic fin. + 14. spinal column. + 15. caudal fin (diphycercal).] + +It is impossible here to give a full discussion of the rival views, +but some of the points which support Gegenbaur's view may be +mentioned. The fact that migration of visceral arches has to be +assumed is no difficulty, as it is obvious that migration in the +opposite direction has taken place in many Teleosteans such as the +Cod, whose pelvic fins are attached to the throat in front of the +pectorals. If migration did take place, the pelvic fins being older +than the pectoral should be the more modified, and this is the case. +Again, if the pectoral girdle is a modified branchial arch, it must at +some period have carried a gill, and in _Protopterus_ it does bear a +vestigial gill. + +According to the view more prevalent at the present time, the paired +fins have been derived from two continuous folds of skin and their +skeletal supports running forward from the anal region along the sides +of the body, their character being similar to the fold that gave rise +to the median fins. In support of this view it may be argued that the +paired and unpaired fins are often identical in structure, and that +some Elasmobranch embryos do show a ridge running between the pectoral +and pelvic fins. Then from this continuous fold two pairs of smaller +folds may have been specialised off, and in each a number of +cartilaginous radiale may have been developed. The fin of +_Cladoselache_ from the Carboniferous of Ohio apparently illustrates +this condition. It consists of certain basal pieces which do not +project beyond the body wall and bear a number of unsegmented +cartilaginous radiale, which show crowding together and are sometimes +bifurcated distally; they extend throughout the whole fin from the +body wall to the margin. From this fin the archipterygium might be +easily derived by the enlargement of one of the middle radiale and the +segmentation and partial fusion of them all. + +Whether the archipterygium be a primitive or secondary type of fin, +when it is once reached it is easy to derive all the other types from +it. The fins of the other living Dipnoi,--_Protopterus_ and +_Lepidosiren_--are simply archipterygia from which the radiale have +almost or completely disappeared, leaving only the segmented axes. +Archipterygia too are found in the pectoral fins of the Ichthyotomi, +but the postaxial radiale are much reduced. + +The =ichthyopterygium=, or type of fin, characteristic of many modern +Elasmobranchs such as _Scyllium_, may have been derived from the +archipterygium by the gradual reduction of the rays on the postaxial +side of the axis and their condensation on the pre-axial side. The +Ichthyotomi such as _Xenacanthus_ show one stage in the reduction of +the postaxial rays, and a further stage is seen in the Notidanidae and +some other sharks like _Scymnus_ and _Acanthias_, in which a few +postaxial rays still remain. The condensation of the pre-axial rays +when further continued leads to one of the rays getting an attachment +to the girdle. Thus the fin comes to articulate with the girdle by two +basalia or basal pieces; a third attachment is formed in the same way +and the three basalia are called respectively pro-, meso-, and +meta-pterygia. By some authors the meta-pterygium and by others the +meso-pterygium is regarded as homologous with the axis of the +archipterygium. + +The pectoral fins of Elasmobranchs vary very much in their mode of +attachment. In some of the sharks, including the Notidanidae and +_Scyllium_, all three basalia articulate with the pectoral girdle, +while in others such as _Cestracion_ the meta-pterygium is excluded. +In Rays the propterygium and the meta-pterygium are long and narrow +and diverge much from one another; other basalia work their way in +between the meso-pterygium and meta-pterygium, and come to articulate +with the pectoral girdle. Sometimes they fuse and form a second +meso-pterygium. The radiale are greatly elongated and are segmented. + +In _Chimaera_ all three basalia are present, but the meso-pterygium +is shifted and does not articulate with the pectoral girdle[49]. + +In _Acipenser_ and _Polyodon_ the pectoral fin is built on the same +type as in Elasmobranchs, but becomes modified from the fact that the +propterygium is replaced by dermal bone which forms a large =marginal +ray=. Extra meso-pterygia are formed in the same way as in Rays. + +In _Polypterus_ the pro-and meta-pterygia have ossified while the +meso-pterygium remains chiefly cartilaginous; the fin-rays are also +chiefly ossified. + +In _Amia_, _Lepidosteus_, and certain Teleosteans like _Salmo_, not +only the propterygium but the meso-pterygium is almost suppressed by +the marginal ray. + +In the great majority of Teleosteans a still further stage is reached, +the endoskeletal elements, the basalia and radiale are almost entirely +suppressed and the fin comes to consist entirely of ossified fin-rays +of dermal origin. + +In some Teleosteans--_Exocaetus_, a herring, and _Dactylopterus_, a +gurnard--the pectoral fins are so enormously developed that by means +of them the fish is able to fly through the air for considerable +distances. The skeleton of these great fins is almost entirely +composed of dermal bone. + +PELVIC FIN. + +The pelvic fin is almost always further removed from the +archipterygial condition, and is in general more modified than is the +pectoral. Thus in the Ichthyotomi, while the pectoral fins are +archipterygia similar to those of _Ceratodus_, the pelvic fins consist +of an axis bearing rays on the postaxial side only, and prolonged +distally into a clasper. In Dipnoi however the pelvic fins are very +similar to the pectoral. In Elasmobranchs the meso-pterygium is +missing, the propterygium is small or absent, and the fin is mainly +composed of the meta-pterygium (generally called basi-pterygium) and +its radiale. The males in Elasmobranchii and Holocephali have the +distal end of the meta-pterygium prolonged into a clasper. + +In Ganoids and in Teleosteans the loss of the pelvic girdle causes the +pelvic fin to be still further removed from the primitive state. There +is always a large basi-pterygium which lies imbedded in the muscles +and meets its fellow at its proximal end. In Cartilaginous Ganoids it +has a secondary segmentation. Its relation to its fellow is subject to +much variation in Teleosteans, sometimes as in the Perch the two are +in contact throughout, sometimes as in the Salmon they meet distally +as well as proximally, but are elsewhere separated by a space, +sometimes as in the Pike and Bony Ganoids they diverge widely. The +radiale are articulated to the basi-pterygium. In Cartilaginous +Ganoids and _Polypterus_ they are well developed, in other Ganoids and +in Teleosteans they are in the main replaced by dermal fin-rays. + +In some Teleosteans such as the Cod the pelvic fins have migrated from +their usual position and come to be attached to the throat in front of +the pectoral fins. Fish with this arrangement are grouped together as +=jugulares=. + + +FOOTNOTES: + +[39] The following general works on fishes may be referred to: +Bashford Dean, _Fishes, Living and Fossil_, New York, 1895. A. +Günther, _An Introduction to the Study of Fishes_, Edinburgh, 1880. +A.A.W. Hubrecht and M. Sagemehl, _Fische_ in Bronn's _Classen und +Ordnungen des Thierreichs_, Band VI. Leipzig, 1876. + +[40] See W.G. Ridewood, _Nat. Sci._ vol. VIII. 1896, p. 380. Full +references are there given to the literature of the subject. + +[41] See H. Gadow and E.C. Abbott, _Phil. Trans._ vol. 186 (1895) B. +pp. 163-221. + +[42] C. Hasse, _Zeitschr. wiss. Zool._ LVII. 1893, p. 76. + +[43] C. Hasse, _Das natürliche System der Elasmobranchier auf +Grundlage des Baues und der Entwickelung ihrer Wirbelsäule_, Jena, +1879 and 1885, and "Die fossilen Wirbel, Morph. Studien I.-IV.," +_Morphol. Jahrb. Bd._ II., III. and IV. 1876-78. + +[44] See H.B. Pollard, _Anat. Anz._ X. 1894. + +[45] T.W. Bridge, "The Cranial Osteology of _Amia calva_," _J. Anat. +Physiol. norm. path._ 1876, vol. XI. p. 605. R. Shufeldt, "The +Osteology of _Amia calva_," _Ann. Rep. of the Commissioner for Fish +and Fisheries_, Washington, 1885. + +[46] A. Smith Woodward, _Nat. Sci._ vol. I. 1892, p. 28. Further +references are here given on the literature of the subject. + +[47] C. Gegenbaur, Ueber das Archipterygium, _Jena Zeitschr. der +Wirbelthiere_, 2^e Heft, 1873, vol. 7, and _Morphol. Jahrb._ XXII. +1894, p. 119. + +[48] The fins of _Ceratodus_ are very variable, no two being exactly +alike. Sometimes even the main axis bifurcates. See W.A. Haswell, +_Linn. Soc. N. S. Wales_, vol. VII. 1882. + +[49] Some of these views with regard to the homologies of the parts of +the fins are not accepted by all anatomists. + + + + +CHAPTER IX. + +CLASS II. AMPHIBIA[50]. + + +AMPHIBIA differ markedly from Pisces in the fact that in the more +abundant and familiar forms the skin is naked, and that when the +integument is prolonged into median fins they are devoid of fin-rays. +The notochord may persist, but bony vertebral centra are always +developed. These are sometimes biconcave, sometimes procoelous, +sometimes opisthocoelous. There is only one sacral vertebra, except in +rare cases. The cartilaginous cranium persists to a considerable +extent but is more or less replaced by cartilage bone, and overlain by +membrane bone. The basi-occipital is not completely ossified, and the +skull articulates with the vertebral column by means of two occipital +condyles formed by the exoccipitals. + +There is a large parasphenoid, but there are no ossifications in the +basisphenoidal, presphenoidal, and alisphenoidal regions. In most +cases the epi-otics and opisthotics are ossified continuously with the +exoccipitals. + +The palato-pterygo-quadrate bar is firmly united to the cranium, so +the skull is autostylic. The palatines and pterygoids are membrane +bones. Teeth are nearly always borne on the vomers and commonly on the +maxillae and premaxillae. There are no sternal ribs, and the sternum is +very intimately related to the pectoral girdle. There are no obturator +foramina. The limbs are as in the higher vertebrata, divisible into +upper arm, fore-arm, and manus (wrist and hand), and into thigh, shin, +and pes (ankle and foot) respectively. The posterior limb is, as a +rule, pentedactylate, but in nearly every case the pollex is vestigial +or absent. + + +_Order 1._ URODELA[51]. + +The Urodela are elongated animals with a naked skin, a persistent +tail, and generally four short limbs. + +The vertebral centra are opisthocoelous or biconcave, and there are +numerous precaudal vertebrae. Portions of the notochord commonly +persist in the intervertebral spaces. In the skull there is no +sphenethmoid forming a ring encircling the anterior end of the brain, +its place being in many cases partly taken by a pair of +orbitosphenoids. There is no quadratojugal, and the quadrate is more +or less ossified. The mandible has a distinct splenial, and the +articular is ossified. + +There is no definite tympanic cavity. The hyoid apparatus is +throughout life connected to the quadrate by ligament, and a large +basilingual plate does not occur. The ribs are short structures with +bifurcated proximal ends. In the pelvis the pubis remains +cartilaginous, and there is a bifid cartilaginous epipubis. The bones +of the fore-arm and shin remain distinct, and the manus never has more +than four digits. + +_Suborder_ (1). ICHTHYOIDEA. + +The vertebrae are amphicoelous, but the notochord remains but little +constricted throughout the whole length of the vertebral column. Three +or four branchial arches nearly always persist in the adult. The +cartilages of the carpus and tarsus remain unossified. + +The Ichthyoidea may be subdivided again into two groups:-- + +A. _Perennibranchiata_, whose chief distinguishing skeletal characters +are that the skull is elongated, the premaxillae are not ankylosed, the +maxillae are vestigial or absent; there are sometimes no nasals, and +the palatines bear teeth; + + e.g. _Siren_, _Proteus_, _Menobranchus_. + +B. _Derotremata_, whose chief distinguishing skeletal characters are +that there are large maxillae and nasals; teeth are borne by both +maxillae and premaxillae; there are no palatines; and both pectoral and +pelvic limbs are always present; + + e.g. _Amphiuma_, _Megalobatrachus_, _Cryptobranchus_. + +_Suborder_ (2). SALAMANDRINA. + +The vertebrae are opisthocoelous. The skull is broad, and teeth are +borne by both premaxillae and dentaries. Nasal bones are present. The +remains of only two branchial arches are found in the adult. The +carpus and tarsus are more or less ossified. + +This suborder includes the Newts (_Molge_), Salamanders +(_Salamandra_), and _Amblystoma_. + + +_Order_ 2. LABYRINTHODONTIA[52]. + +These are extinct Amphibia with a greatly developed dermal +exoskeleton, which is generally limited to the ventral surface. The +body and tail are long and in some cases limbs are absent. The teeth +are pointed and often have the dentine remarkably folded. The +vertebrae are amphicoelous, and are generally well ossified. The skull +is very solid, and has a greatly-developed secondary roof which hides +the true cranium and is very little broken up by fossae. Paired +dermal supra-occipitals are found, and there is an interparietal +foramen. The epi-otics and opisthotics form a pair of bones distinct +from the exoccipitals. Four simple limbs of moderate length are +generally present, and in some cases all four limbs are +pentedactylate. Among the better known genera of Labyrinthodonts are +_Mastodonsaurus_, _Nyrania_, and _Archegosaurus_. + + +_Order 3._ GYMNOPHIONA[53]. + +These animals form a group of abnormal worm-like Amphibia having an +exoskeleton in the form of subcutaneous scales arranged in rings. The +vertebrae are biconcave and are very numerous; very few however belong +to the tail. The skull has a complete secondary bony roof, the +mandible bears teeth and has an enormous backward projection of the +angular. The hyoid arch has very slender cornua and no distinct body, +it is attached neither to the cranium nor to the suspensorium. The +ribs are very long and there are no limbs or limb girdles. + + +_Order 4._ ANURA. + +These are tailless Amphibia, which except in a few instances, are +devoid of an exoskeleton. The vertebrae are as a rule procoelous, and +are very few in number. The post-sacral part of the spinal column +ossifies continuously, forming an unsegmented cylindrical rod, the +urostyle. Remains of the notochord persist, lying _vertebrally_, i.e. +enclosed within the centra of the several vertebrae, and not as in +Urodela lying between one vertebra and the next. The skull is very +short and wide. The mandible is almost always, if not invariably, +toothless. + +The frontals and parietals on each side are united so as to form a +pair of fronto-parietals, and a girdle-like sphenethmoid is present. + +The quadrate is not generally ossified. A predentary or +mento-meckelian bone is commonly present in the mandible, and a single +bone represents the angular and splenial. The branchial arches are +much reduced in the adult, and the distal ends of the cornua unite to +form a flat basilingual plate of a comparatively large size. + +Ribs are very little developed. Clavicles are present. The ilia are +very greatly elongated. The anterior limb has four well-developed +digits and a vestigial pollex, and is of moderate length; the radius +and ulna have fused. The posterior limb is greatly elongated and is +pentedactylate; the tibia and fibula are fused, while the calcaneum +and astragalus are greatly elongated, and it is largely owing to them +that the length of the limb is so great. The group includes the Frogs +and Toads, the predominant Amphibia of the present time. + + +FOOTNOTES: + +[50] T.H. Huxley, _Amphibia_ (_Encyclopaedia Britannica_). + +[51] See R. Wiedersheim, _Morphol. Jahrb._ Bd. III. 1877, p. 459. + +[52] See A. Fritsch, _Fauna der Gaskohle_, Prague, 1883-85-86, also +writings of Cope, Credner, Huxley, H. v. Meyer, Miall. + +[53] See R. Wiedersheim, _Anatomie der Gymnophionen_, Jena, 1879. + + + + +CHAPTER X. + +THE SKELETON OF THE NEWT (_Molge cristata_). + + +I. EXOSKELETON. + +The skin of the Newt is quite devoid of any exoskeletal structures. +The only exoskeletal structures that the animal possesses are the +teeth, and these are most conveniently described with the +endoskeleton. + + +II. ENDOSKELETON. + +The endoskeleton of the Newt, though ossified to a considerable +extent, is more cartilaginous than is that of the frog. It is +divisible into an =axial portion= including the vertebral column, +skull, ribs, and sternum, and an =appendicular portion= including the +skeleton of the limbs and their girdles. + +1. THE AXIAL SKELETON. + +A. THE VERTEBRAL COLUMN. + +This consists of about fifty vertebrae arranged in a regular +continuous series. The first vertebra differs a good deal from any of +the others; the seventeenth or sacral vertebra and the eighteenth or +first caudal also present peculiarities of their own. The remaining +vertebrae are divided by the sacrum into an anterior series of =trunk= +vertebrae which bear fairly large ribs, and a posterior series of +=caudal= vertebrae, all of which except the first few are ribless. + +THE TRUNK VERTEBRAE. + +Any vertebra from the second to the sixteenth may be taken as a type +of the trunk vertebrae. + +The general form is elongated and somewhat hour-glass shaped, and the +=centra= are convex in front and concave behind; an opisthocoelous +condition such as this is quite exceptional in Anura. The =notochord= +may persist intervertebrally[54], but in the centre of each vertebra +it becomes greatly constricted or altogether obliterated, and replaced +by marrow. The superficial portion of the centrum is ossified, while +the articular surfaces are cartilaginous. The =neural arches= are low +and articulate together by means of =zygapophyses= borne on short +diverging processes. The anterior zygapophyses look upwards, the +posterior downwards. Each neural arch is drawn out dorsally into a +very slight cartilaginous =neural spine=. + +On each centrum, at a little behind the middle line, there arise a +pair of short backwardly-directed =transverse processes=; each of +which becomes divided into two slightly divergent portions, a dorsal +portion which meets the tubercular process of the rib and is derived +from the neural arch, and a ventral portion which meets the capitular +process of the rib and is derived from the ventral or haemal arch. The +division between these two parts of the transverse processes can be +traced back as far as the sacrum. + +The =first vertebra= as already mentioned differs much from all the +others. It has no ribs, and presents anteriorly two slightly divergent +concave surfaces which articulate with the occipital condyles of the +skull. Between these surfaces the dorsal portion of the anterior face +of the centrum is drawn out into a prominent =odontoid process=, the +occurrence of which renders it probable that the first vertebra of +the newt is really the axis, and that the atlas with the exception of +the odontoid process has become fused with the skull. The sacral +vertebra or =sacrum= differs from the vertebrae immediately in front +of it only in the fact that its transverse processes are stouter and +more obviously divided into dorsal and ventral portions. + +THE CAUDAL VERTEBRAE. + +The =caudal vertebrae= are about twenty-four in number. The anterior +ones have hour-glass shaped centra, and short backwardly-directed +transverse processes. The middle and posterior ones have rather +shorter centra, and are without transverse processes. The neural +arches resemble those of the trunk vertebrae, but each is drawn out +into a rather high cartilaginous neural spine abruptly truncated +anteriorly. All the caudal vertebrae except the first have also a +haemal arch, which is very similar to the neural arch, and is drawn +out into a haemal spine quite similar to the neural spine. Both neural +and haemal arches are ossified continuously with the centra. + +B. THE SKULL. + +The skull of the newt is divisible into three principal parts:-- + +(1) an axial part, the =cranium proper=, which encloses the brain and +to which + +(2) the =capsules= of the =auditory and olfactory sense organs= are +fused; + +(3) the skeleton of the =jaws and hyoid apparatus=. The skull is much +flattened and expanded, though not so much as in the frog. + +(1) THE CRANIUM PROPER. + +The =cranium proper= or =brain case= is an unsegmented tube which +remains partly cartilaginous, and is partly converted into cartilage +bone, partly sheathed by membrane bone. The roof and floor of the +cartilaginous cranium are, as is the case also in the frog, pierced +by holes or fontanelles, and these are so large that the main part of +the roof and floor comes to be formed by membrane bone. + +Two pairs of large ossifications take place in the cranial walls. Of +these the more posterior on each side represents the =exoccipital= and +all three =periotic= bones. It bears a small convex patch of cartilage +for articulation with the atlas, and with its fellow forms the +boundary of the foramen magnum. + +Two foramina pierce the exoccipital just in front of the occipital +condyle and transmit respectively the glossopharyngeal and +pneumogastric (fig. 21, X) nerves. Lying laterally to these nerve +openings is seen a patch of cartilage, the =stapes=, which is +homologous with the stapes or proximal element of the columellar chain +in the frog. Further forward in front of the stapes is the small +opening for the exit of the facial nerve, and seen in a lateral view +close to the orbitosphenoid, that for the trigeminal (fig. 21, C, 5). + +In front of these large bones the lateral parts of the cranial walls +remain cartilaginous for a short distance, and then there follow two +elongated bones, the =orbitosphenoids= (fig. 21, B and C, 11), pierced +by the foramina for the exit of the optic nerves. These bones partly +correspond to the sphenethmoid of the frog. + +The _membrane bones_ connected with the cranium are the _parietals_, +_frontals_ and _prefronto-lachrymals_ on the dorsal surface, and the +_parasphenoid_ on the ventral surface. + +The _parietals_ (fig. 21, A and C, 6) roof over the posterior part of +the great dorsal fontanelle and overlap the exoccipito-periotics. They +meet one another along a sinuous suture in the middle line, as do also +the _frontals_ which overlap them in front. The _frontals_ and +_parietals_ both extend for a short distance down the sides of the +cranium and meet the orbitosphenoids. The _prefronto-lachrymals_ (fig. +21, A and C, 7) connect the frontals with the maxillae. + +[Illustration FIG. 21. A DORSAL, B VENTRAL, AND C LATERAL VIEWS OF THE +SKULL OF A NEWT (_Molge cristata_) × 2-1/2 (after PARKER). + +The cartilage is dotted, the cartilage bones are marked with dots and +dashes, the membrane bones are left white. + + 1. premaxillae. + 2. anterior nares. + 3. posterior nares. + 4. nasal. + 5. frontal. + 6. parietal. + 7. prefronto-lachrymal. + 8. maxillae. + 9. vomero-palatine. + 10. parasphenoid. + 11. orbitosphenoid. + 12. pterygoid. + 13. squamosal. + 14. pro-otic region of exoccipito-periotic. + 15. quadrate. + 16. quadrate cartilage. + 17. exoccipital region of exoccipito-periotic. + 18. articular. + 19. articular cartilage. + 20. dentary. + 21. splenial. + 22. middle narial passage. + II. V. VII. IX. X. foramina for the exit of cranial nerves.] + +On the ventral surface is the large _parasphenoid_ (fig. 21, B, 10), +which is widest behind and overlapped anteriorly by the +vomero-palatines. + +(2) THE SENSE CAPSULES. + +The =auditory capsules= become almost completely ossified continuously +with the exoccipitals; they have been already described. + +The =nasal capsules= are large and quite unossified though they are +overlain by membrane bone. They appear on the dorsal surface between +the anterior nares and the nasal process of the premaxillae. They +enclose the nasal organs, bound the inner side of the anterior narial +opening, and are connected with one another posteriorly by a +cartilaginous area. + +Developed in connection with the nasal capsules are a pair of rather +large _nasals_ (fig. 21, A and C, 4), which lie on the dorsal surface +immediately in front of the frontals. Each forms part of the posterior +boundary of one of the anterior nares, and the two are separated from +one another in the middle line by the nasal process of the premaxillae +(fig. 21, A, 1), and the opening of the =middle narial passage= (fig. +21, A and B, 22), which passes right through the skull. + +On the ventral surface of the skull and forming the greater part of +the boundary of the posterior nares are two large bones, the +_vomero-palatines_ (fig. 21, B and C, 9). Each consists of a wide +anterior portion, partly separated from its fellow in the middle line +by the ventral opening of the middle narial passage, and of a long +pointed posterior portion which is separated from its fellow by the +_parasphenoid_, and bears a row of small pointed teeth formed of +dentine capped with enamel. + +(3) THE JAWS. + +The =upper jaw= of the newt is a discontinuous structure divided into +two parts, an anterior part which consists of membrane bones, the +_maxillae_ and _premaxillae_, and a posterior part which remains mainly +cartilaginous. + +The _premaxillae_ are united, forming a single bone, which in a +ventral view is seen to meet the maxillae and vomero-palatines, and in +a dorsal view to send back a nasal process (fig. 21, A, 1) between the +nasals. + +The _maxillae_ are large bones, each terminating in a point +posteriorly. A single row of teeth similar to those on the +vomero-palatines runs along the outer margin of the maxillae and +premaxillae. + +The posterior part of the upper jaw forms a mass of cartilage which +extends forwards towards the maxillae as a long pointed process whose +ventral surface and sides are overlapped by a membrane bone, the +_pterygoid_ (fig. 21, 12). + +The suspensorial bones include the =quadrate= and _squamosal_. The +=quadrate= (fig. 21, 15) which forms the true =suspensorium= is +directed forwards and outwards, and is terminated by a patch of +cartilage with which the mandible articulates. + +The lower jaw or mandible remains partly cartilaginous, while its +ossifications include two membrane bones and one cartilage bone. The +cartilage bone is the =articular= (fig. 21, C, 18), it forms the +posterior part of the ramus, extends forwards for some distance along +its inner side, and is terminated posteriorly by a patch of cartilage +which articulates with the quadrate. The _dentary_ (fig. 21, C, 20) is +a large bone which forms the anterior part and nearly all the outer +half of each ramus, and bears teeth similar to those of the upper jaw. +Attached to its inner face is a long slender _splenial_ (fig. 21, C, +21). + +THE HYOID APPARATUS. + +This consists of the hyoid arch and part of the first two branchial +arches. + +The =hyoid arch= (fig. 29, A, 2) consists of a pair of =cornua=, each +of which is divided into two halves. The dorsal half forming the +=cerato-hyal= is mainly ossified though tipped with cartilage, and is +connected by ligament with the suspensorium. The ventral half +(=hypo-hyal=) is cartilaginous, and is connected with the +basibranchial. + +The =branchial arches= consist of a median piece, the =basibranchial=, +which is ossified in the centre and cartilaginous at either end, and +of two pairs of =cerato-branchials= which are attached to the +cartilaginous part (fig. 29, A, 8) of the basibranchial. The first +cerato-branchial is chiefly ossified, the second (fig. 29, A, 4) is a +good deal smaller and is cartilaginous. Both are united dorsally to a +single =epi-branchial=, which is terminated by a small cartilaginous +area at the free end but is elsewhere well ossified. + +C. THE RIBS. + +The ribs are short imperfectly ossified structures, bifid at their +proximal end where they articulate with the transverse processes, and +tipped both proximally and distally with cartilage. The dorsal portion +of the proximal end corresponds to the =tuberculum= of the ribs of +higher animals, and the ventral portion to the =capitulum=. Some of +the anterior ribs have a step-like notch on their dorsal surfaces. + +The second to twelfth ribs are fairly equal in size, but further back +they decrease slightly. The ribs which connect the sacral vertebrae +with the ilia are however large. The short ribs borne on the anterior +caudal vertebrae are cartilaginous. + +D. THE STERNUM. + +The sternum (fig. 22, A, 6) is a rather broad plate of cartilage, +drawn out posteriorly into a median process marked by a prominent +ridge. On its antero-lateral margins it bears surfaces for +articulation with the pectoral girdle. + +2. THE APPENDICULAR SKELETON. + +A. THE PECTORAL GIRDLE. + +This is of a very simple character, and remains throughout life in an +imperfectly ossified condition. It consists of a dorsal =scapular +portion=, and a ventral =coracoid portion= partially divided into an +anterior part, the precoracoid, and a posterior part, the =coracoid=. + +[Illustration FIG. 22. A VENTRAL, AND B LATERAL VIEW OF THE SHOULDER +GIRDLE AND STERNUM OF AN OLD MALE CRESTED NEWT (_Molge cristata_) × 3 +(after PARKER). + + 1. scapula. + 2. suprascapula. + 3. coracoid. + 4. glenoid cavity. + 5. precoracoid. + 6. sternum.] + +The =scapular portion= is a slightly curved oblong plate; its proximal +third the =scapula= (fig. 22, 1) is ossified and bounds part of the +well-marked =glenoid cavity= (fig. 22, 4); its distal portion forms a +large oblong cartilaginous plate, the =suprascapula= (fig. 22, 2). + +The =precoracoid= (fig. 22, 5) forms a small forwardly-directed +cartilaginous plate. The =coracoid= (fig. 22, 3) forms a much larger +plate, the greater part of which is unossified and overlaps its fellow +in the middle line, the two being overlapped by the sternum. Around +the glenoid cavity is an area which is mainly ossified and is +continuous with the scapula. + +B. THE ANTERIOR LIMB. + +This is divisible into three parts, the =upper arm= or =brachium=, the +=fore-arm= or =antibrachium=, and the =manus=. + +The =upper arm= includes a single bone, the =humerus=. + +The =humerus= is a slender bone cylindrical in the middle and expanded +at either end, the proximal part forms a rounded =head= which +articulates with the glenoid cavity. Along the proximal part of the +anterior or pre-axial surface runs a strong =deltoid ridge=. The +proximal part of the postaxial surface also bears a small outgrowth. + +The =fore-arm= contains two bones, the =radius= and =ulna=, both of +which are small and imperfectly ossified at their terminations. + +The =radius= (fig. 23, B, 11) or pre-axial bone is rather the larger +of the two, and is considerably expanded at its proximal end. The +=ulna= or postaxial bone is somewhat expanded distally, but is not +drawn out proximally into an olecranon process. + +The =manus= consists of two parts, a group of small bones forming the +=carpus= or =wrist=, and the =hand=. + +The =carpus= is in a very simple unmodified condition as compared with +that of the Frog. It consists of a proximal row of two bones and a +distal row of four, with one, the =centrale=, interposed between. All +these bones are small and polygonal and are imbedded in a plate of +cartilage. + +The bones of the proximal row are a smaller pre-axial bone, the +=radiale= (fig. 23, B, 13), and a larger postaxial bone, which +represents the fused =ulnare= and =intermedium= of the very simple +carpus described on pp. 26 and 27. + +The four bones of the distal row are respectively =carpalia= 2, 3, 4 +and 5. + +The =hand= consists of four digits, that corresponding to the thumb of +the human hand, judging from the analogy of the frog probably being +the one that is absent. + +[Illustration FIG. 23. A RIGHT POSTERIOR, AND B RIGHT ANTERIOR LIMB OF +A NEWT × 1-1/2 (_Molge cristata_). + + 1. femur. + 2. tibia. + 3. fibula. + 4. tibiale. + 5. intermedium. + 6. fibulare. + 7. centrale of tarsus. + 8. tarsale 1. + 9. tarsalia 4 and 5 fused. + 10. humerus. + 11. radius. + 12. ulna. + 13. radiale. + 14. intermedium and ulnare fused. + 15. centrale of carpus, the pointing line passes across carpale 2. + 16. carpale 3. + 17. carpale 5. + I. II. III. IV. V. digits.] + +Each digit consists of a somewhat elongated =metacarpal= and of two or +three phalanges. The metacarpals are contracted in the middle and +expanded at either end. They are connected with the carpus by +cartilage, and the articulations between the several phalanges, and +between the metacarpals and phalanges are also cartilaginous. The +second, third, and fifth digits have two phalanges apiece, the fourth, +which is the longest, has three. The second metacarpal in the +specimen examined and figured articulates partly with carpale 2, +partly with carpale 3. + +C. THE PELVIC GIRDLE. + +The pelvic girdle of the Newt is in a much less modified condition +than is that of the Frog (see p. 165). It consists of a dorsal +element, the =ilium=, a posterior ventral element, the =ischium=, and +an anterior ventral element, the =pubis=, to which is attached an +=epipubis=. + +The =ilium= is a somewhat cylindrical bone which at its ventral end +meets the ischium, and forms part of the =acetabulum=. It is then +directed upwards and slightly backwards, and is attached to the ribs +of the sacral vertebra. + +The =ischia= are a pair of somewhat square bones which meet one +another in the middle line; they form part of the acetabulum, and are +united to the ilia above. + +In front of the ischia is a narrow cartilaginous area which represents +the =pubes=. Projecting forwards from it is a bifid cartilaginous +=epipubis=. + +D. THE POSTERIOR LIMB. + +This is divisible into a proximal portion, the =thigh=, a middle +portion, the =crus= or =shin=, and a distal portion, the =pes=. + +The =thigh= consists of a single bone, the =femur= (fig. 23, A, 1), +which has a thin shaft and expanded ends. The anterior part of the +pre-axial border and posterior part of the postaxial border bear +slight outgrowths. + +The =crus= or =shin= includes two short bones, the =tibia= and +=fibula=, which are nearly equal in length. The pre-axial bone or +tibia is a straight bone thickest at its proximal end, the postaxial +bone or =fibula= (fig. 23, A, 3) is a rather stouter curved bone of +nearly equal diameter throughout. + +The =pes= includes the =tarsus= or =ankle=, and the =foot=. + +The =tarsus= consists of eight small bones arranged in a proximal row +of three, the =tibiale=, =intermedium= and =fibulare=, and a distal +row of four =tarsalia=, with one bone, the =centrale= (fig. 23, A, 7), +interposed between the two rows. In the specimen examined, the +=tibiale=, is a small bone articulating with the tibia, the +=intermedium= (fig. 23, A, 5) is larger and articulates with both +tibia and fibula, the =fibulare= is the largest of the three and +articulates with the fibula. + +The bones of the distal row are =tarsalia 1=, =2=, =3=, and a bone +representing =4= and =5= fused. In the specimen examined tarsale 1 is +pushed away dorsally (fig. 23, A, 8), so as to lie between the tibiale +and tarsale 2. All the tarsal bones are small and somewhat polygonal, +and are connected with one another, and with the tibia and fibula on +the one hand, and with the metatarsals on the other by a thin layer of +cartilage. + +The five =digits= of the foot each consist of a =metatarsal= and of a +certain number of =phalanges=. In the specimen examined, owing to the +shifting of tarsale 1, the first metatarsal as well as the second +articulates with tarsale 2, while the fifth metatarsal articulates +partially with the bone representing the fused tarsalia 4 and 5, +partially with the fibulare. All the bones of the digits except the +distal phalanges are terminated at each end by cartilaginous +epiphyses, the distal phalanx of each digit has a cartilaginous +epiphysis only on its proximal end. + +The first, second, and fifth digits have two phalanges apiece, the +third and fourth have three. + +Figure 31 B, showing a Newt's tarsus copied from Gegenbaur, has +precisely the arrangement generally regarded as primitive for the +higher vertebrates, except that tarsalia 4 and 5 are fused. + + +FOOTNOTES: + +[54] i.e. between one vertebra and the next. + + + + +CHAPTER XI. + +THE SKELETON OF THE FROG[55] (_Rana temporaria_). + + +I. EXOSKELETON. + +The skin of the frog is smooth and quite devoid of scales or other +exoskeletal structures. The only exoskeletal structures met with in +the frog are:-- + +1. The =teeth=, which are most conveniently described with the +endoskeleton. + +2. The horny covering of the calcar or prehallux (see p. 167). + + +II. ENDOSKELETON. + +The endoskeleton of the adult frog consists partly of cartilage, +partly of bone and each of these types of tissue occurs in two forms. +The cartilage may be hyaline, as in the omosternum and xiphisternum, +or may be more or less calcified as in part of the suprascapula and +the epiphyses of the limb bones. The bone may be cartilage bone, or +membrane bone. + +The skeleton is divisible into an =axial portion= consisting of the +skull, vertebral column, and sternum, and an =appendicular portion= +consisting of the skeleton of the limbs and their girdles. + +1. THE AXIAL SKELETON. + +A. THE VERTEBRAL COLUMN. + +The vertebral column is a tube, formed of a series of ten bones which +surround and protect the spinal cord. Of these ten bones nine are +vertebrae, while the tenth is a straight rod, the =urostyle=, and is +almost as long as all the vertebrae put together. The second to eighth +vertebrae inclusive have a very similar structure, but the first and +ninth differ from the others. + +Any one of the second to eighth vertebrae forms a bony ring with a +somewhat thickened floor, the =centrum= or body, which articulates +with the centra of the immediately preceding and succeeding vertebrae. +The articulating surfaces are covered with cartilage and are +procoelous, or convex in front and concave behind. The eighth vertebra +is however amphicoelous or biconcave. The centrum of each vertebra +encloses an isolated vestige of the notochord. The =neural arch= forms +the roof and sides of the neural canal, which is very spacious in the +anterior vertebrae, but becomes more depressed in the posterior ones. +The arch bears the =neural spine=, a low median ridge of variable +character, and is drawn out in front and behind, forming the two pairs +of articulating surfaces or =zygapophyses= by means of which the +vertebrae are attached together. Of these the anterior articulating +surfaces or =prezygapophyses= look upwards and slightly inwards, while +the posterior articulating surfaces or =postzygapophyses= look +downwards and slightly outwards. The sides of the neural arches are +drawn out into a pair of prominent =transverse processes=. Those of +the second vertebra look somewhat forwards, those of the third look +directly outwards or somewhat forwards, while those of the fourth, +fifth, and sixth are directed slightly backwards, and those of the +seventh and eighth nearly straight outwards. All the transverse +processes are terminated by very small cartilaginous =ribs=. + +SPECIAL VERTEBRAE. + +The =first vertebra= is a ring-like structure with a much depressed +centrum. It bears in front two oval concave surfaces for articulation +with the condyles of the skull, while the centrum is terminated behind +by a prominent convex surface. There are as a rule no transverse +processes, and the postzygapophyses look downwards and outwards. +Occasionally however transverse processes do occur. Projecting +forwards from the centrum is a minute process better developed in the +Newt. This resembles an odontoid process, and it has hence been +supposed that the first vertebra is homologous with the axis of +mammalia, and that the atlas of the frog is fused with the skull. + +The =ninth vertebra= has very stout transverse processes directed +backwards and somewhat upwards. They articulate with the pelvic girdle +and hence this vertebra is regarded as the =sacrum=. The neural arch +is much depressed, the centrum is convex in front and bears on its +posterior surface two short rounded processes for articulation with +the urostyle. + +The =urostyle= is a long rod-like bone forming the posterior +unsegmented continuation of the vertebral column. It is probably +equivalent to three vertebrae, the tenth, eleventh, and twelfth fused +together, and to an unsegmented rod of cartilage which lies ventral to +the notochord. The anterior end is expanded and bears two concave +articular surfaces by means of which it articulates with the sacrum. A +prominent ridge runs along the dorsal surface, but gradually +diminishes when traced back. The anterior portion contains a canal +which is a continuation of the neural canal. At a point not far from +the anterior end, this canal communicates with the exterior by a pair +of minute holes which correspond with the intervertebral foramina. + +B. THE SKULL[56]. + +The skull of the Frog consists of three principal parts:-- + +(1) an axial part, the =cranium proper=, which encloses the brain. To +it are firmly fused + +(2) the =capsules of the olfactory and auditory sense organs=, + +(3) lastly there is the =hyoid apparatus= and the =skeleton of the +jaws=. + +The skull is by no means so completely ossified as is the vertebral +column, but in addition to the cartilage bone, there is a great +development of membrane bone in connection with it. + +The skull has a peculiarly flattened and expanded form depending on +the wide lateral separation of the jaws from the cranium. + + +(1) THE CRANIUM PROPER or Brain case. + +This is an unsegmented tube, which is widest behind. It remains to a +considerable extent cartilaginous, but is partly converted into +cartilage bone, partly sheathed in membrane bone. Its roof is +imperfect, being pierced by three holes or =fontanelles=, one large +anterior fontanelle (fig. 25, A, 9), and two smaller posterior +fontanelles (fig. 25, A, 10). + +The cartilage bones of the cranium proper are the two =exoccipitals= +and the =sphenethmoid=. + +The =exoccipitals= (figs. 24, 25, and 26, 6) are a pair of irregular +bones bounding the foramen magnum at the posterior end of the skull. +They almost completely surround the foramen magnum, and bear a pair of +oval convex surfaces, the =occipital condyles=, with which the first +vertebra articulates. The bones generally called the exoccipitals of +the frog include the =epi-otic= and =opisthotic= elements of many +skulls, in addition to the exoccipitals. + +[Illustration FIG. 24. A DORSAL, AND B VENTRAL VIEWS OF THE CRANIUM OF +A COMMON FROG (_Rana temporaria_) × 2 (after PARKER). + +In this and the next two figs. cartilage is dotted, cartilage bones +are marked with dots and dashes, membrane bones are left white. + + 1. sphenethmoid. + 2. fronto-parietal. + 3. pterygoid. + 4. squamosal. + 6. exoccipital. + 7. parasphenoid. + 8. pro-otic. + 9. quadratojugal. + 10. maxillae. + 11. nasal. + 12. premaxillae. + 13. anterior nares. + 14. vomer. + 15. posterior nares. + 16. palatine. + 18. columella. + 19. quadrate. + 20. occipital condyle. + II. optic foramen. + V. VII. foramen for exit of trigeminal and facial nerves. + IX. X. foramina for exit of glossopharyngeal and pneumogastric + nerves.] + +The patch of unossified cartilage immediately external to the +occipital condyle is pierced by two small foramina, through which the +ninth and tenth nerves leave the cranial cavity. The ninth nerve +passes through the more external of these foramina, the tenth through +the one nearer the condyle. The foramina lie however very close +together and are sometimes confluent. The cranial walls for a +considerable distance in front of the occipitals are unossified, but +the anterior end of the cranial cavity is encircled by another +cartilage bone, the =sphenethmoid= (figs. 24 and 25, 1) or girdle +bone. This partly corresponds to the orbitosphenoids of the Newt's +skull. Anteriorly it is pierced by a pair of small foramina through +which the ophthalmic branches of the trigeminal nerve pass out. + +The anterior part of the cranial cavity is divided into two halves by +a vertical plate, the =mesethmoid=. Some little distance behind the +sphenethmoid the ventro-lateral walls of the cartilaginous cranium are +pierced by a pair of rather prominent holes, the =optic foramina= +(figs. 24 and 25, B, II), and at a similar distance further back, +occupying a kind of notch in the pro-otic are the large =trigeminal +foramina=, through which the fifth and seventh nerves leave the +cranium. Between the trigeminal and optic foramina are the very small +foramina for the sixth nerves (fig. 25, B, VI). + +The _membrane bones_ of the cranium proper include the +_fronto-parietals_ and the _parasphenoid_. + +The _fronto-parietals_ (figs. 24 and 26, A, 2) form a pair of long +flat bones closely applied to one another in the middle line, the line +of junction being the =sagittal suture=. They cover over the +fontanelles and overlap the sphenethmoid in front. + +The _parasphenoid_ (figs. 24 and 26, B, 7) is a bone shaped like a +dagger with a very short handle. It lies on the ventral surface of the +cranium, the blade being directed forwards and underlying the +sphenethmoid; its lateral processes underlie the auditory capsules. + + +(2) THE SENSE CAPSULES. + +The sense capsules are cartilaginous or bony structures which surround +the olfactory and auditory organs and are closely united to the +cranium. + +[Illustration FIG. 25. A DORSAL AND B VENTRAL VIEW OF THE CRANIUM OF A +COMMON FROG (_Rana temporaria_) from which the membrane bones have +mostly been removed. × 2 (after PARKER). + + 1. sphenethmoid. + 2. palatine. + 3. pterygoid. + 4. quadrate. + 5. columella. + 6. exoccipital. + 7. ventral cartilaginous wall of cranium. + 8. pro-otic. + 9. anterior fontanelle. + 10. right posterior fontanelle. + 11. quadratojugal. + 12. nasal capsule. + II. V. VI. IX. X. foramina for exit of cranial nerves.] + +The =auditory capsules= are fused with the sides of the posterior +end of the cranium just in front of the exoccipitals. They are +largely cartilaginous, but include in their anterior walls a pair +of irregular cartilage bones, the =pro-otics= (figs. 24 and 25, 8). +The cartilaginous area lying ventral to the pro-otic and external +to the exoccipital is pierced by a rather prominent hole, the +=fenestra ovalis=, which forms a communication between the internal +ear cavity, and a space the tympanic cavity, which lies at the side +of the head, and is bounded externally by the tympanic membrane. The +fenestra ovalis is occupied by a minute cartilaginous structure, the +=stapes=, and articulated partly to this and partly to a slight recess +in the pro-otic is the =columella= (fig. 25, B, 5), a rod in part +bony and in part cartilaginous, whose outer end is attached to the +tympanic membrane. The columella and stapes are together homologous +with the mammalian auditory ossicles and with the hyomandibular of +Elasmobranchs. Sometimes the term columella is used to include the +whole ossicular chain,--the columella together with the stapes. + +The =olfactory= or =nasal capsules= (fig. 25, B, 12) are fused with +the anterior end of the cranium and differ from the auditory capsules +in being to a great extent unossified. There are however two pairs of +membrane bones developed in connection with them, the _vomers_ and the +_nasals_. They are drawn out into three pairs of cartilaginous +processes, on the dorsal surface into the =prenasal= and =alinasal= +processes which bound the external nares, and on the ventral surface +towards the middle line into the forwardly-projecting =rhinal= +processes. + +The _nasals_ (figs. 24 and 26, 11) form a pair of triangular bones +lying dorsolaterally in front of the fronto-parietals. Their bases are +turned towards one another and their apices are directed outwards and +backwards. They correspond in position with the prefrontals of the +reptilian skull as well as with the nasals. + +The _vomers_ are a pair of irregular bones lying on the ventral +surface of the olfactory capsules. Each bears on its inner and +posterior angle a group of minute pointed teeth, while its outer +border is drawn out into three or four small slightly diverging +processes, the two posterior of which form the inner boundary of the +=posterior nares= (fig. 24, B, 15). + +(3) THE JAWS. + +The =upper jaw= consists of a rod of cartilage connected with the +cranium near its two ends, but widely separated from it in the middle. +It is almost completely overlain by membrane bone. With its posterior +end the lower jaw articulates. + +The membrane bones of the upper jaw include first the _premaxillae_, a +small bone meeting its fellow in the middle line, and forming the +extreme anterior end of the upper jaw. It gives off on its dorsal +surface a backwardly-projecting process. It is connected behind with +the _maxillae_ (figs. 24 and 26, 10), a long flattened bone which forms +the greater part of the margin of the upper jaw, and gives off near +its anterior end a short process which projects upwards and meets the +nasal. + +[Illustration FIG. 26. A, LATERAL VIEW OF THE SKULL, B, POSTERIOR VIEW +OF THE CRANIUM OF A COMMON FROG (_Rana temporaria_) × 2 (after +PARKER). + + 1. sphenethmoid. + 2. fronto-parietal. + 3. pterygoid. + 4. squamosal. + 5. tympanic membrane. + 6. exoccipital. + 7. parasphenoid. + 8. pro-otic. + 9. quadratojugal. + 10. maxillae. + 11. nasal. + 12. premaxillae. + 13. anterior nares. + 14. mento-meckelian. + 15. dentary. + 16. angulo-splenial. + 17. basilingual plate. + 19. quadrate. + 20. columella. + 21. occipital condyle. + 22. anterior cornu of the hyoid (cerato-hyal). + 23. foramen magnum. + + II. IX. X. foramina for the exit of cranial nerves.] + +Both maxillae and premaxillae are grooved ventrally, and bear, attached +to the outer more prominent margin of the groove, a row of minute +conical teeth. These teeth are =pleurodont=, that is, are ankylosed by +their bases and outer sides to the margin of the jaw. Each tooth is a +hollow cone, the basal part of which is formed of bone, the apical +part of dentine, capped by a very weak development of enamel. + +The posterior end of the maxillae is overlapped by a small bone, the +_quadratojugal_ (figs. 24 and 26, 9), whose posterior end forms part +of the articular surface for the lower jaw. Just behind the +quadratojugal there is a small unossified area which lies at the angle +of the mouth, and is connected by a narrow bar of cartilage with the +cranium; this forms the =quadrate= (figs. 24 and 26, 19). A +backwardly-directed outgrowth from the cartilaginous bar more or less +completely surrounds the tympanic membrane, forming the tympanic ring. +When followed back the maxillae and quadratojugal diverge further and +further from the cranium, till the angle of the mouth comes to be +separated from the foramen magnum by a space nearly double the width +of the cranium. This space is bridged over to a considerable extent by +two triradiate bones, the _pterygoid_ and _squamosal_. + +The _pterygoid_ (figs. 24 and 26, 3) is a large bone, whose anterior +limb runs forwards meeting the maxillae and palatine; while its inner +limb meets the auditory capsule and parasphenoid, and its outer limb +runs backwards and outwards to the angle of the mouth. The _palatine_ +is a small transversely-placed bone, which connects the pterygoid with +the anterior part of the sphenethmoid. The _squamosal_ (figs. 24 and +26, 4) is a T-shaped bone whose anterior arm is pointed and passes +forwards to meet the pterygoid. The posterior upper arm is closely +applied to the pro-otic, while the posterior lower arm meets the +pterygoid and quadratojugal at the angle of the jaw, and surrounds the +narrow cartilaginous bar of the quadrate which goes to join the +cranium. The squamosal is probably homologous with the squamosal +together with the pre-opercular of Bony Ganoids. + +The quadrate and squamosal form the =suspensorium= by which the lower +jaw is connected with the cranium. + +The =lower jaw= or =mandible= consists of a pair of cartilaginous rods +(=Meckel's cartilages=) in connection with each of which there are +developed two membrane bones and one cartilage bone. The cartilage +bone is the =mento-meckelian= (fig. 26, A, 14), a very small +ossification at the extreme anterior end. The membrane bones are the +_angulo-splenial_ and the _dentary_. The _angulo-splenial_ is a strong +flat bone which forms the inner and lower part of the mandible for the +greater part of its length. Its dorsal surface is produced into a +slight =coronoid process=. The _dentary_ (fig. 26, A, 15) is a flat +plate which covers the outer surface of the anterior half of the +mandible, as far forwards as the mento-meckelian. The lower jaw is +devoid of teeth. The part of Meckel's cartilage which in most +vertebrates ossifies, forming the articular bone, remains unossified +in the Frog. + +THE HYOID APPARATUS. + +The =hyoid= of the adult Frog is formed of the modified hyoid and +branchial arches of the tadpole. It consists of a broad thin plate of +cartilage, the =basilingual plate= (fig. 29, B, 1), drawn out into two +pairs of long processes, the =cornua=. The basilingual plate is +broader in front than behind, and is formed from the fused ventral +ends of the hyoid and branchial arches of the tadpole. + +The =anterior cornua= (fig. 29, B, 2) form a pair of long slender +cartilaginous rods which project from the body of the hyoid at first +forwards, then backwards, and finally upwards and somewhat forwards +again, to be united to the auditory capsules just below the fenestrae +ovales. They are formed from the dorsal portion of the hyoid arch of +the tadpole and are homologous with the cerato-hyals of the Dogfish. + +The =posterior cornua= form a pair of straight bony rods diverging +outwards from the posterior end of the basilingual plate. They are +formed from the fourth branchial arches of the tadpole, and differ +from the rest of the hyoid apparatus in being well ossified. + +The =columellar chain=, which has been already described (p. 157), +should be mentioned with the hyoid as it is homologous to the +hyomandibular of fishes. + +The =sternum= of the =Frog=, though regarded as part of the axial +skeleton, is so intimately connected with the pectoral girdle, that it +will be described with the appendicular skeleton. + + +2. THE APPENDICULAR SKELETON. + +This consists of the skeleton of the two pairs of limbs and their +respective girdles. It is at first entirely cartilaginous but the +cartilage becomes later on mainly replaced by bone. The only bone +developed in connection with the appendicular skeleton, which has no +cartilaginous predecessor, is the _clavicle_. + + +A. THE PECTORAL GIRDLE. + +This consists originally of two half rings of cartilage encircling the +sides of the body a short way behind the head. These two halves meet +one another in the ventral middle line, and separate the anterior +elements of the sternum from the posterior ones. + +Each half-ring bears on the middle of its outer and posterior surface +a prominent cup, the =glenoid cavity=, with which the proximal +arm-bone articulates. This cup divides the half-arch into a dorsal +=scapular= and a ventral =coracoid= portion. + +The =scapular portion= consists of two parts, the =suprascapula= and +the =scapula=. + +The =suprascapula= (fig. 30, A, 2) is a wide, thin plate attached by +its ventral and narrowest border to the scapula. Its proximal and +anterior half is imperfectly ossified, its whole border or sometimes +only its dorsal and posterior borders consist of unaltered hyaline +cartilage, while the rest of it is composed of calcified cartilage. +The =scapula= (fig. 30, A, 3) is a fairly stout rod of bone +constricted in the middle, and forming the dorsal half of the glenoid +cavity. + +The =coracoid portion= consists of three parts, the =coracoid=, +=precoracoid= and _clavicle_. + +The largest and most posterior of these is the _coracoid_ (fig. 30, A, +4) which like the scapula, is contracted in the middle and expanded at +the ends, especially at the ventral end. It forms a large part of the +glenoid cavity. The ventral ends of the coracoids which meet one +another in the middle line are unossified, and form narrow strips of +calcified cartilage, the =epicoracoids= (fig. 30, A, 5); these are +often regarded as sternal elements. + +The =precoracoid= forms a narrow strip of cartilage lying in front of +the coracoid, from which it is separated by the wide =coracoid +foramen= (fig. 30, A, 9). The dorsal end is continuous with an area of +unossified cartilage which separates the coracoid and scapula and +forms part of the glenoid cavity. + +The _clavicle_ is a narrow membrane bone closely attached to the +anterior surface of the precoracoid, its dorsal end is expanded. + + +THE STERNUM. + +The sternum consists of four parts arranged in two groups; two parts +to each group. The anterior members are the episternum and omosternum. + +The =episternum= (fig. 30, A, 10) is a thin almost circular plate of +cartilage much of which remains hyaline. + +The =omosternum= (fig. 30, A, 11) is a slender bony rod widest at its +posterior end; it connects the episternum with the ventral ends of the +precoracoids. + +The =sternum proper= is a short rod of cartilage sheathed in bone; it +is contracted in the middle and expanded at each end. It bears +attached to its posterior end a broad somewhat bilobed plate of +partially calcified cartilage, the =xiphisternum= (fig. 30, A, 13). + +B. THE ANTERIOR LIMB. + +This is divisible into three parts, the =upper arm= or =brachium=, the +=fore-arm= or =antibrachium=, and the =manus=. + +All the larger bones have their ends formed by prominent epiphyses +which do not unite with the shaft till late in life. Their +articulating surfaces are covered by hyaline cartilage. + +In the =upper arm= there is a single bone, the =humerus=. + +This has a more or less cylindrical shaft and articulates by a +prominent rounded =head= with the glenoid cavity. The distal end shows +a large rounded swelling on either side of which is a =condylar +ridge=, the inner or postaxial one being the larger. A prominent +=deltoid ridge= runs along the proximal half of the anterior surface, +and in the male frog a second equally prominent ridge runs along the +distal half of the posterior surface. + +The =fore-arm= consists of two bones, the =radius= and =ulna=, united +together and forming the =radio-ulna=. The two bones are quite fused +at their proximal ends where they form a deep cup which articulates +with the distal end of the humerus, and is drawn out into a rather +prominent backwardly-projecting =olecranon process=, which ossifies +from a centre distinct from that of the shaft. The distal end is +distinctly divided by a groove into an anterior radial and a posterior +ulnar portion. + +The =manus= consists of two parts, the =wrist= or =carpus= and the +=hand=. + +The =carpus=[57] consists of six small bones arranged in two rows. The +three bones of the proximal row are the =ulnare=, =radiale= and +=centrale=. The =ulnare= and =radiale= are about equal in size and +articulate regularly with the radio-ulna. The =centrale= is pushed out +of its normal position and lies partly on the pre-axial side, partly +in front of the radiale. Of the three bones of the distal row the two +pre-axial ones, =carpalia 1= and =2=, are small; carpale 2 articulates +with the second metacarpal, carpale 1 with both the first and second. +The third bone is large and articulates with the third, fourth and +fifth metacarpals, it represents =carpalia 3-5=, with probably in +addition the representative of a second centrale. + +The =hand= consists of four complete digits, and a vestigial =pollex= +reduced to a short metacarpal. + +Each of the four complete digits consists of a =metacarpal= and a +variable number of =phalanges=. The first digit, as just mentioned, +has no phalanges, the second and third have two, and the fourth and +fifth have three. + +C. THE PELVIC GIRDLE. + +The pelvic girdle of the Frog is much modified from the simple or +general type found in the Newt (p. 149). + +It is a V-shaped structure consisting of two halves which are fused +together in the middle line posteriorly, while in front they are +attached to the ends of the transverse processes of the sacral +vertebra. Each half bears at its posterior end a deep cup, the +=acetabulum=, with which the head of the femur articulates. + +Each half of the pelvis ossifies from two centres. The anterior and +upper half of the acetabulum, and the long laterally compressed bar +extending forwards to meet the sacral vertebra ossify from a single +centre and are generally called the =ilium=; it is probable however +that they represent both the =ilium= and =pubis= of mammals[58]. The +posterior part of this bone meets its fellow in a median symphysis. + +The posterior third of the acetabulum is formed by a small bone, the +=ischium=, which likewise meets its fellow in a median symphysis. + +The ventral portion of the pelvic girdle never ossifies, even in old +animals being formed only of calcified cartilage. This is generally +regarded as the pubis, but it perhaps corresponds to the =acetabular +bone= of mammals. + + +D. THE POSTERIOR LIMB. + +This corresponds closely to the anterior limb and, like it, is +divisible into three parts, the =thigh=, the =shin= or =crus= and the +=pes=. + +As was the case with the anterior limb, all the long bones have their +ends formed by prominent epiphyses which do not unite with the shaft +till late in life. + +In the =thigh= there is only a single bone, the =femur=. + +The =femur= is a moderately long, slender bone with a well-ossified +hollow shaft slightly curved in a sigmoid manner. Both ends are +expanded, the proximal end is hemispherical and articulates with the +acetabulum, the distal end is larger and more laterally expanded. + +The =shin= likewise includes a single bone, the =tibio-fibula=, but +this, as can be readily seen by the grooves at the proximal and distal +ends of the shaft, is formed by the fusion of two distinct bones, the +=tibia= and =fibula=. The tibio-fibula is longer and straighter than +the femur. + +The =pes= consists of two parts, the =ankle= or =tarsus= and the +=foot=. + +The =tarsus= consists of two rows of structures, very different in +size. The proximal row consists of two long bones, the =tibiale= and +=fibulare=, which are united by common epiphyses at the two ends, +while in the middle they are widely separated. The tibiale lies on the +tibial or pre-axial side, and the fibulare which is the larger of the +two bones on the fibular or postaxial side. The distal row of tarsals +consists of three very small pieces of calcified cartilage. The +postaxial of these is the largest, it articulates with the second and +third metatarsals and is probably homologous with tarsalia 2 and 3 +fused. The middle one is very small, it articulates with the first +metatarsal and is probably tarsale 1. The pre-axial one articulates +with the metatarsal of the calcar, a structure to be described +immediately, and has been regarded as a =centrale=. + +The =foot= includes five complete digits and a supplemental toe as +well. Each of the five digits consists of a long =metatarsal= with +epiphyses at both ends, and of a variable number of phalanges. The +first digit or =hallux= and the second have two phalanges, the third +three, the fourth, which is the largest, four, and the fifth, three. +The distal phalanges have epiphyses only at their proximal ends, the +others at both ends. + +On the pre-axial side of the hallux is the supplemental digit, the +=prehallux= or =calcar=. It consists of a short metatarsal and one or +two phalanges, and is terminated distally by a horny covering of +epidermal origin. + + +FOOTNOTES: + +[55] See A. Ecker, _Die anatomie des Frosches_, Braunschweig 1864, +translated by G. Haslam, Oxford, 1889, also A.M. Marshall, _The Frog_, +5th edition, Manchester and London, 1894. + +[56] W.K. Parker, _Phil. Trans._ 161, 1871, p. 137, and W.K. Parker +and G.T. Bettany, _The Morphology of the Skull_, London, 1877, p. 136. + +[57] See G.B. Howes and W. Ridewood, _P.Z.S._, 1888, p. 141. + +[58] See bottom of p. 187. + + + + +CHAPTER XII. + +GENERAL ACCOUNT OF THE SKELETON IN AMPHIBIA. + + +EXOSKELETON. + +The exoskeleton, at any rate in most living forms, is very slightly +developed in Amphibia. The only representatives of the epidermal +exoskeleton are (1) the minute horny beaks found coating the +premaxillae and dentaries in _Siren_ and the tadpoles of most Anura, +(2) the nails borne by the first three digits of the pes in _Xenopus_ +and by the Japanese Salamander _Onychodactylus_, (3) the horny +covering of the calcar or prehallux of frogs. The Urodela and nearly +all the Anura, which form the vast majority of living Amphibia, have +naked skins. A few Anura belonging to the genera _Ceratophrys_ and +_Brachycephalus_ have bony dermal plates developed in the skin of the +back, and these plates become united with some of the underlying +vertebrae. + +In the Gymnophiona the integument bears small cycloid scales arranged +in rings which are equal in number to the vertebrae. These scales +contain calcareous concretions. Scales also occur between the +successive rings. + +In the Labyrinthodontia the dermal exoskeleton is in many genera +greatly developed. It is generally limited to the ventral surface and +consists principally of a buckler formed of three bony plates, one +median and two lateral. These plates protect the anterior part of the +thorax, and are closely connected with the adjacent endoskeleton. They +probably represent the interclavicle and clavicles. Behind this +buckler numerous scutes are generally developed, which often cover the +whole ventral surface, and may cover the whole body. + + +TEETH[59]. + +In Amphibia teeth are generally present on the maxillae, premaxillae and +vomers, and except in Anura on the dentaries; sometimes they occur on +the palatines as in many Urodela, most Labyrinthodontia, and the +Gymnophiona; less commonly on the pterygoids as in _Menobranchus_, +_Siredon_, some Labyrinthodontia, and _Pelobates cultripes_[60], or on +the splenials as in _Siren_ and _Menobranchus_, or parasphenoid as in +_Pelobates cultripes_, _Spelerpes belli_ and _Batrachoseps_. In some +Anura such as _Bufo_ and _Pipa_ the jaws are toothless. + +In Gymnophiona, _Menobranchus_, and _Siredon_, the teeth are arranged +in two concentric curved rows. The teeth of the outer row are borne on +the premaxillae and maxillae if present, (the maxillae are absent in +_Menobranchus_), the teeth of the second row on the vomers and +pterygoids in _Menobranchus_ and _Siredon_, and on the vomers and +palatines in Gymnophiona. In some Gymnophiona there is a double row of +mandibular teeth. The vomerine, palatine and parasphenoid teeth of all +forms are numerous and are not arranged in rows. + +The teeth of all living Amphibia are simple conical structures +ankylosed to the bone, and consisting of dentine, coated or capped +with a thin layer of enamel. In the Labyrinthodontia teeth of more +than one size are sometimes present. The dentine of the basal part of +the larger teeth is in some genera very greatly folded, causing the +structure to be highly complicated. These folds, the intervals between +which are filled with cement, radiate inwards from the exterior and +outwards from the large pulp cavity. The basal part of the teeth of +_Ceratophrys_ (Anura) has a similar structure. + + +ENDOSKELETON. + +VERTEBRAL COLUMN. + +Four regions of the vertebral column can generally be recognised in +Amphibia, viz. the cervical, the trunk or thoraco-lumbar, the sacral +and the caudal regions. In the limbless Gymnophiona, however, only +three regions, the cervical, thoracic, and post-thoracic can be made +out. The cervical region is limited to a single vertebra which +generally differs from the others in having no transverse processes or +indication of ribs. It is generally called the atlas, but it commonly +bears a small process arising from the anterior face of the centrum +which resembles the odontoid process of higher animals, and renders it +probable that the first vertebra of Amphibia corresponds to the axis, +not to the atlas. Amphibia generally have a single sacral vertebra. + +Three elements go to make up the vertebral column in Amphibia, viz. + +1. the notochord, + +2. the long vertebral centra, + +3. intervertebral cartilage which forms the joints between successive +centra. + +The relations which these three elements bear to one another are +subject to much variation. The successive stages can be well traced in +the Urodela. + +1. The first stage is found in larval Urodeles in general and in adult +Ichthyoidea, and some Salamandrina. In these forms the notochord +persists and retains approximately the same diameter throughout the +whole length of the vertebral column. Bony biconcave centra are +present and constrict it to a certain extent vertebrally, while +intervertebrally there is a development of cartilage. The connection +between the bony vertebrae is effected mainly by the expanded +notochord. + +2. In the next stage, as seen in _Gyrinophilus porphyriticus_, the +growth of intervertebral cartilage has caused the almost complete +obliteration of the notochord intervertebrally, and its entire +disappearance vertebrally, i.e. in the centre of each vertebra. The +intervertebral cartilage now forms the main connection between +successive vertebrae, and sometimes cases are found whose condition +approaches that of definite articulations. Readily recognisable +remains of the notochord are still found at each end of the +intervertebral constriction. + +3. In the third stage differentiation and absorption of the +intervertebral cartilage has given rise to definitely articulating +opisthocoelous vertebrae. These are found in most adult Salamandrina. + + * * * * * + +The transverse processes of the earlier trunk vertebrae are divided +into two parts, a dorsal part which meets the tubercular process of +the rib and is derived from the neural arch, and a ventral part which +meets the capitular process of the rib, and is derived from the +ventral or haemal arch. In the caudal vertebrae and often also in the +posterior trunk vertebrae the two processes are fused. + +_Siren_ and _Proteus_, although they possess minute posterior limbs, +have no sacral vertebrae, while _Cryptobranchus lateralis_ has two. +The caudal vertebrae, except the first, have haemal arches very +similar to the neural arches. + +In Labyrinthodontia the centra of the vertebrae are generally well +ossified biconcave discs. In some forms however, like _Euchirosaurus_, +the centra are imperfectly ossified, and consist of bony rings +traversed by a wide notochordal canal. Each ring is formed of four +pieces, a large well-ossified neural arch, a basal piece, and a pair +of lateral pieces. Vertebrae of this type are called _rachitomous_. + +In the tail region of other forms each vertebra consists of an +anterior centrum bearing the neural arch, and a posterior +intercentrum[61] bearing chevron bones. Vertebrae of this type are +called _embolomerous_. Haemal arches similar to the neural arches are +often found as in Urodela. The transverse processes are sometimes well +developed and are divided into tubercular and capitular portions. + +In Gymnophiona the vertebrae are biconcave and are very numerous, they +sometimes number about two hundred and thirty. Only quite the last few +are ribless and so can be regarded as post-thoracic vertebrae. The +first vertebra has nothing of the nature of an odontoid process. + +In Anura the number of vertebrae is very greatly reduced, only nine +and the urostyle being present. Of these, eight are presacral and one +sacral. The urostyle is post-sacral and corresponds to three or more +modified vertebrae. The first vertebra is without transverse +processes, the remaining presacral vertebrae have the transverse +processes fairly large, while the sacral vertebra has them very large, +forming in some genera widely expanded plates. The urostyle is a long +cylindrical rod which articulates with the sacrum generally by two +facets. Ankylosed to its anterior end are the remains of two neural +arches. + +In Anura remains of the notochord are found in the centre of each +vertebra, i.e. vertebrally, while in the Urodela they only occur +intervertebrally. + +The vertebrae in Anura are, as a rule, procoelous. The eighth vertebra +is however generally amphicoelous, while the ninth commonly has one +convexity in front, and two behind. + +In some forms such as _Bombinator_, _Pipa_, _Discoglossus_ and +_Alytes_ they are opisthocoelous; in others like _Pelobates_ they are +variable. + + +THE SKULL[62]. + +CRANIUM AND MANDIBLE. + +In the Amphibian skull there are as a rule far fewer bones than in the +skull of bony fish. The primordial cartilaginous cranium often +persists to a great extent. Only quite a few ossifications take place +in it; namely in the occipital region--the exoccipitals, further +forwards--the pro-otics, and at the boundary of the orbital and +ethmoidal regions--the sphenethmoid. The basi-occipital and +basisphenoid are never ossified. As in Mammalia there are two +occipital condyles formed by the exoccipitals. + +Large vacuities commonly occur in the cartilage of both floor and roof +of the primordial cranium. These are roofed over to a greater or less +extent by the development of membrane bone. Thus on the roof of the +cranium there are paired parietals, frontals, and nasals, and on its +floor are paired vomers, and a median unpaired parasphenoid. + +In all living forms the parietals meet and there is no interparietal +foramen, though this exists in Labyrinthodonts. + +The palato-pterygo-quadrate bar is united at each end with the +cranium, but elsewhere in most cases forms a wide arch standing away +from it. The suspensorium is, as in Dipnoi and Holocephali, +autostylic. The palato-pterygo-quadrate bar sometimes remains entirely +cartilaginous, sometimes its posterior half is ossified forming the +quadrate. In connection with it a number of membrane bones are +generally developed, viz. the maxillae, premaxillae, palatines, +pterygoids, quadratojugals, and squamosals. The pterygoids are, +however, sometimes partially formed by the ossification of cartilage. +The cartilage of the lower jaw and its investing membrane bones +generally have much the same relations as in bony fishes. + +URODELA. The skulls of the various Urodeles show an interesting series +of modifications and differ much from one another, but all agree in +the absence of the quadratojugals, in the fact that the palatines lie +parallel to the axis of the cranium, and in the large size of the +parasphenoid. + +The lower types _Menobranchus_, _Siren_, _Proteus_, and _Amphiuma_ +have longer and narrower skulls than do the higher types. + +_Menobranchus_ has a very low type of skull which remains throughout +life in much the same condition as that of a young tadpole or larval +salamander. The roof and floor of the cranium internal to the membrane +bones are formed of fibrous tissue, not of well-developed cartilage. +The epi-otic regions of the skull are ossified, forming a pair of +large bones which lie external to, and distinct from, the +exoccipitals. _Proteus_ and the Labyrinthodonts are the only other +Amphibia which have these elements separately ossified. The parietals +send a pair of long processes forwards along the sides of the +frontals. Nasals and maxillae are absent, as is likewise the case in +_Proteus_. Teeth are borne on the vomers, premaxillae, pterygoids, +dentaries and angulo-splenials. The suspensorium is forwardly +directed. + +The skull of _Siren_ resembles that of _Menobranchus_ in several +respects, as in the forward direction of the suspensorium and in the +absence of maxillae, but differs in the possession of nasals, in the +toothless condition of the premaxillae and dentaries, and in the fusion +and dentigerous condition of the vomers and palatines. + +_Amphiuma_ has a skull which, though narrow and elongated, differs +from those of _Menobranchus_, _Proteus_, and _Siren_, and resembles +those of higher types in the following respects:-- + +(1) the suspensorium projects nearly at right angles to the cranium +instead of being directed forwards, (2) the maxillae are well +developed, and the premaxillae are completely ankylosed together, (3) +there are no palatines. + +The skulls of _Megalobatrachus_, _Cryptobranchus_ and _Siredon_ +resemble those of the highest Urodeles the Salamanders in their wide +form, in having the pro-otics distinct from the exoccipitals which are +ossified continuously with the epi-otics and opisthotics, and in +having no palatines, but differ in having the two premaxillae +separate, and in the arrangement of the vomerine teeth which in +_Megalobatrachus_ and _Cryptobranchus_ are placed along the anterior +boundaries of the bones, these meeting in the middle line. In +_Siredon_ the vomers are separated by the very large parasphenoid. + +The suspensorium in _Megalobatrachus_ and _Cryptobranchus_ projects at +right angles to the cranium; in _Siredon_ it projects somewhat +downwards and forwards as in the Salamandrina. + +Modifications of the vomers, pterygoids and palatines accompany the +changes of the larval ichthyoid _Siredon_ into the adult salamandroid +_Amblystoma_, the vomers especially come to resemble to a much greater +extent those of the Salamandrina. + +The ossification of the skull in the Salamandrina is carried further +than in the Ichthyoidea, though the supra-occipital and basi-occipital +are not ossified. The skull differs from that in the Ichthyoidea in +the size of the part of the vomero-palatines which lies in front of +the teeth, in the frequent union of the two premaxillae and in the +ossification of all the periotic bones continuously with the +exoccipital. + +The skull differs from that of Anura in the following respects:-- + +(1) the bones of the upper jaw do not form a complete arch standing +away from the cranium, and the maxillae are not united to the quadrates +by quadratojugals, (2) the long axis of the suspensorium passes +obliquely downwards and forwards instead of downwards and backwards, +(3) there is no sphenethmoid encircling the anterior end of the +brain, its place being partly taken by a pair of orbitosphenoids, (4) +there is no definite tympanic cavity. + +[Illustration FIG. 27. DORSAL VIEW OF THE SKULL OF A LABYRINTHODONT +(_Capitosaurus nasutus_) × 1/9 (from VON ZITTEL). + + 1. premaxillae. + 2. nasal. + 3. maxillae. + 4. anterior nares. + 5. frontal. + 6. prefrontal. + 7. lachrymal. + 8. jugal. + 9. orbit. + 10. parietal. + 11. postfrontal. + 12. postorbital. + 13. interparietal foramen. + 14. squamosal. + 15. supratemporal. + 16. quadratojugal. + 17. quadrate. + 18. epi-otic. + 19. dermo-supra-occipital. + 20. exoccipital. + 21. foramen magnum.] + +LABYRINTHODONTIA. The skull in Labyrinthodontia is remarkable for its +extreme solidity, the large number of bones which are present, and the +extent to which the roofing over of the temporal and other fossae has +taken place. In many forms the surface of the bones is as in +Crocodiles, strongly sculptured (fig. 27, right half) with ridges and +grooves which probably lodged sensory organs. The bones forming the +roof of the skull are generally very uniform in size, perhaps the most +noticeable of them being the paired dermo-supra-occipitals (fig. 27, +19). Paired dermo-supra-occipitals occur also in certain Ganoids. The +Labyrinthodont skull also bears resemblance to that of many fish in +the development of a pair of long pointed epi-otics (fig. 27, 18), +which remain permanently distinct from the surrounding bones. The +parietals are small and enclose between them the interparietal foramen +(fig. 27, 13). In some forms in which the head is protected with an +armour of scutes, these do not roof over the interparietal foramen, +and from this fact it has been inferred that the Labyrinthodonts had a +functional pineal eye. Both supra- and infra-temporal fossae are +partially or completely roofed over by the postorbitals and large +supra-temporals (fig. 27, 15). + +There is generally a ring of bones in the sclerotic coat of the eye. +The pterygoids do not meet in the middle line, being separated by the +parasphenoid. The palatines bear teeth, and in some genera +(_Archegosaurus_) form long splints lying along the inner side of the +maxillae and more or less surrounding the posterior nares. In others +(_Nyrania_) they lie in the normal position near the middle line, one +on each side of the parasphenoid. The vomers bear teeth and sometimes +meet in the middle line; they are sometimes confluent with the +parasphenoid. On the ventral surface of the cranium there are +generally large palatal vacuities. + +In the mandible there is often a well-marked postglenoid process, and +the articular is generally completely ossified. + +[Illustration FIG. 28. A, VENTRAL VIEW OF THE CRANIUM; B, LATERAL VIEW +OF THE CRANIUM AND MANDIBLE OF _Siphonops annulatus_ (after +WIEDERSHEIM). + + 1. anterior nares. + 2. naso-premaxillae. + 3. frontal. + 4. parietal. + 5. maxillae. + 6. vomer. + 7. orbit. + 8. quadrate united with the pterygoid in front. + 9. squamosal. + 10. exoccipital. + 11. dentary. + 12. angular. + 13. basi-occipital and basisphenoid fused. + 14. posterior narial opening surrounded by the palatine. + X. pneumogastric foramen.] + +GYMNOPHIONA. The skull bears a considerable resemblance to that of +Labyrinthodonts, especially in the arrangement of the bones which +bound the mouth cavity. The cranium is very hard, and is covered by +a complete bony roof formed mainly of the exoccipitals, parietals, +frontals, prefrontals, nasals and premaxillae. The nasals and +premaxillae are sometimes ossified continuously. There is a median +unpaired ethmoid whose dorsal end appears at the surface wedged in +between the frontals and parietals. The bone generally regarded as the +squamosal[63] is very large, and it and the maxillae generally together +surround the orbit, which, in _Epicrium_, has in it a ring of bones. +The palatines form long tooth-bearing bones fused with the inner sides +of the maxillae; they nearly surround the posterior nares. + +The quadrate bears the knob, and the angular the cup for the +articulation of the mandible,--a very primitive feature. The mandible +is also noticeable for the enormous backward projection of the +angular. + + * * * * * + +ANURA. In Anura the skull is very short and wide owing to the +transverse position of the suspensorium. There is often a small +ossification representing the quadrate. Sometimes as in _Hyla_ and +_Alytes_ there is a fronto-parietal fontanelle. + +As compared with the skull in Urodela the chief characteristics of the +skull of Anura are:-- + +1. the presence of a sphenethmoid, + +2. the union of the frontals and parietals on each side, + +3. the occasional occurrence of small supra- and basi-occipitals, + +4. the backward growth of the maxillae and its connection with the +suspensorium by means of the quadratojugal, + +5. the dagger-like shape of the parasphenoid, + +6. the occurrence of a definite tympanic cavity, + +7. the frequent occurrence of a predentary or mento-meckelian +ossification in the mandible. + +The skull of _Pipa_ is abnormal, being greatly flattened and +containing little cartilage. The fronto-parietals are fused, and there +is no sphenethmoid. The quadrates are well developed and the +squamosals and parasphenoid differ much from those of other Anura. + + +HYOID AND BRANCHIAL ARCHES. + +In larval Amphibia the hyoid and four branchial arches are generally +present, and in adult Ichthyoidea they are frequently almost as well +represented as in the larva, and are of use in strengthening the +swallowing apparatus. They are very well seen in _Siredon_, and +consist of a hyoid attached by ligaments to the suspensorium, followed +by four branchial arches of which the first and second are united by a +copula (fig. 29, D, 8), while the third and fourth are not. The hyoid +is not always the largest and best preserved of the arches, for +sometimes as in _Spelerpes_ one of the branchials is far larger than +the hyoid. Four branchial arches occur in _Siren_ as in _Siredon_, but +in _Proteus_ there are only three. + +In some larval Labyrinthodontia (_Branchiosaurus_) four branchial +arches are known to occur, and their arrangement is almost precisely +similar to that in _Siredon_. + +In Gymnophiona the remains of only three branchial arches occur in +addition to the hyoid. The four arches are all very similar to one +another, each consists of a curved rod of uniform diameter throughout. +The hyoid is united with the first branchial arch, but has no +attachment to the cranium. + +In larval Anura (fig. 29, C) the arrangement of the hyoid and +branchial arches is much as in Urodela. In the adult, however, the +ventral parts of all the arches unite, forming a compact structure, +the _basilingual plate_ (fig. 29, B, 1). + +[Illustration FIG. 29. VISCERAL ARCHES OF AMPHIBIA. + + A. _Molge cristata_ (after PARKER). + B. _Rana temporaria_ adult (after PARKER). + C. Tadpole of _Rana_ (after MARTIN ST ANGE). + D. _Siredon pisciformis_ (after CREDNER). + +In each case the ossified portions are slightly shaded, while the +cartilaginous portions are left white. + + 1. basilingual plate. + 2. hyoid arch. + 3. first branchial arch. + 4. second do. + 5. third branchial arch. + 6. fourth do. + 7. thyro-hyal. + 8. copula.] + +The dorsal parts of the first three branchial arches disappear, but +those of the fourth become ossified and form the short, stout +thyro-hyals or posterior cornua. The dorsal parts of the hyoid arch in +the adult form a pair of long bars, the anterior cornua, which are +united to the periotic region of the skull in front of the fenestra +ovalis either by short ligaments or by fusion as in _Bufo_. In _Pipa_ +and _Xenopus_ the first and second branchial arches persist as well as +the fourth (thyro-hyal), but in _Pipa_ the hyoid is wanting. + + +RIBS. + +Ribs are generally very poorly developed in Amphibia. In Anura they +are in most cases absent; when present they generally form minute +unossified appendages attached to the transverse processes, but in +_Discoglossus_ and _Xenopus_ the anterior vertebrae are provided with +distinct ribs. In Urodela and Labyrinthodontia they are generally +short structures, each as a rule attached to the vertebra by a +bifurcated proximal end. The number of rib-bearing vertebrae varies, +but the first and the posterior caudal vertebrae are always ribless. +The anterior caudal vertebrae too are generally ribless, but sometimes +a few of them bear small ribs. In _Spelerpes_ the last two trunk +vertebrae are ribless, and hence may be regarded as lumbar vertebrae. + +In Gymnophiona ribs are better developed than in any other Amphibia; +they occur on all the vertebrae except the first and last few, and are +attached to the transverse processes, sometimes by single, sometimes +by double heads. + +Sternal ribs are almost unknown in Amphibia, but traces of them occur +in _Menobranchus._ + + +STERNUM. + +In Amphibia the sternum is not very well developed; sometimes as in +Gymnophiona and _Proteus_ no traces of it occur, and in the Urodela it +is never ossified. It is always very intimately related to the +pectoral girdle. In the Salamandrina it has the form of a broad thin +plate of cartilage, grooved and overlapped by the coracoid. + +[Illustration FIG. 30. SHOULDER-GIRDLE AND STERNUM OF + + A. An old male common Frog (_Rana temporaria_). + B. An adult female _Docidophryne gigantea_ (after PARKER). + +In both A and B the left suprascapula is removed. The parts left +unshaded are ossified; those marked with small dots consist of hyaline +cartilage, those marked with large dots of calcified cartilage. + + 1. calcified cartilage of suprascapula. + 2. ossified portion of suprascapula. + 3. scapula. + 4. coracoid. + 5. epicoracoid. + 6. precoracoid. + 7. clavicle. + 8. glenoid cavity. + 9. coracoid foramen. + 10. episternum. + 11. omosternum. + 12. sternum. + 13. xiphisternum.] + +In most Anura the sternum consists of a number of parts arranged in +series. At the anterior end is a flat cartilaginous plate with a +bony basal stalk. This plate is called the episternum, and its stalk +the omosternum. The continuity of the sternum is now interrupted +by a pair of cartilaginous structures, the epicoracoids, which are +shoulder-girdle elements, and represent the unossified ventral ends of +the coracoids. In some cases cartilaginous epiprecoracoids can also +be distinguished. Further back is the long sternum proper, while last +comes the xiphisternum, a broad expanded plate of cartilage. + +In some Anura such as _Pipa_ and _Hyla_ the number of sternal elements +is considerably reduced. + + +APPENDICULAR SKELETON. + + +PECTORAL GIRDLE. + +The most primitive Amphibian shoulder-girdle is found in the Urodela. +It consists of a dorsal element, the scapula, a posterior ventral +element, the coracoid, and an anterior ventral element, the +precoracoid. The clavicle is not developed, and the two coracoids +overlap in the middle line. The shoulder-girdle remains largely +cartilaginous but the proximal end of the scapula is ossified, and the +ossification may extend through part of the coracoid and precoracoid. + +In Labyrinthodontia there is an exoskeletal ventral buckler formed of +three plates, a median one, which probably represents an +interclavicle, and two lateral ones, which are probably clavicles. +Traces of endoskeletal structures, probably corresponding to the +precoracoid and scapula, are also known in some cases. The Gymnophiona +and some of the Labyrinthodontia have lost the pectoral girdle and +limbs. + +The ossification of the shoulder-girdle has gone on much further in +Anura than it has in Urodela. Clavicles are present and the scapula +and coracoid of each side are ossified from separate centres. The +distal part of the scapula forms a large imperfectly ossified plate, +the suprascapula. + +The shoulder-girdle of Anura is however subject to considerable +variations. In the Toads (Bufonidae) the epicoracoids or unossified +ventral ends of the coracoids and precoracoids overlap in the middle +line (fig. 30, B, 5). This arrangement is called _Arciferous_. In the +Frogs,--Ranidae, and other forms belonging to the group +_Firmisternia_,--the epicoracoids do not overlap but form a narrow +cartilaginous bar separating the ventral ends of the coracoids (fig. +30, A, 5). + + +ANTERIOR LIMB. + +In many Amphibia and especially in the Urodela the anterior limb has a +very simple unmodified arrangement. The humerus is straight and of +moderate length, its ends are rounded for articulation on the one hand +with the shoulder-girdle, and on the other hand with the radius and +ulna. In the Urodela the radius and ulna are distinct. In the Anura +they have fused, though the line of junction of the two is not +obliterated. Their proximal ends are hollowed for articulation with +the convex end of the humerus. + +The manus in all recent Amphibia agrees in never having more than four +complete digits, but is subject to considerable variation, this +statement applying especially to the carpus. + +In the larva of Salamandra (fig. 31, A), except that the pollex is +absent[64], the manus retains completely the condition which is +generally regarded as primitive for the higher Vertebrata. It consists +of an anterior row of three elements, the ulnare, intermedium, and +radiale, and a posterior row of four, the carpalia 2, 3, 4, and 5. +Interposed between the two rows is a centrale. _Menobranchus_ has a +similar very simple carpus. In most other Amphibia this simplicity is +lost. This loss may be due to:-- + +(_a_) fusion of certain structures, e.g. in the adult _Salamandra_ the +intermedium and ulnare have fused, + +(_b_) displacement of structures, e.g. in _Bufo viridis_, the centrale +has been pushed up till it comes to articulate with the radius, + +(_c_) the development of supernumerary elements, especially of extra +centralia. In _Megalobatrachus_ two or even three centralia sometimes +occur. + +[Illustration FIG. 31. A, RIGHT ANTIBRACHIUM AND MANUS OF A LARVAL +SALAMANDER (_Salamandra maculosa_) (after GEGENBAUR). + +B, RIGHT TARSUS AND ADJOINING BONES OF _Molge sp._ (after GEGENBAUR). + + 1. radius. + 2. ulna. + 3. radiale. + 4. intermedium. + 5. ulnare. + 6. centrale. + 7. carpale 2. + 8. " 3. + 9. " 4. + 10. " 5. + 11. tibia. + 12. fibula. + 13. tibiale. + 14. intermedium. + 15. fibulare. + 16. centrale. + 17. tarsale 1. + 18. tarsalia 4 and 5 fused. + I. II. III. IV. V. digits.] + +In the great majority of Amphibia while one digit, probably the first, +is absent, the other four digits are well developed. In the forms +however with degenerate limbs like _Amphiuma_, _Siren_ and _Proteus_ +the number of digits is still further reduced. In _Siren_ there are +three or four, in _Proteus_ three, and in _Amphiuma_ two or three +digits in the manus. + +In Anura the pollex is represented only by a short metacarpal. There +are sometimes traces of a prepollex. The carpus often has two +centralia and the intermedium is absent. + +In Labyrinthodontia the limbs are generally very simple and resemble +those of Urodela. In some forms, however, the manus differs from that +of all living Amphibia in possessing five well-developed digits. + + +PELVIC GIRDLE. + +The simplest Amphibian pelvis is that of some of the Labyrinthodontia; +thus in _Mastodonsaurus_ it consists dorsally of a short broad ilium +placed vertically and attached to the sacrum, and ventrally of a small +pubis and of a large ischium meeting its fellow in the middle line. In +some Labyrinthodonts the pubes as well as the ischia meet in a ventral +symphysis, and in many there are no obturator foramina. In _Siren_, +Gymnophiona and some Labyrinthodontia the pelvic girdle and limbs are +absent. + +In Urodela the ventral element of the pelvis on each side forms a flat +plate which meets its fellow of the opposite side. The anterior part +of the plate, representing the pubis, generally remains cartilaginous +throughout life; the posterior part representing the ischium is in +almost every case well ossified. Attached to the anterior end of the +pubes there is an unpaired bifid cartilaginous structure, the +epipubis. The ilia are vertically placed. + +In most Anura the pelvis is peculiarly modified in correlation with +the habits of jumping. The long bone generally called the ilium is +placed horizontally and is attached at its extreme anterior end to the +sacrum. The ischium is ossified and distinct. Ventrally in front of +the ischium there is a tract of unossified cartilage which is often +regarded as the pubis. In _Xenopus_, however, the bone corresponding +to the ilium of the Frog is seen to ossify from two centres, one +forming the ilium, the other, which lies at the symphysis, being +apparently the pubis. This makes it probable that the so-called ilium +of the Frog is really to be regarded as an ilio-pubis, and renders the +homology of the cartilaginous part uncertain, but it probably +corresponds to the acetabular bone of mammals. In _Xenopus_ also there +is a minute epipubis similar to that of Urodeles. + + +POSTERIOR LIMB. + +In Urodela the posterior limb (fig. 31, B) closely resembles the +anterior limb, but is even less removed from the primitive condition +of the higher vertebrates in the fact that all five digits are +commonly present. The tibia and fibula are short bones approximately +equal in size. In some cases the number of digits is reduced. Thus in +_Menobranchus_ the pes has four digits, in _Proteus_ it has two, and +in _Amphiuma_ two or three, while in _Siren_ the posterior limbs have +atrophied. + +In correlation with their habits of jumping, the posterior limbs in +Anura are much lengthened and considerably modified. The tibia and +fibula are completely fused. The intermedium is absent, while the +tibiale and fibulare are greatly elongated. Tarsalia 4 and 5 are +absent. Five digits are always present, and there is a prehallux +formed of two or more segments. + +In general the posterior limbs in Labyrinthodontia bear the closest +resemblance to the anterior limbs; in some cases three centralia are +found. + +In Ichthyoidea, and in most Labyrinthodontia, the cartilages of the +carpus and tarsus remain unossified; in Salamandrina and in Anura they +are generally ossified. + + +FOOTNOTES: + +[59] O. Hertwig. Ueber das Zahnsystem der Amphibien. _Arch. mikr. +Anat._ supplem. Bd. XI. 1875. + +[60] G.A. Boulenger, _P.Z.S._ 1890, p. 664. + +[61] See p. 14. + +[62] See many papers by W.K. Parker published in the _Phil. Trans._ of +the Royal Soc. + +[63] Perhaps this bone includes supra-orbital and postorbital +elements. + +[64] The first digit present is sometimes regarded as the pollex, but +from analogy with Anura it is probable that the pollex is the missing +digit. + + + + +CHAPTER XIII. + +SAUROPSIDA. + + +This great group includes the Reptiles and Birds and forms the second +of the three into which the Gnathostomata may be divided. There is +nearly always a strongly-developed epiblastic exoskeleton which has +the form of scales or feathers, and in some cases a dermal exoskeleton +is also well developed. In living forms the notochord never persists, +being replaced by vertebrae, but in some extinct forms the centra are +notochordal. The vertebral centra are ossified, and only in +exceptionally rare cases have terminal epiphyses. The skull is well +ossified and has membrane bones incorporated in its walls. + +The occipital segment is completely ossified, and an interorbital +septum or bony partition separating the two orbits is usually +developed to a greater or less extent. The skull generally articulates +with the vertebral column by a single occipital condyle into the +composition of which the exoccipitals and basi-occipital enter in +varying proportions. The pro-otic ossifies, and either remains +distinct from the epi-otic[65] and opisthotic throughout life, or +unites with them only after they have fused with the adjacent bones. +The hyoid and branchial arches are much reduced; and the +representative of the hyomandibular is connected with the auditory +apparatus, forming the auditory ossicles[66]. Each ramus of the +mandible always consists of a cartilage bone, the articular, and +several membrane bones. The mandible articulates with the cranium by +means of a quadrate. The ribs in Birds and some Reptiles bear +_uncinate processes_, i.e. small, flat, bony or cartilaginous plates +projecting backwards from their posterior borders. The sternum is not +transversely segmented as in mammals, and there are commonly distinct +cervical ribs. The ankle joint is intertarsal, or situated between the +proximal and distal row of tarsal bones, not cruro-tarsal as in +Mammalia. + + +CLASS I. REPTILIA[67]. + +The axial skeleton is generally long, and that of the limbs frequently +comparatively short, or sometimes absent. + +The exoskeleton generally has the form of epidermal scales, which are +often combined with underlying bony dermal plates or scutes and may +sometimes form a continuous armour. Neither feathers nor true hairs +are ever present. The vertebral column is generally divisible into the +five usual regions. The centra of the vertebrae vary enormously, and +may be amphicoelous, procoelous, opisthocoelous or flat, but they +never have saddle-shaped articulating surfaces. The quadrate is always +large, and is sometimes fixed, sometimes movable. A transpalatine bone +uniting the pterygoid and maxillae is generally present. + +Free ribs are often borne along almost the whole length of the trunk +and tail, and often occur attached to the cervical vertebrae. The +sacrum is generally composed of two vertebrae which are united with +the ilia by means of expanded ribs. The sternum is rhomboidal, and may +either be cartilaginous or formed of cartilage bone, but never of +membrane bone; it differs from that of birds also in the fact that it +does not ossify from two or more centres. An interclavicle is +generally present. There are always more than three digits in the +manus, and never less than three in the pes. In all living reptiles +the ilia are prolonged further behind the acetabula than in front of +them, and the bones of the pelvis remain as a rule, distinct from one +another throughout life. + +The pubes (pre-pubes) and ischia both commonly meet in ventral +symphysis, and the acetabula are wholly or almost wholly ossified. The +metatarsals are not ankylosed together. + + +_Order 1._ THEROMORPHA[68]. + +This order includes a number of mainly terrestrial, extinct reptiles, +which differ much from one another, and show remarkable points of +affinity on the one hand with the Labyrinthodont Amphibia, and on the +other with the Mammalia. The vertebrae are nearly always amphicoelous +and sometimes have notochordal centra. The skull is short and has the +quadrate immovably fixed. There is an interparietal foramen, and +generally large supratemporal fossae bounded by supratemporal[2] +arcades, but with no infratemporal[69] arcades; _Elginia_ however has +the whole of the temporal region completely roofed over. + +The teeth are placed in distinct sockets and are very variable in +form, the dentition sometimes resembling the heterodont dentition of +mammals. The humerus has distinct condyles and an ent-epicondylar +foramen[70] as in many mammals. + +The pubis is fused with the ischium, and both pectoral and pelvic +girdles are remarkably solid. The obturator foramen is remarkably +small or even absent. The anterior ribs have two articulating +surfaces, and each articulates by its tuberculum with the transverse +process, and by its capitulum with the centrum as in mammals. + +These reptiles occur chiefly in deposits of Triassic and Permian age. +Some of the best known genera are _Dicynodon_, _Udenodon_, _Placodus_, +_Pariasaurus_ and _Galesaurus_. They will be noticed in the general +account of the skeleton in reptiles. + + +_Order 2._ SAUROPTERYGIA. + +This order includes a number of extinct marine reptiles, devoid of an +exoskeleton. The tail is short, the trunk long, and the neck in the +most typical forms extremely long. The vertebrae have slightly +biconcave, or nearly flat centra. The skull is relatively small and +has large supratemporal fossae. The teeth are placed in distinct +sockets, and are generally confined to the margins of the jaws; they +are sharp and curved and are coated with grooved enamel. The +premaxillae are large, and there is an interparietal foramen. The +quadrate is firmly united to the cranium. The anterior nares are +separate and are placed somewhat close to the orbits. There is no +ossified sclerotic ring. The palatines and pterygoids meet the vomers, +and more or less completely close the palate, and in some forms, e.g. +_Plesiosaurus_, there is a distinct parasphenoid. Thoracic ribs are +strongly developed and each articulates with its vertebra by a single +head. The cervical vertebrae have well-marked ribs, which articulate +only with the centra, in this respect differing from those of +Crocodiles. The caudal vertebrae bear both ribs and chevron bones, and +abdominal splint-ribs are largely developed. + +In the shoulder-girdle the coracoids are large and meet in a ventral +symphysis; precoracoids and a sternum are apparently absent, but parts +generally regarded[71] as the clavicles and interclavicle are well +developed. In the pelvis, the pubes and ischia meet in a long +symphysis. The limbs are pentedactylate, and in the best known forms, +the Plesiosauridae, form swimming paddles. + +The Sauropterygia occur in beds of Secondary age, and some of the best +known genera are _Plesiosaurus_, _Pliosaurus_ and _Nothosaurus_. + + +_Order 3._ CHELONIA. + +In the Tortoises and Turtles the body is enclosed in a bony box, +formed of the dorsal carapace, and a flat ventral buckler, the +plastron. Except in _Dermochelys_ the carapace is partly formed from +the vertebral column and ribs, partly from dermal bones. Both carapace +and plastron are, except in _Dermochelys_, _Trionyx_ and their allies, +covered with an epidermal exoskeleton of horny plates, which are +regularly arranged, though their outlines do not coincide with those +of the underlying bones. The thoracic vertebrae have no transverse +processes, and are quite immovably fixed, but the cervical and caudal +vertebrae are very freely movable. There are no lumbar vertebrae. The +skull is extremely solid, and frequently has a very complete false +roof. Teeth have been detected in embryos of _Trionyx_ but with this +exception the jaws are toothless, and are encased in horny beaks. The +quadrate is firmly fixed. The facial part of the skull is very short, +and the alisphenoidal and orbitosphenoidal regions are unossified. In +living forms there are no separate nasal bones, while large +prefrontals and postfrontals are developed. There is a comparatively +complete bony palate chiefly formed of the palatines and pterygoids. +The anterior nares are united and placed at the anterior end of the +skull, and the premaxillae are very small. There is no transpalatine +bone and the vomer is unpaired. The dentaries are generally fused +together. + +There are ten pairs of ribs, and each rib has only a single head and +is partially attached to two vertebrae; there are no cervical or +sternal ribs. There is no true sternum. + +The three anterior elements of the plastron are respectively +homologous with the interclavicle and two clavicles of other reptiles, +while the remaining elements of the plastron are probably homologous +with the abdominal ribs of Crocodiles. The pectoral girdle lies within +the ribs, and the precoracoids and coracoids do not meet in ventral +symphyses. The scapula and precoracoid are ossified continuously. The +pubis probably corresponds with the pre-pubis of Dinosaurs. There are +four limbs each with five digits. + +The order includes three suborders:-- + + +_Suborder (1)._ TRIONYCHIA. + +The carapace and plastron have a rough granular surface covered with +skin and without any horny shields. + +The plastron is imperfectly ossified, and marginal bones may be +absent, or if present are confined to the posterior portion of the +carapace. The pelvis is not united to the plastron. The cranium has +not a complete false roof and the head can be drawn back under the +carapace. + +The first three digits of both manus and pes bear claws, and the +fourth digit in each case has more than three phalanges. The most +important genus is _Trionyx_. + + +_Suborder (2)._ CRYPTODIRA. + +The carapace and plastron vary in the extent to which they are +ossified, and except in _Dermochelys_[72] and its allies are covered +by horny plates. Marginal bones are always present. The head can +generally be drawn back under the carapace. The pelvis is not firmly +united to the plastron. The cranium often has a complete false roof, +and in the mandibular articulation the cup is borne by the cranium, +and the knob by the mandible. Among the more important genera are +_Dermochelys_, _Chelone_, and _Testudo_. + + +_Suborder (3)._ PLEURODIRA. + +The carapace and plastron are strongly ossified, and firmly united to +the pelvis. The head and neck can be folded laterally under the +carapace, but cannot be drawn back under it. The cranium has a more or +less complete false roof, and in the mandibular articulation the knob +is borne by the cranium, and the cup by the mandible. _Chelys_ is a +well-known genus. + + +_Order 4._ ICHTHYOSAURIA[73]. + +The order includes a number of large extinct marine reptiles whose +general shape is similar to that of the Cetacea. The skull is +enormously large, and the neck short. The tail is very long, and is +terminated by a large vertically-placed bilobed fin, the vertebral +column running along the lower lobe. The very numerous vertebrae are +short and deeply biconcave. The vertebral column can be divided into +caudal and precaudal regions only, as the ribs which begin at the +anterior part of the neck are continued to the posterior end of the +trunk without being connected with any sternum or sacrum. The precaudal +vertebrae bear two surfaces for the articulation of the ribs, while +in the caudal vertebrae the two surfaces have coalesced. The caudal +region is also distinguished by its chevron bones. The vertebrae have +no transverse processes, and the neural arches are not firmly united +to the centra, and have only traces of zygapophyses. The atlas and +axis are similar to the other vertebrae, but there is a wedge-shaped +intercentrum between the atlas and the skull, and another between +the atlas and the axis. The skull is greatly elongated (fig. 32) and +pointed, mainly owing to the length of the premaxillae. The orbits +are enormous, and there is a ring of bones in the sclerotic (fig. 32, +15). The anterior nares are very small; and are placed far back just +in front of the orbits. There is an interparietal foramen, and the +supratemporal fossae (fig. 32, 9) are very large, while there are no +infratemporal fossae. An epipterygoid occurs. The quadrate is firmly +fixed to the cranium, and there is a large parasphenoid. There are +large prefrontals, but the frontals are very small. The very numerous +teeth are large and conical, and are placed in continuous grooves +without being ankylosed to the bone. They are confined to the jaw-bones. + +[Illustration FIG. 32. LATERAL (BELOW) AND DORSAL (ABOVE) VIEWS OF THE +SKULL OF AN _Ichthyosaurus_. (Modified from Deslongchamps.) + + 1. premaxillae. + 2. maxillae. + 3. nasal. + 4. prefrontal[1]. + 5. frontal. + 6. postfrontal[74]. + 7. anterior nares. + 8. orbit. + 9. supratemporal fossa. + 10. interparietal foramen. + 11. parietal. + 12. squamosal. + 13. supratemporal. + 14. quadratojugal. + 15. sclerotic ring. + 16. postorbital. + 17. jugal. + 18. lachrymal. + 19. dentary. + 20. articular. + 21. angular.] + +The ribs are long, and the anterior ones have capitula and tubercula. +There is no sternum, but the ventral body wall is strengthened by a +complex system of abdominal splint ribs. + +The pectoral girdle is strongly developed, the scapulae are narrow, +the coracoids broad, and meet ventrally without overlapping. There are +probably no precoracoids, but clavicles and a T-shaped interclavicle +are well developed. + +The limbs are very short, and completely modified into swimming +paddles. The humerus and femur are both short, while the radius and +ulna, tibia and fibula are generally still further reduced to the form +of short polygonal bones. + +The digits are formed of longitudinal series of very numerous small +bones. The number of digits is five, but there sometimes appear to be +more owing to the bifurcation of certain of them, or to the addition +of marginal bones, either to the radial or ulnar side of the limb. The +humerus has no foramen, and both humerus and femur are unique in that +they are distally terminated by concave surfaces instead of by convex +condyles. The pelvic limb is much smaller than the pectoral. The +pelvis has no bony connection with the vertebral column, and all the +component bones are small and rod-like. + +The Ichthyosauria are confined to beds of Secondary age and by far the +best known genus is _Ichthyosaurus_. + + +_Order 5._ RHYNCHOCEPHALIA. + +This order includes the living _Sphenodon_ (_Hatteria_) and various +extinct forms. The general shape of these animals is lizard-like and +the tail is long. + +The vertebrae are amphicoelous or sometimes nearly flat, and the +notochord sometimes persists to some extent. _Proterosaurus_ differs +from the other members of the order in having opisthocoelous cervical +vertebrae. + +The sacrum is composed of two vertebrae. Ossified inter centra +(interdorsalia) generally occur in the cervical and caudal regions, +and sometimes throughout the whole vertebral column. In the skull the +quadrate is immovably fixed and united to the pterygoid. The palate is +well ossified, while the premaxillae which are often beak-like are +never ankylosed together. The jaws may be toothless or may be provided +with teeth which are usually acrodont (see p. 199). The palatines +frequently bear teeth, and in _Proterosaurus_ teeth occur also on the +pterygoids and vomers. The rami of the mandible are united by ligament +at the symphysis except in the Rhynchosauridae, in which the union is +bony. Superior and inferior temporal arcades occur. + +The ribs have capitula and tubercula, and often uncinate processes +(see p. 190) as in birds. A pectoral girdle and sternum, with +clavicles and a T-shaped interclavicle are developed, and abdominal +ribs are always found. The precoracoid is however absent. The limbs +are pentedactylate. + +_Sphenodon_[75] (Hatteria) now living in some of the islands of the +New Zealand group, is certainly the most generalised of all living +reptiles. Though lizard-like in form it differs from all living +lizards in the possession of two temporal arcades, abdominal ribs and +a fixed quadrate; and is often considered to be nearly allied in many +respects to the type of reptile from which all the others took their +origin. + +Among the better known extinct forms are _Proterosaurus_ of Permian +and _Hyperodapedon_ of Triassic age. + + +_Order 6._ SQUAMATA. + +This order includes the extinct Mosasaurians, and the lizards and +snakes which form the vast majority of living reptiles. The trunk may +be moderately elongated and provided with four short limbs as in +lizards, or it may be limbless, extremely elongated, and passing +imperceptibly into the tail. The surface is generally completely +covered with overlapping horny epidermal scales, below which bony +dermal scutes may be developed. + +The vertebrae are procoelous, rarely amphicoelous. There are no inter +centra, and the neural arches are firmly united to the centra. +Additional articulating surfaces, the zygosphenes and zygantra, are +often developed[76]. The sacrum is formed of two or rarely three +vertebrae, or may be wanting as in Ophidia. In the skull an +infratemporal arcade forming the lower boundary of the infratemporal +fossa is absent, and the quadrate, except in the Chamaeleons, is +movably articulated to the squamosal. The palatal vacuities are large +and the nares are separate. There is often a distinct parasphenoid. +The teeth are either _acrodont_ (i.e. ankylosed to the summit of the +jaw), or _pleurodont_, i.e. ankylosed to the inner side of the jaw. +The thoracic ribs each have a single head which articulates with the +centrum of the vertebra; while uncinate processes and abdominal ribs +never occur. + +A pectoral girdle and sternum may be present, or may be completely +absent as in snakes. Except in snakes there are generally four +pentedactylate limbs which may either form paddles or be adapted for +walking. + + +_Suborder (1)._ LACERTILIA[77]. + +The body is elongated, and as a rule four short pentedactylate limbs +are present, but sometimes limbs are vestigial or absent. The +exoskeleton generally has the form of horny plates, spines, or scales; +while sometimes as in the Chamaeleons and Amphisbaenians it is absent. +In other forms such as _Tiliqua_ and _Scincus_, the body has a +complete armour of bony scutes, whose shape corresponds with that of +the overlying horny scales. + +The vertebrae are procoelous, rarely as in the Geckos amphicoelous; +they are usually without zygosphenes and zygantra, but these +structures occur in the Iguanidae. The sacral vertebrae of living +forms are not ankylosed together, and the caudal vertebrae usually +have well-developed chevron bones. + +In the skull[78] the orbits are separated from one another, only by an +imperfectly developed interorbital septum, the cranial cavity not +extending forwards between them, while the alisphenoidal region is +unossified. The premaxillae may be paired or united (Amphisbaenidae), +and there is usually an interparietal foramen. There may be a complete +supratemporal[79] arcade bounding the lower margin of the +supratemporal fossa, or the supratemporal fossa may be open below. The +quadratojugal is not ossified, and the quadrate articulates with the +exoccipital. There is no infratemporal arcade. There is commonly a +rod-like epipterygoid[80] (fig. 33, 14) connecting the pterygoid and +parietal. + +Teeth are always present, and may be confined to the jaws or may be +developed also on the pterygoids and rarely on the palatines; they are +either acrodont or pleurodont. The rami of the mandible are suturally +united. + +A pectoral girdle is always present, and generally also a sternum. +Clavicles and a T-shaped interclavicle are commonly present, but are +absent in the Chamaeleons. + +[Illustration FIG. 33. A, LATERAL VIEW, AND B, LONGITUDINAL SECTION OF +THE SKULL OF A LIZARD (_Varanus varius_). × 3/5. (Brit. Mus.) + + 1. premaxillae. + 2. maxillae. + 3. nasal. + 4. lateral ethmoid. + 5. supra-orbital. + 6. lachrymal. + 7. frontal. + 8. postfrontal. + 9. prefrontal. + 10. basisphenoid. + 11. pro-otic. + 12. epi-otic. + 13. pterygoid. + 14. epipterygoid (columella cranii). + 15. jugal. + 16. transpalatine. + 17. parasphenoid. + 18. quadrate. + 19. parietal. + 20. squamosal. + 21. supratemporal. + 22. exoccipital. + 23. dentary. + 24. splenial. + 25. supra-angular. + 26. angular. + 27. coronoid. + 28. articular. + 29. vomer. + 30. basi-occipital. + 31. orbitosphenoid.] + +There is no separate precoracoid but a precoracoidal process (fig. 34, +7) of the coracoid is generally prominent. + +[Illustration FIG. 34. LATERAL VIEW OF THE SHOULDER-GIRDLE OF +_Varanus._ × 3/5. + +(Brit. Mus.). + + 1. suprascapula. + 2. scapula. + 3. glenoid cavity. + 4. coracoid. + 5. clavicle. + 6. interclavicle. + 7. precoracoidal process.] + +Sternal ribs are present in chamaeleons and scinks. The limbs are in +the great majority of cases pentedactylate and the digits are clawed. +The phalanges articulate by means of condyles. Sometimes one or both +pairs of limbs are absent. When the posterior limbs are absent the +pelvis is also wanting, though the loss of the anterior limbs does not +lead to a corresponding loss of the pectoral girdle. + +The pubis corresponds to the pre-pubis of Dinosaurs, and both pubes +and ischia meet in ventral symphyses. + +The suborder includes the Lizards, Chamaeleons and Amphisbaenians. + + +_Suborder (2)._ OPHIDIA[81]. + +The Ophidia or snakes are characterised by their greatly elongated +body and want of limbs. The body is covered with overlapping horny +scales and bony dermal scutes are never present. The vertebrae are +procoelous, and are distinguishable into two groups only, precaudal or +rib-bearing, and caudal or ribless. The atlas vertebra is also +ribless. The neural arches are always provided with zygosphenes and +zygantra. Many of the vertebrae have strong hypapophyses, and the +caudal vertebrae are without chevron bones. + +In the skull the cranial cavity extends forwards between the orbits, +and is closed in front by downgrowths from the frontals and parietals +which meet the well-ossified alisphenoids and orbitosphenoids[82]. The +cranium is strongly ossified, and there are no parotic processes or +interparietal foramen. There are no temporal arcades and no +epipterygoid. The premaxillae if present are very small (fig. 51, 1) +and usually toothless. The quadrates articulate with the squamosals, +and do not as in Lacertilia meet the exoccipitals. The palatines do +not unite directly with the vomers or with the base of the cranium, +and the whole palato-maxillary apparatus is more loosely connected +with the cranium than it is in Lacertilia. The pterygoids, and in most +cases also the palatines, bear teeth. The dentition is acrodont, and +the rami of the mandible are united only by an elastic ligament--an +important point serving to distinguish the Ophidia from the +Lacertilia. There is an imperfectly developed interorbital septum, the +ventral part of which is formed by the parasphenoid. The postfrontal +is generally well developed, while the jugals and quadratojugals are +absent. There are never any traces of the anterior limbs or pectoral +girdle, but occasionally there are vestiges of a pelvis and posterior +limbs. + + +_Suborder (3)._ PYTHONOMORPHA[83]. + +This suborder includes _Mosasaurus_ and its allies, a group of +enormous extinct marine reptiles found in beds of Cretaceous age. + +The skin is in most forms at any rate unprovided with dermal scutes. +The vertebrae may be with or without zygosphenes and zygantra. The +skull resembles that of lizards, having an interparietal foramen, and +a cranial cavity open in front. The squamosal takes part in the +formation of the cranial wall, and the quadrate articulates with the +squamosal, not as in Lacertilia with the exoccipital. There are large +supratemporal fossae, bounded below by supratemporal arcades. The +teeth are large and acrodont, and occur on the pterygoids as well as +on the jaws. The two rami of the mandible are united by ligament only. +Pectoral and pelvic girdles are present, but clavicles are wanting, +and the pelvis is not as a rule united to any sacrum. + +The limbs are pentedactylate, and are adapted for swimming, while all +the limb bones except the phalanges are relatively very short. The +number of phalanges is not increased beyond the normal, and they +articulate with one another by flat surfaces. The terminal phalanges +are without claws. + + +_Order 7._ DINOSAURIA[84]. + +The extinct reptiles comprising this order were all terrestrial, and +include the largest terrestrial animals known. They vary greatly in +size and in the structure of the limbs, some approach close to the +type of structure met with in birds, others are allied to crocodiles. + +Passing to the more detailed characters:--there is sometimes a +well-developed exoskeleton having the form of bony plates or spines. +The vertebrae may be solid or their centra may be hollowed internally; +their surfaces may be flat, biconcave or opisthocoelous. The sacrum is +composed of from two to six vertebrae. + +As regards the skull, the quadrate is large and fixed, and +supratemporal and infratemporal fossae bounded by bone occur. The +teeth are more or less laterally compressed, and often have serrated +edges; they may be placed in distinct sockets or in a continuous +groove. The ribs have capitula and tubercula, and sternal ribs often +occur. The scapula is very large, the coracoid small, and there is no +precoracoid, or T-shaped interclavicle. Clavicles are only known in a +few cases. In the pelvis the ilium is elongated both in front of, and +behind, the acetabulum, sometimes the pre-pubis, sometimes the +post-pubis is the better developed. The anterior limbs are shorter +than the posterior, and the long bones are sometimes solid, sometimes +hollow. + +There are three well-marked suborders of the Dinosauria. + + +_Suborder (1)._ SAUROPODA[85]. + +The reptiles belonging to this group were probably quadrupedal and +herbivorous. + +They have the cervical and anterior trunk vertebrae opisthocoelous, +while the posterior vertebrae are biconcave; all the presacral, and +sometimes the sacral vertebrae are hollowed internally. The teeth are +spatulate and without serrated edges, they are always planted in +distinct sockets, and some of them are borne by the premaxillae. + +[Illustration FIG. 35. RESTORED SKELETON OF _Ceratosaurus nasicornis_. +× 1/30. (After Marsh.) + + 1. anterior nares. + 2. prominence on the nasal bones which probably carried a horn. + 3. pre-orbital vacuity. + 4. orbit. + 5. scapula. + 6. coracoid. + 7. ilium. + 8. pubis (pre-pubis). + 9. ischium.] + +The nares have the form of long slits and there are large pre-orbital +vacuities. + +The limb bones are solid, and the anterior limbs are not much shorter +than the posterior ones. All the limbs are plantigrade and +pentedactylate, and the digits of the pes are clawed. There is a large +pre-pubis directed downwards and forwards, meeting its fellow in a +ventral symphysis, but there is no post-pubis. + +The Sauropoda are found in the secondary rocks of Europe and N. +America and include the largest land animals that are known to have +existed. Many of the best known forms such as _Brontosaurus_ and +_Morosaurus_ are North American. + + +_Suborder (2)._ THEROPODA. + +The members of this suborder were all carnivorous, and from the small +comparative size of the anterior limbs many of them were probably +bipedal. + +The vertebrae are opisthocoelous or amphicoelous, their neural arches +are provided with zygosphenes and zygantra, and their centra are +frequently hollowed internally; the limb bones are also hollow, and in +fact the whole skeleton is extremely light. The tail is of great +length. The teeth are pointed and recurved, and have one or both +borders serrated; they are always planted in distinct sockets, and +some of them are borne by the premaxillae. There are large pre-orbital +vacuities. The digits of both manus and pes are terminated by pointed +ungual phalanges which must have borne claws. In the pelvis the +pre-pubes and ischia are slender bones, the former meeting in a +ventral symphysis. The ilia are very deep vertically and there are no +post-pubes. The astragalus is closely applied to the tibia, in front +of which it sends an ascending process, sometimes the two bones appear +to have been ankylosed together, as in birds. The metatarsals are +elongated and the feet digitigrade. + +The Theropoda vary greatly in size, one of the best known genera +_Compsognathus_ was about as large as a cat, another, _Megalosaurus_, +perhaps as large as an elephant. _Ceratosaurus_ is the name of a +well-known North American form regarded by many authorities as +identical with _Megalosaurus_. + + +_Suborder (3)._ ORTHOPODA. + +This suborder includes the most specialised of the Dinosaurs, certain +of which resemble the Theropoda in being bipedal. In some of them such +as _Stegosaurus_ the exoskeleton is strongly developed, in others such +as _Iguanodon_ it is absent. + +The vertebrae are solid and may be opisthocoelous, biconcave, or flat. +The teeth are compressed and serrated, often irregularly, and are +frequently not set in distinct sockets. The anterior part of the +premaxillae is without teeth, and a toothless predentary or +mento-meckelian bone is present. The pre-orbital vacuities are small +or absent, and the nares are large and placed far forwards. + +The most characteristic features of the group are found in the pelvis +which, except in the Ceratopsia, bears a striking resemblance to that +of birds. The ischium and post-pubis are long slender bones directed +backwards parallel to one another, and the pre-pubis is also well +developed. The ischium has an obturator process. The limb bones are +sometimes hollow, sometimes solid. The anterior limbs are much shorter +than the posterior, pointing to a bipedal method of progression. The +pes is digitigrade or plantigrade, and has three, rarely four, +digits. + +The suborder Orthopoda may be further subdivided into three +sections:-- + + +A. STEGOSAURIA. + +A dermal exoskeleton is strongly developed. The vertebral centra are +flat or biconcave, and neither they nor the limb bones are hollowed +out by internal cavities. The limbs are plantigrade, the anterior ones +short, the posterior ones very large and strong. The post-pubis is +well developed; + +e.g. _Stegosaurus_ from the Upper Jurassic of Colorado. + + +B. CERATOPSIA. + +There is sometimes a well-developed dermal exoskeleton formed of small +granules and plates of bone. The bones are solid, and the vertebral +centra flat. The cranium bears a pair of enormous pointed frontal +horns, and the parietal is greatly expanded and elevated behind, +forming with the squamosals a shield which overhangs the anterior +cervical vertebrae. The premaxillae are united, and in front of them is +a pointed beak-like bone which bites upon a toothless predentary +ossification of the mandible. The teeth have two roots. The anterior +limbs are but little shorter than the posterior ones. There is no +post-pubis; + +e.g. _Polyonax_ from the uppermost Cretaceous of Montana. + + +C. ORNITHOPODA[86]. + +There is no dermal exoskeleton. The cervical vertebrae are +opisthocoelous, and so are sometimes the thoracic. The limb bones are +hollow and the anterior limbs are much shorter than the posterior +ones. The feet are digitigrade and provided with long pointed claws. +The post-pubis is long and slender and directed back parallel to the +ischium; + +e.g. _Iguanodon_ from the European Cretaceous. + + +_Order 8._ CROCODILIA[87]. + +This order includes the Crocodiles, Alligators and Garials and various +extinct forms, some of which are closely allied to the early +Dinosaurs. + +There is always a more or less complete exoskeleton formed of bony +scutes overlain by epidermal scales; these bony scutes are specially +well developed on the dorsal surface but may occur also on the +ventral. The vertebral column is divisible into the five regions +commonly distinguishable. In all living forms the vertebrae, with the +exception of the atlas and axis, the two sacrals, and first caudal, +are procoelous, but in many extinct forms they are amphicoelous. The +atlas (fig. 71) is remarkable, consisting of four pieces, and the +first caudal is biconvex. + +The teeth are, in the adult, planted in separate deep sockets. The +skull is very dense and solid, and all the component bones including +the quadrate are firmly united. The dorsal surface of the skull is +generally characteristically sculptured. There is an interorbital +septum, and the orbitosphenoidal and presphenoidal regions are +imperfectly ossified. Supratemporal, infratemporal, and post-temporal +fossae occur, but no interparietal foramen. In living genera there is +a long secondary palate formed by the meeting in the middle line of +the palatines, pterygoids and maxillae (fig. 43, A). + +Cervical ribs (fig. 41, 8 and 9) are well developed, and articulate +with rather prominent surfaces borne on the neural arches and centra +respectively. The thoracic ribs articulate with the long transverse +processes, and sternal ribs and abdominal splint ribs (fig. 46, 4) +occur. The sternum is cartilaginous, and both it and the +shoulder-girdle are very simple. The precoracoid is represented by +merely a small process on the coracoid, while the clavicles are +absent, except in the Parasuchia. In the pelvis (fig. 49) there is a +large ilium, and an ischium meeting its fellow in a ventral symphysis; +these two bones form almost the whole of the acetabulum. In front of +the acetabulum, in the Eusuchia, projects a bone which is generally +called the pubis, but is in reality rather an epipubis (fig. 49, 4), +the true pubis being probably represented by a fourth element which +remains cartilaginous for some time, and later on ossifies and +attaches itself to the ischium. The limbs are small in proportion to +the size of the body, and are adapted for swimming or for shuffling +along the ground; they are plantigrade and the bones are all solid. In +living forms the anterior limbs have five digits and the posterior +four, the fifth being represented only by a short metatarsal. The +first three digits in each case are clawed. The calcaneum has a large +backwardly-projecting process. + +The order Crocodilia may be subdivided into two suborders. + + +_Suborder (1)._ PARASUCHIA. + +The vertebral centra are flat or biconcave. The premaxillae are very +large, and the nares are separated, and placed far back. The posterior +narial openings lie comparatively far forward between the anterior +extremities of the palatines. + +The palatines and pterygoids do not form a secondary palate. The +supratemporal fossae are small, and open posteriorly, the lateral +temporal fossae are very large. The parietals and frontals are paired. +Clavicles are present. The best known and most important genus of +these extinct crocodiles is _Belodon_. + + +_Suborder_ (2). EUSUCHIA. + +The vertebrae are either biconcave or procoelous. The premaxillae are +small, and the anterior nares are united and placed far forwards. The +posterior nares lie far back, the palatines and in living genera the +pterygoids, meeting in the middle line, and giving rise to a closed +palate. The supratemporal fossae are surrounded by bone on all sides, +and the parietals, and often also the frontals are united. There are +no clavicles. The suborder includes the genera _Crocodilus_, +_Alligator_, _Garialis_ and others living and extinct. + + +_Order 9._ PTEROSAURIA[88]. + +These animals, called also the pterodactyles or Ornithosauria, are a +group of extinct reptiles, whose structure has been greatly modified +from the ordinary reptilian type for the purpose of flight. + +The skin was naked and they vary greatly in size and in the length of +the tail. The vertebrae and limb bones are pneumatic just as in birds. +The presacral vertebrae are procoelous and have their neural arches +firmly united to the centra. The neck is long, the caudal vertebrae +are amphicoelous, and from three to five vertebrae are fused together +in the sacral region. The skull is large and somewhat bird-like, the +facial portion being much drawn out anteriorly, and the sutures being +obliterated. It resembles that of other reptiles in having large +supratemporal fossae; large pre-orbital vacuities also occur. The jaws +may be toothed or toothless, and the teeth, when present, are imbedded +in separate sockets. The premaxillae are large, and the quadrate is +firmly attached to the skull. The rami of the mandible are united at +the symphysis, and there is an ossified ring in the sclerotic. The +occurrence of a postfrontal and its union with the jugal behind the +orbit, are characteristic reptilian features. + +The ribs have capitula and tubercula, and sternal and abdominal ribs +occur. The sternum has a well-developed keel, and the scapula and +coracoid are large and bird-like. There are no clavicles or +interclavicle. + +The anterior limbs are modified to form wings by the great elongation +of the fifth digit, to which a membrane was attached. The second, +third and fourth digits are clawed and are not elongated in the way +that they are in bats. The pollex, if present at all, is quite +vestigial. + +The pelvis is weak and small, and though the ilia are produced both in +front of and behind the acetabula, in other features the pelvis is not +bird-like. The ischia are short and wide, and the pubes are +represented only by the pre-pubes. The posterior limbs are small and +the fibula is much reduced. The pes is quite reptilian in type, and +has five separate slender metatarsals. The two best known genera are +_Pterodactylus_, in which the tail is short, and _Rhamphorhynchus_, in +which it is long. The Pterosauria are found throughout the Jurassic +and Cretaceous formations in both Europe and North America. + + +FOOTNOTES: + +[65] According to Baur a distinct epi-otic is not recognisable in the +reptilian skull. + +[66] H. Gadow, _Phil. Trans._, vol. 179, 1888. + +[67] See G. Baur, _J. Morph._, vol. I., 1887. R. Lydekker, _Catalogue +of the Fossil Reptilia and Amphibia in the British Museum_, Parts I. & +II. C.K. Hoffmann, _Reptilien_, in Bronn's _Classen und Ordnungen des +Thierreichs_, Bd. VI., 3 abth. 1879-90. + +[68] T.H. Huxley, _Quart. J. Geol. Soc._, vol. XV. p. 649, 1859. R. +Owen, _Catalogue of Fossil Reptiles of S. Africa in the British +Museum_, London, 1876. H.G. Seeley, various papers published in the +_P.R. Soc. London_, and _Phil. Trans._ + +[69] See pp. 281-283. + +[70] An ent-epicondylar foramen is one piercing the humerus on its +inner side just above the condyle. + +[71] According to Hulke they should be regarded as the +omosternum,--the clavicles and interclavicle being wanting. + +[72] See p. 272. + +[73] R. Lydekker, _Nat. Sci._ vol. I. p. 514, 1892. Further references +are there given. + +[74] The exact position of the suture between the prefrontal and +postfrontal is not known. + +[75] A. Günther, On the Anatomy of Hatteria, _Phil. Trans_, vol. 157, +1867, p. 595. + +[76] =Zygosphenes= are extra articulating surfaces borne upon the +anterior face of the neural arch; they fit into corresponding +structures, the =zygantra=, which are borne on the posterior surface +of the neural arch of the preceding vertebra. Ordinary zygapophyses +always accompany them. + +[77] See E.D. Cope, _P. Amer. Phil. Soc_. vol. xxx. p. 185. + +[78] See W.K. Parker, _Phil. Trans._ vol. 170, 1879, p. 595. + +[79] See p. 281. + +[80] Often called the columella cranii. + +[81] See C.K. Hoffmann, in Bronn's _Klassen und Ordnungen des +Thierreichs_, Bd. VI., 3 abth. 1885-90. + +[82] Some anatomists consider that the closing in of the brain case in +front is entirely due to the frontals and parietals. + +[83] E.D. Cope, _Rep. U.S. Geol. Surv._, 1875, vol. II., The +vertebrata of the Cretaceous formations of the west. E.D. Cope, _P. +Boston Soc._ 1862, XII. p. 250. O.C. Marsh, _Amer. J. Sci._, 1872, +vol. 3. R. Owen, _Quart. J. Geol. Soc._, 1877, and 1878. + +[84] J.W. Hulke, _Presidential address to the Geol. Soc. of London_, +1883 and 1884. O.C. Marsh, many papers in the _Amer. J. Sci._ from +1878 onwards, also in the _Geol. Mag._ R. Owen, _History of British +fossil reptiles: Dinosauria_ (Palaeont. Soc.). + +[85] The diagnostic characters of the different groups of Dinosaurs +are in the main those given by von Zittel. + +[86] See O.C. Marsh, _Amer. J. Sci._ (3), vol. 48, 1894, p. 85. + +[87] See C.B. Brühl, _Das Skelet der Krokodiliden_, Wien, 1862. C.K. +Hoffmann in Bronn's _Klassen und Ordnungen des Thierreichs_, Bd. VI. +Abth. III. 1881-85. T.H. Huxley, _Proc. Linn. Soc._ (Zoology) 1860 +vol. IV. p. 1. R. Owen, _History of British fossil Reptiles_. +_Crocodilia_ (Palaeont. Soc.). A. Smith Woodward, _Geol. Mag._ 1885, +3rd dec. II. p. 496. A. Smith Woodward, _Proc. of Geologists' Assoc._ +vol. IX. p. 288, 1886. + +[88] See H.G. Seeley, On the Organisation of the Ornithosauria, +_Journ. Linn. Soc._ (Zoology) vol. XIII. p. 84. K.A. Zittel, Ueber +Flugsaurier aus dem lithographischen schiefer, _Palaeontograph._ XXIX. +p. 49. + + + + +CHAPTER XIV. + +THE SKELETON OF THE GREEN TURTLE. + +(_Chelone midas._) + + +The most striking feature as regards the skeleton of the Turtle is +that the trunk is enveloped in a bony box, the dorsal portion of which +is called the =carapace=, while the ventral portion is the =plastron=. + + +I. EXOSKELETON. + +_a._ The =epidermal exoskeleton= in the Green Turtle as in all other +Chelonia except _Dermochelys_, _Trionyx_ and their allies is strongly +developed, its most important part consisting of a series of horny +=shields= which cover over the bony plates of the carapace and +plastron but do not at all correspond to them in size and arrangement. + +The shields covering over the =carapace= consist of three rows of +larger central shields,--five (=vertebral=) shields being included in +the middle row and four (=costal=) in each lateral row,--and of a +number of smaller =marginal= shields. + +Of the marginal shields, that lying immediately in front of the first +vertebral is termed the =nuchal=, while the two succeeding the last +vertebral are called sometimes =pygal=, sometimes =supracaudal=; the +remainder are the marginal shields proper. + +The epidermal covering of the =plastron= consists principally of six +pairs of symmetrically arranged shields, called respectively the +=gular=, =humeral=, =pectoral=, =abdominal=, =femoral=, and =anal=, +the gular being the most anterior. In front of the gular shields is an +unpaired =intergular=, and the shields of the plastron are connected +laterally with those of the carapace, by five or six pairs of rather +irregular =infra-marginal= shields. Smaller horny plates occur on +other parts of the body, especially on the limbs and head. + +Two other sets of structures belong also to the epidermal exoskeleton, +viz. (_a_) horny =beaks= with denticulated edges which ensheath both +upper and lower jaws, (_b_) =claws=, which as a rule are borne only by +the first digit of each limb. Sometimes in young individuals the +second digit is also clawed. + +_b._ The =dermal exoskeleton= is strongly developed, and is combined +with endoskeletal structures derived from the ribs and vertebrae to +form the carapace. + +The =Carapace= (fig. 36) consists of a number of plates firmly united +to one another by sutures. They have a very definite arrangement and +include: + +(_a_) the =nuchal= plate (fig. 36, 1), a wide plate forming the whole +of the anterior margin of the carapace. It is succeeded by three +series of plates, eight in each series, which together make up the +main part of the carapace. Of these the small + +(_b_) =neural plates=[1] (fig. 36, A, 2) form the middle series. They +are closely united with the neural arches of the underlying vertebrae; + +(_c_) the =costal plates=[89] (fig. 36, A, 3) are broad arched plates +united to one another by long straight sutures. They are united at +their inner extremities with the neural plates, but the boundaries of +the two sets of plates do not regularly correspond. Each is united +ventrally with a rib which projects beyond it laterally for some +distance; (_d_) the =marginal plates= (fig. 36, 4) are twenty-three +in number, eleven lying on each side, while an unpaired one lies in +the middle line posteriorly. Many of them are marked by slight +depressions into which the ends of the ribs fit; + +(e) the =pygal= plates (fig. 36, 5) are two unpaired plates lying +immediately posterior to the last neural. + +[Illustration Fig. 36. A, DORSAL AND B, VENTRAL VIEW OF THE CARAPACE +OF A LOGGERHEAD TURTLE (_Thalassochelys caretta_), (after Owen). + + 1. nuchal plate. + 2. first neural plate. + 3. second costal plate. + 4. marginal plate. + 5. pygal plate. + 6. rib. + 7. thoracic vertebra. + 8. first vertebral shield. + 9. costal shield.] + +The sculpturing due to the epidermal shields is very obvious on the +carapace. + +The =plastron= (fig. 37) consists of one unpaired ossification, the +=entoplastron=, and four pairs of ossifications called respectively +the =epiplastra=, =hyoplastra=, =hypoplastra=, and =xiphiplastra=. + +The =epiplastra= (fig. 37, 1) are the most anterior, they are expanded +and united to one another in the middle line in front, while behind +each tapers to a point which lies external to a process projecting +forwards from the hyoplastron. They are homologous with the +_clavicles_ of other vertebrates. + +The =entoplastron= or =episternum= (fig. 37, 2) which is homologous +with the _interclavicle_ of other reptiles, is expanded at its +anterior end and attached to the symphysis of the epiplastra, while +behind it tapers to a point and ends freely. + +The =hyoplastra= are large irregular bones each closely united +posteriorly with the corresponding hypoplastron, and drawn out +anteriorly into a process which lies internal to that projecting +backwards from the epiplastron. Each gives off on its inner surface a +slender process which nearly meets its fellow, while the anterior half +of the outer surface is drawn out into several diverging processes. + +The =hypoplastra= (fig. 37, 4) are flattened bones resembling the +hyoplastra, with which they are united by long sutures; the posterior +half of both outer and inner surfaces is drawn out into a number of +pointed processes. + +The =xiphiplastra= are small flattened elongated bones meeting one +another in the middle line posteriorly. In front they are notched and +each interlocks with a process from the hypoplastron of its side. The +hyoplastra, hypoplastra and xiphiplastra are homologous with the +abdominal ribs of Crocodiles. + +[Illustration FIG. 37. THE PLASTRON OF A GREEN TURTLE (_Chelone +midas_). × 1/7. (Camb. Mus.) + + 1. epiplastron (clavicle). + 2. entoplastron (interclavicle). + 3. hyoplastron. + 4. hypoplastron. + 5. xiphiplastron.] + + +II. ENDOSKELETON. + +1. THE AXIAL SKELETON. + +The axial skeleton includes the vertebral column, the ribs, and the +skull. + +A. THE VERTEBRAL COLUMN AND RIBS. + +The number of vertebrae in the Green Turtle is thirty-eight, not a +great number as compared with that in many reptiles, and of these +eighteen are caudal. + +The vertebral column is divisible into four regions only--=cervical=, +=thoracic=, =sacral=, and =caudal=. + + +THE CERVICAL VERTEBRAE. + +These are eight in number, and are chiefly remarkable for the great +variety of articulating surfaces which their centra present, and for +their mobility upon one another. + +The first or =atlas= vertebra differs much from all the others and +consists of the following parts:-- + +_a._ the =neural arch=, formed of two separate ossifications united in +the mid-dorsal line; + +_b._ the =inferior arch=; + +_c._ the =centrum=, which is detached from the rest and forms the +odontoid process of the second vertebra. + +Each half of the =neural arch= consists of a ventral portion, the +=pedicel=, which lies more or less vertically and is united ventrally +to the inferior arch, and of a dorsal portion, the =lamina=, which +lies more or less horizontally and meets its fellow in the middle line +in front, partially roofing over the neural canal. Each pedicel bears +a facet on its anterior surface, which, with a corresponding one on +the inferior arch, articulates with the occipital condyle of the +skull. Three similar facets occur also on the posterior surface of the +pedicel and inferior arch, and articulate with the odontoid process. +The laminae meet one another in front, but do not fuse, while behind +they are separated by a wide triangular space. They bear a pair of +small downwardly-directed facets, the =postzygapophyses=, for +articulation with the prezygapophyses of the second vertebra. + +The =inferior arch= is a short irregular bone bearing two converging +facets for articulation with the occipital condyle and odontoid +process respectively. + +The =centrum= or =odontoid process= has a convex anterior surface for +articulation with the neural and inferior arches, and a concave +posterior surface by which it is united with the centrum of the second +or axis vertebra. It bears posteriorly a small epiphysis which is +really a detached portion of the inferior arch. + +The second or =axis= and following five cervical vertebrae, though +showing distinct differences, resemble one another considerably, each +having a fairly elongated centrum with a keel-like =hypapophysis=, +each having also a neural arch with prominent articulating surfaces, +the anterior of which, or =prezygapophyses=, look upwards and inwards, +while the posterior ones, the =postzygapophyses=, look downwards and +outwards. They however, as was previously mentioned, differ very +remarkably in the character of the articulating surfaces of the +centra. Thus the second and third vertebrae are convex in front and +concave behind, the fourth is biconvex, the fifth is concave in front +and convex behind. The sixth is concave in front and attached to the +seventh by a flat surface behind, the seventh has a flat anterior face +and two slightly convex facets behind. The vertebrae all have short +blunt transverse processes and the second has a prominent =neural +spine=. + +The =eighth cervical vertebra= is curiously modified, the centrum is +very short, has a rather prominent hypapophysis, and is convex behind, +while in front it articulates with the preceding centrum by two +concave surfaces. The neural arch is deeply notched in front and bears +two upwardly-directed prezygapophyses, while behind it is very massive +and is drawn out far beyond the centrum, bearing a pair of flat +postzygapophyses. The top of the neural arch almost or quite meets a +blunt outgrowth from the nuchal plate. + + +THE THORACIC VERTEBRAE. + +These are ten in number and are all firmly united with the ribs and +elements forming the carapace. + +The first thoracic vertebra differs from the others, the centrum is +short and has a concave anterior surface articulating with the centrum +of the last cervical vertebra, and a pair of prezygapophyses borne on +long outgrowths. The neural spine arises only from the anterior half +of the centrum, and is not fused to the carapace. Arising laterally +from the anterior part of the centrum are a small pair of ribs each of +which is connected with a process arising from the rib of the +succeeding vertebra. + +The next seven thoracic vertebrae are all very similar, each has a +long cylindrical centrum, expanded at the ends, and firmly united to +the preceding and succeeding vertebrae. The neural arches are +flattened and expanded dorsally, and are united to one another and to +the overlying neural plates; each arises only from the anterior half +of its respective centrum, and overlaps the centrum of the vertebra in +front of it. Between the base of the neural arch and its successor is +a small foramen for the exit of the spinal nerve. There are no +transverse processes or zygapophyses. + +To each thoracic vertebra from the second to ninth inclusive, there +corresponds a pair of =ribs= (fig. 36, 6) of a rather special +character. Each is suturally united with the anterior half of the edge +of its own vertebra, and overlaps on to the posterior half of the edge +of the next preceding vertebra. The ribs are much flattened, and each +is fused with the corresponding costal plate, beyond which it projects +to fit into a pit in one of the marginal plates. + +The tenth thoracic vertebra is smaller than the others, and its neural +arch does not overlap the preceding vertebra, it bears a pair of small +ribs which are without costal plates, but meet those of the ninth +vertebra. + +There are no =lumbar= vertebrae. + + +THE SACRAL VERTEBRAE. + +The =sacral vertebrae= are two in number, they are short and wide, +their centra are ankylosed together, and their neural arches are not +united to the carapace. + +The first has the anterior face of the centrum concave and the +posterior flat, while both faces of the second are flat. Each bears a +pair of short ribs which meet the ilia, but are not completely +ankylosed either with them or the centra. + + +THE CAUDAL VERTEBRAE. + +The =caudal vertebrae= are eighteen in number. The centrum of the +first is flat in front and is ankylosed to the second sacral; behind +it is convex. The others are all very similar to one another, and +decrease gradually in size when followed back. Each has a moderately +long centrum, concave in front and convex behind, both terminations +being formed by epiphyses. The neural arch arises only from the +anterior half of the vertebra; it bears a blunt truncated neural spine +and prominent pre- and post-zygapophyses. The first seven caudal +vertebrae bear short ribs attached to their lateral margins, the +similar outgrowths on the succeeding vertebrae do not ossify from +distinct centres, and are transverse processes rather than ribs. + + +B. THE SKULL. + +The skull of the Turtle is divisible into the following three parts:-- + +(1) the cranium; + +(2) the lower jaw or mandible; + +(3) the hyoid. + +(1) THE CRANIUM. + +The =cranium= is a very compact bony box, containing a cavity in which +the brain lies, and which is a direct continuation of the neural canal +of the vertebrae. + +[Illustration FIG. 38. THE SKULL OF THE GREEN TURTLE (_Chelone +midas_). × 1/2. A, POSTERIOR HALF, B, ANTERIOR HALF. (Brit. Mus.) + + 1. parietal. + 2. squamosal. + 3. quadrate. + 4. basisphenoid. + 5. basi-occipital. + 6. quadratojugal. + 7. opisthotic. + 8. exoccipital. + 9. foramen magnum. + 10. splenial. + 11. articular. + 12. dentary. + 13. angular. + 14. supra-angular. + 15. premaxilla. + 16. maxilla. + 17. jugal. + 18. postfrontal. + 19. vomer. + 20. prefrontal. + 21. frontal. + 22. external auditory meatus leading into tympanic cavity.] + +Like those of the skull as a whole its component bones may be +subdivided into three sets:-- + +1. those forming the brain-case or =cranium proper=; + +2. those developed in connection with the special sense organs; + +3. those forming the upper jaw and suspensorial apparatus. + +Both cartilage and membrane bones take part in the formation of the +skull, and a considerable amount of cartilage remains unossified, +especially in the ethmoidal and sphenoidal regions. + + +1. THE CRANIUM PROPER OR BRAIN-CASE. + +The cartilage and membrane bones of the brain-case when taken together +can be seen to be more or less arranged in three rings or segments, +called respectively the occipital, parietal, and frontal segments. + +The =occipital segment= is the most posterior of these, and consists +of four cartilage bones, the =basi-occipital=, the two =exoccipitals= +and the =supra-occipital=; these bound the =foramen magnum=. + +The =basi-occipital= (figs. 38 and 39, 5) lies ventral to the foramen +magnum and only bounds a very small part of it; it forms one-third of +the =occipital condyle= by which the skull articulates with the atlas +vertebra. It unites dorsally with the exoccipitals and anteriorly with +the basisphenoid. + +The =exoccipitals= are rather small bones, which form the sides and +the greater part of the floor of the foramen magnum, and two-thirds of +the occipital condyle. Laterally each is united with the pterygoid and +opisthotic of its side. At the sides of the occipital condyle each +exoccipital is pierced by a pair of foramina, the more dorsal and +posterior of which transmits the hypoglossal nerve. + +The =supra-occipital= (fig. 39, 14) is a larger bone than the others +of the occipital segment. It forms the upper border of the foramen +magnum and is drawn out dorsally into a large crest which extends back +far beyond the occipital condyle. In the adult the supra-occipital is +completely ankylosed with the epi-otics. + + +The =Parietal segment=. + +The ventral portion of the parietal segment is formed by the +=basisphenoid= (figs. 38 and 39, 4) which lies immediately in front of +the basi-occipital. A triangular portion of it is seen in a ventral +view of the skull, but it is quickly overlapped by the pterygoids. It +gives off dorsally a pair of short processes which meet the pro-otics. + +The alisphenoidal region is unossified and the only other constituents +of the parietal segment are the _parietals_ (fig. 39, 1). These are +large bones which, after roofing over the cranial cavity, extend +upwards and become expanded into a pair of broad plates which unite +with the squamosal and bones of the frontal segment to form a wide, +solid, false roof to the skull. Each also sends ventralwards a plate +which meets an upgrowth from the pterygoid and acts as an alisphenoid. + + +The =Frontal segment=. + +Of the frontal segment the basal or presphenoidal and lateral or +orbitosphenoidal portions do not become ossified, the dorsal portion +however includes three pairs of membrane bones, the _frontals_, +_prefrontals_ and _postfrontals_. + +The _frontals_ are a pair of small bones lying immediately in front of +the parietals, and in front of them are the _prefrontals_ (figs. 38 +and 39, 20), a pair of similar but still smaller bones, which are +produced ventrally to meet the vomer and palatines. They form also the +dorsal boundary of the anterior nares. The _postfrontals_ (figs. 38 +and 39, 18) are larger bones, united dorsally to the frontals and +parietals, posteriorly to the squamosals, and ventrally to the jugals +and quadratojugals. All three pairs of frontal bones, especially the +postfrontals, take part in the bounding of the orbits. + +[Illustration FIG. 39. LONGITUDINAL VERTICAL SECTION THROUGH THE +CRANIUM OF A GREEN TURTLE (_Chelone midas_). × 2/3. (Camb. Mus.) + + 1. parietal. + 2. squamosal. + 3. quadrate. + 4. basisphenoid. + 5. basi-occipital. + 6. quadratojugal. + 7. pro-otic. + 8. opisthotic. + 9. pterygoid. + 10. palatine. + 11. rod passed into narial passage. + 12. exoccipital. + 13. epi-otic fused to supra-occipital. + 14. supra-occipital. + 15. premaxillae. + 16. maxillae. + 17. jugal. + 18. postfrontal. + 19. vomer. + 20. prefrontal. + 21. frontal. + V, VII, VIII, IX, X, XI, XII, + foramina for the exit of + cranial nerves.] + + +2. THE SENSE CAPSULES. + +Skeletal structures occur in connection with each of the three special +sense organs of hearing, sight, and smell. + + +The =Auditory capsules=. + +The auditory or periotic capsule of the turtle is rather large and its +walls are well ossified, epi-otic, pro-otic and opisthotic bones being +present. + +The =epi-otic= (fig. 39, 13) is the more dorsal of the three bones, +and in the adult is completely ankylosed with the supra-occipital. + +The =opisthotic= (fig. 39, 8) is the ventral posterior element. On its +inner side it is united to the supra-occipital above, and to the +exoccipital below; it sometimes becomes completely fused with the +exoccipital. In front it meets the pro-otic, and on its outer side the +squamosal and quadrate. Its anterior portion is hollowed out by the +cavity in which the auditory organ lies, it gives off also a process +which is separated from the exoccipital by an oval foramen through +which the glossopharyngeal, pneumogastric, and spinal accessory nerves +leave the cranial cavity. + +The =pro-otic= is the anterior element; it meets the supra-occipital +and opisthotic posteriorly, while anteriorly it is separated from the +alisphenoidal plate of the parietal and pterygoid by a large oval +foramen through which the maxillary and mandibular branches of the +trigeminal nerve pass out (fig. 39, V 1 & 2). It is hollowed out +posteriorly by the cavity in which the auditory organ lies, and its +inner wall as seen in longitudinal section is pierced by a foramen +through which the external carotid artery and facial nerve leave the +cranial cavity,--the nerve finally leaving the skull through a small +oval foramen on the anterior face of the pro-otic near its junction +with the quadrate. + +Between the pro-otic and opisthotic as seen in a longitudinal section +of the skull is a large opening constricted in the middle. This is +the =internal auditory meatus= (fig. 39, VIII.). Through it the +auditory nerve leaves the cranial cavity and enters the ear. The ramus +vestibularis leaves through the dorsal part of the hole, the ramus +cochlearis through the ventral. + +The cavity of the auditory or periotic capsule communicates with the +exterior by a fairly large hole, the =fenestra ovalis=, which lies +between the opisthotic and pro-otic, and opens into a deep depression, +the =tympanic cavity=, which is seen in a posterior view of the skull +lying just external to the exoccipital. The cavity communicates with +the exterior by a large opening, the =external auditory meatus= (fig. +38, 22). + +Several other openings are seen in the tympanic cavity; through one at +the extreme posterior end the pneumogastric and spinal accessory +nerves leave the skull, and through another, a little further +forwards, the glossopharyngeal. + +The auditory ossicles consist of a long bony =columella=, whose inner +end fits into the fenestra ovalis, while the outer end is attached to +a small cartilaginous plate, the =extra-columella=, which is united to +the tympanum. + + +The =Optic capsules=. + +The skeletal structures developed in connection with the optic capsule +do not become united to the skull. They consist of:-- + +(_a_) the =sclerotic=, a cartilaginous sheath investing the eye and +bearing + +(_b_) a ring of ten small bony scales. + +There is no _lachrymal_ bone. + + +The =Olfactory= or =Nasal capsules=. + +The basicranial axis in front of the basisphenoid remains +cartilaginous, neither presphenoid nor mesethmoid bones are developed, +and the orbits in a dry skull communicate by a wide space through +which the second, third, fourth, and sixth cranial nerves pass out. +Separate _nasal_ bones also do not occur, the large prefrontals +extending over the area usually occupied by both nasals and +lachrymals. + +The only bone developed in connection with the nasal capsules is the +_vomer_ (fig. 39, 19), an unpaired bone lying ventral to the +mesethmoid cartilage, and in contact laterally with the maxillae, +premaxillae and palatines. + + +3. THE UPPER JAW AND SUSPENSORIAL APPARATUS. + +A number of pairs of bones are developed in connection with the upper +jaw and suspensorial apparatus, one pair, the =quadrates=, being +cartilage bones, while the rest are all membrane bones. + +The _squamosals_ (fig. 38, 2) are large bones which, lying external to +the auditory bones, extend dorsalwards to meet the parietals and +postfrontals, and form a large part of the false roof of the skull. +They are united ventrally with the quadrates and quadratojugals. + +Each =quadrate= (fig. 38, 3) forms the outer boundary of the tympanic +cavity, and is firmly united on its inner side with the opisthotic, +exoccipital, and pterygoid. Dorsally it is fixed to the squamosal and +anteriorly to the quadratojugal. Its outer surface is marked by a deep +recess, and it ends below in a strong condyle with which the mandible +articulates. In front of the quadrates are a pair of thin plate-like +bones, the _quadratojugals_ which are united in front to the jugals or +malars. + +The _jugals_ (fig. 38, 17) are also thin plate-like bones, and form +part of the posterior boundary of the orbit. They are attached +dorsally to the postfrontals, and anteriorly to the maxillae, while +each also sends inwards a horizontal process which meets the pterygoid +and palatine. + +The _maxillae_ (figs. 38 and 39, 16) are a pair of large +vertically-placed bones, each drawn out ventrally into a straight, +sharp, cutting edge. They form the lateral boundaries of the anterior +nares, and each sends dorsalwards a process which meets the +postfrontal. Each also sends inwards a horizontal =palatine process=, +which meets the palatine and vomer, and also forms much of the floor +of the narial passage. + +The _premaxillae_ (figs. 38 and 39, 15) are a pair of very small bones +forming the floor of the anterior narial opening, they are wedged in +between the two maxillae, and send back processes which meet the vomer +and palatines. + +The _palatines_ (fig. 39, 10) are a pair of small bones firmly united +with the pterygoids behind, with the maxillae and jugals externally, +and with the vomer in the middle line. Each also gives off a palatine +plate which unites with the expanded lower edge of the vomer, and +forms the ventral boundary of the posterior nares. Anteriorly the +palatines form the posterior boundary of a large foramen through which +the ophthalmic branches of the fifth and seventh nerves pass to the +olfactory organs. + +The _pterygoids_ (fig. 39, 9) are a pair of large bones which unite +with one another by a long median suture. They are united also with +the palatines in front, and with the quadrate, basisphenoid, +basi-occipital, and exoccipitals behind. Each also sends dorsalwards a +short =alisphenoid plate= which meets that from the parietal. + +Piercing the posterior end of the _pterygoid_ is the prominent opening +of the carotid canal; a bristle passed into this hole emerges through +a foramen lying between the pro-otic and the alisphenoid process of +the pterygoid. + + +(2) THE LOWER JAW OR MANDIBLE. + +The =mandible= consists of one unpaired bone, formed by the fusion of +the two _dentaries_, and five pairs of bones, called respectively the +=articular=, _angular_, _supra-angular_, _splenial_ and _coronoid_. + +The fused _dentaries_ (fig. 38, 12) form by far the largest of the +bones; they constitute the flattened anterior part of the mandible, +and extend back below the other bones almost to the end of the jaw. + +The _coronoid_ is the most anterior of the paired bones, it forms a +prominent process to which the muscles for closing the jaw are +attached. + +The =articular= (fig. 38, 11) is expanded, and with the +_supra-angular_ forms the concave articulating surface for the +quadrate. + +The _splenial_ (fig. 38, 10) is a thin plate applied to the inner +surface of the posterior part of the mandible. + +The _angular_ (fig. 38, 13) is a slender plate of bone lying below the +supra-angular and splenial. + + +(3) THE HYOID. + +The hyoid apparatus is well developed, parts of the first two +branchial arches being found, as well as of the hyoid proper. It +consists of a more or less oblong flattened =basilingual plate= or +=body of the hyoid= which represents the fused ventral ends of the +hyoid and branchial arches of the embryo, and is drawn out into a +point anteriorly. The greater part is formed of unossified cartilage, +but at the posterior end it is bilobed, and a pair of ossified tracts +occur. To its sides are attached three pairs of structures, which are +portions of the hyoid and first and second branchial arches +respectively. + +The free part of the =hyoid= consists of a small piece of cartilage +attached to the anterior part of the basilingual plate at its widest +portion (fig. 53, 2). + +The =anterior cornu= or free part of the =first branchial arch= is +much the largest of the three structures. Its proximal portion +adjoining the basilingual plate is cartilaginous, as is its distal +end; the main part is however ossified. + +The =posterior cornu= or free part of the =second branchial arch= +(fig. 53, 4) consists of a short flattened cartilaginous bar arising +from the bilobed posterior end of the basilingual plate. + +The hyoid apparatus has no skeletal connection with the rest of the +skull. + + +2. THE APPENDICULAR SKELETON. + +This includes the skeleton of the two pairs of limbs and their +girdles. + + +THE PECTORAL GIRDLE. + +The pectoral girdle has an anomalous position, being situated +internal or ventral to the ribs. It consists of three bones, a dorsal +bone, the =scapula=, an anterior ventral bone, the =precoracoid=, and +a posterior ventral bone, the =coracoid=. + +The =scapula= is a small somewhat rod-shaped bone forming about +two-thirds of the glenoid cavity. At its proximal end it is closely +united with the precoracoid, the two bones ossifying continuously. It +tapers away distally, and is directed dorsalwards towards the +carapace. + +The =precoracoid= forms an angle of about 130° with the scapula, with +which it is completely fused at its proximal end. Its distal end is +somewhat expanded and flattened, and is terminated by a +fibrocartilaginous =epiprecoracoid= which meets its fellow. It takes +no part in the formation of the glenoid cavity. + +The =coracoid= is a large flattened blade-shaped bone forming about +one-third of the glenoid cavity. It does not meet its fellow in a +ventral symphysis, and is terminated by a cartilaginous =epicoracoid=. +The glenoid articulating surfaces of both scapula and coracoid are +lined by a thick pad of cartilage. + + +THE ANTERIOR LIMB. + +This is divisible into three portions, the =upper arm=, =fore-arm= and +=manus=. + +The =upper arm= contains a single bone, the =humerus=. + +The =humerus= (fig. 40, A, 1) is a stout, nearly straight, somewhat +flattened bone widely expanded at both ends. At the proximal end is +the large hemispherical =head=, which articulates with the glenoid +cavity. Behind the head the bone is drawn out into another large +rounded process. Below the head the shaft bears a small outgrowth +which is continuous with a larger one on the flexor surface (see p. +29). The bone is terminated distally by the =trochlea=, consisting of +three partially distinct convex surfaces which articulate with the +bones of the fore-arm. + +The =fore-arm= includes two bones, the =radius= and =ulna=; both these +are small bones, and are immovably fixed to one another proximally and +distally. + +The =radius= or pre-axial bone is the larger of the two, and is a +rod-like bone terminated at either end by an epiphysis. It articulates +at its proximal end with the humerus, and at its distal end with the +radiale or scaphoid bone of the carpus. + +The =ulna= (fig. 40, A, 3) or postaxial bone is shorter than the +radius, and more expanded at its proximal end, where it articulates +with the humerus. It articulates distally with the intermedium (lunar) +and the ulnare (cuneiform) bones of the carpus. All three bones of the +arm have their terminations formed by epiphyses which ossify from +centres distinct from those forming the shafts. + +The =Manus= consists of the =carpus= or =wrist= and the =hand= which +includes the metacarpals and phalanges. + +The =carpus= consists of a series of ten small bones, one of which, +the =pisiform= (fig. 40, A, 10), differs from the others in being +merely an ossification in the tendon of a muscle. The remaining nine +bones are arranged in a proximal row of three, the =ulnare= (fig. 40, +A, 6), =intermedium=, and =radiale=, and a distal row of five +(carpalia 1-5), each of which supports one of the metacarpals. A ninth +bone, the =centrale= (fig. 40, A, 7), is wedged in between the two +rows. The ulnare, intermedium and pisiform are comparatively large +flattened bones, the others are small and cubical. + +The =hand=. This is composed of five digits, each of which consists of +a metacarpal and of a varying number of phalanges. + +The =metacarpals=. The first metacarpal (fig. 40, A, 11) is a short +flattened bone, the others are all elongated and cylindrical, and are +terminated proximally by slightly concave surfaces, and distally by +slightly convex ones. + +The =phalanges=. The first and fifth digits both have two phalanges, +the second, third, and fourth have each three. The distal phalanx of +the first digit is stout and curved, and bears a horny claw; those of +the other digits are flattened and more or less pointed. + +[Illustration FIG. 40. A. ANTERIOR LIMB OF A YOUNG HAWKSBILL TURTLE +(_Chelone imbricata_) × 1/4 (Brit. Mus.). B. POSTERIOR LIMB OF A LARGE +GREEN TURTLE (_Chelone midas_) × 1/8 (Camb. Mus.). + + 1. humerus. + 2. radius (almost hidden by the ulna). + 3. ulna. + 4. radiale. + 5. intermedium. + 6. ulnare. + 7. centrale. + 8. carpale I. + 9. carpale IV. + 10. pisiform. + 11. first metacarpal. + 12. femur. + 13. tibia. + 14. fibula. + 15. tibiale intermedium and centrale fused. + 16. fibulare. + 17. tarsale 1. + 18. tarsale 2. + 19. tarsalia 4 and 5 fused. + 20. first metatarsal. + 21. fifth metatarsal. + I, II, III, IV, V, digits.] + + +THE PELVIC GIRDLE. + +The pelvic girdle consists of three bones; a dorsal bone, the =ilium=, +an anterior ventral bone, the =pubis=, and a posterior ventral bone, +the =ischium=. All three bones contribute largely to the formation of +the =acetabulum=, with which the head of the femur articulates. + +The =ilium= is a small slightly curved bone, which unites ventrally +with the pubis and ischium, and extends dorsalwards and backwards to +meet the distal ends of the sacral ribs. + +The =pubis= is the largest bone of the three; its distal end forms a +wide bilobed plate, the inner lobe meeting its fellow in a median +symphysis, while the other lobe or lateral process extends outwards. +Attached to the symphysis in front is a cartilaginous =epipubis=, +while behind, the two pubes are terminated by a wide rounded +cartilaginous area. + +The =ischium=, the smallest bone of the three, is flattened and like +the pubis meets its fellow in a median symphysis. A narrow band of +cartilage connects the symphysis pubis with the symphysis ischii, and +separates the two =obturator foramina= from one another. + + +THE POSTERIOR LIMB. + +This is divisible into three portions, the =thigh=, the =crus= or +=shin=, and the =pes=. + +The =thigh= includes a single bone, the =femur=. + +The =femur= (fig. 40, B, 12) is a short thick bone, with a prominent +rounded =head= articulating with the acetabulum. Behind this head is a +deep pit, beyond which is a roughened area corresponding with the +great trochanter of mammals. The distal end is expanded and somewhat +convex. + +The bones of the =crus= or =shin= are the =tibia= and =fibula=. These +are both straight rod-like bones with expanded terminations which +closely approach one another, while elsewhere the bones diverge +considerably. + +The terminations of all three of the leg bones are formed by +epiphyses. + +The =Pes= consists of the =tarsus= or =ankle=, and the =foot=, which +is made up of five digits. + +The =tarsus=. The tarsal bones of the Turtle do not retain their +primitive arrangement to such an extent as do the carpals. They are +arranged in a proximal row of two and a distal row of four. Of the +bones in the proximal row the postaxial one is much the smaller and is +the =fibulare=; the larger pre-axial one (fig. 40, B, 15) represents +the =tibiale=, =intermedium=, and =centrale= fused, and articulates +with both tibia and fibula. The first three distal tarsalia are all +small bones and are very similar in size, and each articulates +regularly with the corresponding metatarsal. The fourth bone (fig. 40, +B, 19) is much larger, and represents tarsalia 4 and 5 fused. The +first two distal tarsalia articulate with the pre-axial tarsal of the +proximal row, the third only with its neighbours the second, and the +fused fourth and fifth. The latter articulates with both bones of the +proximal row. + +Each =digit= consists of a metatarsal and of a varying number of +phalanges. + +The =metatarsals=. The first metatarsal (fig. 40, B, 20) is broad and +flattened, the second, third and fourth, are all elongated bones with +nearly flat terminations formed by small epiphyses. The fifth is large +and flattened, and the articular surface for the phalanx is situated +somewhat laterally. + +The =phalanges=. The first digit has two phalanges and is the stoutest +of them all; its distal phalanx is sheathed in a large horny claw. The +other digits, of which the third is the longest, have each three +phalanges. The distal phalanges of the second and third digits are +flattened and pointed and bear small horny claws. + + +FOOTNOTES: + +[89] Another view commonly held is that the neural and costal plates +are respectively formed by the expanded neural arches and ribs. + + + + +CHAPTER XV. + +THE SKELETON OF THE CROCODILE. + + +The species chosen for description is _C. palustris_, a form occurring +throughout the Oriental region, but the description would apply almost +equally well to any of the other species of the genus _Crocodilus_, +and with comparatively unimportant modifications to any of the living +Crocodilia. + + +I. EXOSKELETON. + +The exoskeleton of the Crocodile is strongly developed and includes +elements of both epidermal and dermal origin. + +_a._ The =epidermal exoskeleton= is formed of a number of horny +=scales= or plates of variable size covering the whole surface of the +body. Those covering the dorsal and ventral surfaces are oblong in +shape, and are arranged in regular rows running transversely across +the body. The scales covering the limbs and head are mostly smaller +and less regularly arranged, and are frequently raised into a more or +less obvious keel. Those covering the dorsal surface of the tail are +very prominently keeled. + +The epidermal exoskeleton also includes the horny =claws= borne by the +first three digits of both manus and pes. + +_b._ The =dermal exoskeleton=. This has the form of bony =scutes= +which underlie the epidermal scales along the dorsal surface of the +trunk and anterior part of the tail. Except in very young individuals +the epidermal scales are rubbed off from these scutes, which +consequently come to project freely on the surface of the body. Each +scute is a nearly square bony plate, deeply pitted or sculptured, and +marked by a strong ridge on its dorsal surface, while its ventral +surface is smooth. Contiguous scutes are united to one another by +interlocking sutures. + +The scutes are arranged in two distinct areas, viz. (1) a small +anterior =nuchal shield= which lies just behind the head and is formed +of six large scutes more or less firmly united together, and (2) a +larger posterior =dorsal shield= covering the whole of the back and +anterior part of the tail, and formed of smaller scutes, which are +arranged in regular transverse rows, and progressively diminish in +size when followed back. + +The =teeth= are exoskeletal structures, partly of dermal, partly of +epidermal origin. They lie along the margins of the jaws and are +confined to the premaxillae, maxillae and dentaries. They are simple +conical structures, without roots; each is in the adult placed in a +separate socket, and is replaced by another which as it grows comes to +occupy the pulp cavity of its predecessor. In the young animal the +teeth are not placed in separate sockets but in a continuous groove. +This feature is met with also in the Ichthyosauria. The groove +gradually becomes converted into a series of sockets by the ingrowth +of transverse bars of bone. The anterior teeth are sharply pointed and +slightly recurved, the posterior ones are more blunt. + +The upper jaw bears about nineteen pairs of teeth, the lower jaw about +fifteen pairs. The largest tooth in the upper jaw is the tenth, and in +the lower jaw the fourth. + +The three living families of Crocodilia, the Crocodiles, Alligators +and Garials, can be readily distinguished by the characters of the +first and fourth lower teeth. In Alligators both first and fourth +lower teeth bite into pits in the upper jaw; in Garials they both bite +into notches or grooves in the upper jaw. In Crocodiles the first +tooth bites into a pit, the fourth into a notch in the upper jaw. + + +II. ENDOSKELETON. + +1. THE AXIAL SKELETON. + +This includes the vertebral column, the skull, and the ribs and +sternum. + +A. THE VERTEBRAL COLUMN. + +The vertebral column is very long, consisting of some sixty vertebrae. +It can be divided into the usual five regions, the cervical, thoracic, +lumbar, sacral, and caudal regions. + +[Illustration FIG. 41. FIRST FOUR CERVICAL VERTEBRAE OF A CROCODILE +(_C. vulgaris_). (Partly after VON ZITTEL.) + + 1. pro-atlas. + 2. lateral portion of atlas. + 3. odontoid process. + 4. ventral portion of atlas. + 5. neural spine of axis. + 6. postzygapophysis of fourth vertebra. + 7. tubercular portion of fourth cervical rib. + 8. first cervical rib. + 9. second cervical rib. + 10. convex posterior surface of centrum of fourth vertebra.] + +THE CERVICAL VERTEBRAE. + +Counting as cervical all those vertebrae which are anterior to the +first one whose ribs meet the sternum, there are nine cervical +vertebrae, all of which bear ribs. + +As a type of the cervical vertebrae the fifth may be taken. It has a +short cylindrical =centrum= deeply concave in front and convex behind. +From the anterior part of the ventral surface of the centrum arises a +short =hypapophysis=, and on each side is a facet with which the lower +limb (=capitulum=) of the cervical rib articulates. The =neural arch= +is strongly developed and drawn out dorsally into a long =neural +spine=, in front of which are a pair of upstanding processes bearing +the prominent upwardly and inwardly directed =prezygapophyses=. At the +sides and slightly behind the neural spine are a corresponding pair of +processes bearing the =postzygapophyses=, which look downwards and +outwards. At the point where it joins the centrum the neural arch is +drawn out into a short blunt =transverse process= with which the upper +limb (=tuberculum=) of the cervical rib articulates. The sides of the +neural arch are slightly notched behind for the exit of the spinal +nerves. + +The first or =atlas= vertebra differs much from any of the others, and +consists of four quite detached portions, a ventral arch, with two +lateral portions and one dorsal. The =ventral arch= (fig. 41, 4) is +flat below and slightly concave in front, forming together with two +flattened surfaces on the lateral portions a large articulating +surface for the occipital condyle of the skull. Its posterior face is +bevelled off and forms with a second pair of facets on the lateral +portions a surface with which the odontoid process of the second +vertebra articulates. The postero-lateral surfaces of the ventral arch +also bear a pair of little facets with which the cervical ribs +articulate. The lateral portions are somewhat flattened and expanded, +and bear in addition to those previously mentioned a pair of small +downwardly directed facets, the postzygapophyses, which articulate +with the prezygapophyses of the second vertebra. The dorsal portion +(fig. 41, 1) is somewhat triangular in shape, and overhangs the +occipital condyle. It is often regarded as the neural arch of a +vertebra in front of the atlas and is called the _pro-atlas_; but as +it is a membrane bone it is not properly a vertebral element. + +The second or =axis vertebra= also differs a good deal from the other +cervicals. The centrum is massive, and is terminated in front by a +very large slightly concave articulating surface formed by the +=odontoid process= (fig. 41, 3) which is united with the centrum by +suture only, and is really the detached centrum of the first vertebra. +The cervical rib (fig. 41, 9) articulates with two little +irregularities on the odontoid process. The posterior surface of the +centrum is convex. The neural arch is strongly developed and +terminated dorsally by a long neural spine (fig. 41, 5), its sides are +notched, slightly in front and more prominently behind for the exit of +the spinal nerves. It is drawn out in front into two little processes +bearing a pair of upwardly and outwardly directed prezygapophyses, +while the postzygapophyses are similar to those of the other cervical +vertebrae. + +The last two cervical vertebrae resemble the succeeding thoracic +vertebrae, in the increased length of the transverse processes, and +the shifting dorsalwards of the facet with which the capitulum of the +rib articulates. + +THE THORACIC VERTEBRAE. + +The thoracic vertebrae commence with the first of those that bears +ribs reaching the sternum. They are ten in number, and the first eight +are directly connected with the sternum by ribs. + +The =third= of them may be taken as a type. It has a thick cylindrical +centrum, concave in front and convex behind, there is a slight +hypapophysis, and the centrum is suturally united with a strong neural +arch enclosing a narrow neural canal. The neural arch is drawn out +dorsally into a wide truncated neural spine, and laterally into two +prominent transverse processes, with the ends of which the tubercula +of the ribs articulate, while the capitulum articulates in each case +with a step-like facet (fig. 42, A, 3) on the anterior face of the +transverse process. The prezygapophyses (fig. 42, A, 2) are borne on +outgrowths from the bases of the transverse processes, and the +postzygapophyses on outgrowths at the base of the neural spine. + +The thoracic vertebrae behind the third have no hypapophyses, and the +capitular facets gradually come to be placed nearer and nearer the +ends of the transverse processes, at the same time becoming less +prominent; otherwise these vertebrae are just like the third. + +[Illustration FIG. 42. ANTERIOR VIEW OF A, A LATE THORACIC AND B, THE +FIRST SACRAL VERTEBRA OF A YOUNG CROCODILE (_C. palustris_). × 1/3. + + 1. neural spine. + 2. process bearing prezygapophysis. + 3. facet for articulation with the capitulum of the rib. + 4. sacral rib. + 5. surface which is united with the ilium. + 6. concave anterior face of centrum.] + +In the first and second thoracic vertebrae the capitulum of the rib +articulates, not with a facet on the transverse process, but with a +little elevation borne at the line of junction of the centrum and +neural arch. + +THE LUMBAR VERTEBRAE. + +These are five in number, and are precisely like the posterior +thoracic vertebrae, except in the fact that the transverse processes +have no facets for the articulation of ribs. + +THE SACRAL VERTEBRAE. + +These are two in number, and while the centrum of the first is concave +in front (fig. 42, B, 6) and nearly flat behind, that of the second is +flat in front and concave behind. Each has a pair of strong =ribs= +(fig. 42, B, 4) firmly ankylosed in the adult with a wide surface +furnished partly by the centrum, partly by the neural arch. The distal +ends of these ribs are united with the ilia. The character of the +neural spines and zygapophyses is the same as in the thoracic +vertebrae. + +THE CAUDAL VERTEBRAE. + +These are very numerous, about thirty-four in number. The first +differs from all the other vertebrae of the body in having a biconvex +centrum. The succeeding ones are procoelous and are very much like the +posterior thoracic and lumbar vertebrae, having high neural spines and +prominent straight transverse processes. They differ however in having +the neural spines less strongly truncated above, and the transverse +processes arise from the centra and not from the neural arches. When +followed further back the centra and neural spines gradually lengthen +while the transverse processes become reduced, and after the twelfth +vertebra disappear. Further back still the neural spines and +zygapophyses gradually become reduced and disappear, as finally the +neural arch does also, so that the last few vertebrae consist simply +of cylindrical centra. + +Each caudal vertebra, except the first and the last eleven or so, has +a =V=-shaped =chevron bone= attached to the postero-ventral edge of +its centrum. The anterior ones are the largest and they gradually +decrease in size till they disappear. + +B. THE SKULL[90]. + +The skull of the Crocodile is a massive depressed structure +presenting a number of striking characteristics, some of the more +important of which are:-- + +1. All the bones except the mandible, hyoid, and columella are firmly +united by interlocking sutures. In spite of this, however, growth of +the whole skull and of the component bones goes on continuously +throughout life, this growth being especially marked in the case of +the facial as opposed to the cranial part of the skull. + +2. All the bones appearing on the dorsal surface are remarkable for +their curious roughened and pitted character; this feature is +prominent also in many Labyrinthodonts. + +3. The size of the jaws and teeth is very great. + +4. The mandibular condyle is carried back to some distance behind the +occipital condyle. + +5. The occipital plane (see p. 386) of the skull is vertical. + +6. The length of the secondary palate is remarkably great, and the +vomer takes no part in its formation. + +7. The posterior nares are placed very far back, the nasal passages +being as in mammals separated from the mouth by the long secondary +palate. + +8. There is a complicated system of Eustachian passages communicating +at one end with the tympanic cavity and at the other end with the +mouth cavity. + +9. The interorbital septum is mainly cartilaginous, the presphenoidal +and orbitosphenoidal regions remaining unossified. + +The =skull= is divisible into three parts:-- + +(1) the cranium, (2) the lower jaw, (3) the hyoid. + +The =cranium= may again for purposes of description be divided into:-- + +1. the cranium proper or brain case; + +2. the bones connected with the several special sense organs; + +3. the bones of the upper jaw, and suspensorial apparatus. + + +1. THE CRANIUM PROPER OR BRAIN CASE. + +[Illustration FIG. 43. PALATAL ASPECT A, OF THE CRANIUM, B, OF THE +MANDIBLE OF AN ALLIGATOR (_Caiman latirostris_). × 1/3. (Brit. Mus.) + + 1. premaxillae. + 2. maxillae. + 3. palatine. + 4. pterygoid. + 5. posterior nares. + 6. transpalatine. + 7. posterior palatine vacuity. + 8. anterior palatine vacuity. + 9. basi-occipital. + 10. opening of median Eustachian canal. + 11. jugal. + 12. quadratojugal. + 13. quadrate. + 14. dentary. + 15. splenial. + 16. coronoid. + 17. supra-angular. + 18. angular. + 19. articular. + 20. lateral temporal fossa. + 21. openings of vascular canals leading into alveolar sinus.] + +The cartilage and membrane bones of the cranium proper when taken +together can in most vertebrates be seen to be more or less arranged +in three rings or segments called respectively the =occipital=, +=parietal= and =frontal= segments; in the Crocodile however only the +occipital and parietal segments are clearly seen. + +The =occipital segment= consists of four cartilage bones, three of +which together surround the =foramen magnum=. + +The most ventral of these, the =basi-occipital= (figs. 43 and 45, 9), +forms the single convex =occipital condyle= for articulation with the +atlas, bounds the base of the foramen magnum, and is continuous +laterally with two larger bones, the =exoccipitals= (fig. 45, 24), +which meet one another dorsally and form the remainder of the boundary +of the foramen magnum. Each is drawn out externally into a strong +process, which is united below with the quadrate, and above with the +squamosal by a surface seen in a disarticulated skull to be very rough +and splintered. In a longitudinal section the anterior face of the +exoccipital is seen to be closely united with the opisthotic. + +The exoccipital is pierced by a number of foramina, four lying on the +posterior surface. Just external to the foramen magnum is a small +foramen for the exit of the hypoglossal nerve (figs. 44 and 45, XII). +External to this is the foramen for the pneumogastric (fig. 44, X), +while more ventrally still is the foramen (fig. 44, 15) through which +the internal carotid artery enters the skull. Some distance further to +the side, and more dorsally, is a larger foramen which gives passage +to the facial nerve and certain blood-vessels. + +In a median longitudinal section of the skull the hypoglossal foramen +is seen, and just in front of it a small foramen for a vein. Further +forwards the long slit-like opening between the exoccipital and +opisthotic is the =internal auditory meatus= (fig. 45, VIII) through +which the auditory nerve leaves the cranial cavity and enters the +internal ear. + +The =supra-occipital= (fig. 45, 5) is a small bone which takes no part +in the formation of the foramen magnum, and is closely united in front +with the epi-otic. It is characteristic of Crocodiles that all the +bones of the occipital segment have their longer axes placed +vertically, and that they scarcely if at all appear on the dorsal +surface. + +In front of the occipital segment is the =parietal segment=. The +dorsal and ventral portions of the two segments are in contact with +one another, but the lateral portions are widely separated by the +interposition of the =auditory= and =suspensorial bones=. + +The =basisphenoid= (fig. 45, 12) is an unpaired wedge-shaped bone, +united along a deep vertical suture with the basi-occipital. The two +bones are, however, partially separated in the mid-ventral line by a +foramen, the opening of the =median Eustachian canal=, which leads +into a complicated system of Eustachian passages ultimately +communicating with the tympanic cavity. + +The dorsal surface of the basisphenoid is well seen in a section of +the skull, but owing to the way it tapers ventrally, it appears on the +ventral surface only as a very narrow strip of bone wedged in between +the basi-occipital and pterygoids. In a lateral view it is seen to be +drawn out in front into an abruptly truncated process, the =rostrum=, +which forms part of the =interorbital septum.= On the anterior part of +the dorsal surface is a deep pit, the =pituitary fossa= or =sella +turcica=, at the base of which are a pair of foramina, through which +the carotid arteries pass. Dorsolaterally the basisphenoid articulates +with the =alisphenoids=. + +[Illustration FIG. 44. LATERAL VIEW OF THE SKULL OF AN ALLIGATOR +(_Caiman latirostris_). × 1/3. (Brit. Mus.) + + 1. premaxillae. + 2. maxillae. + 3. lachrymal. + 4. prefrontal. + 5. jugal. + 6. postfrontal. + 7. squamosal. + 8. quadrate. + 9. palatine. + 10. pterygoid. + 11. transpalatine. + 12. quadratojugal. + 13. exoccipital. + 14. basi-occipital. + 15. foramen by which carotid artery enters skull. + 16. external auditory meatus. + 17. frontal. + 18. supra-angular. + 19. articular. + 20. dentary. + 21. coronoid. + 22. angular. + III, VI, opening for exit of oculomotor and abducens nerves. + V, foramen ovale. + X, pneumogastric foramen. + XII, hypoglossal foramen.] + +The =alisphenoids= (fig. 45, 13) are a pair of irregular bones which +arise from the basisphenoid antero-laterally, and are united dorsally +with the parietal, frontal, and postfrontals. They bound most of the +anterior part of the brain case, and each presents on its inner face a +deep concavity which lodges the cerebral hemisphere of its side. +Viewed from the ventral side the two alisphenoids are seen to almost +or quite meet one another immediately below the frontal, and then to +diverge, forming an irregular opening--partially closed by cartilage +in the fresh specimen,--through which the optic nerves leave the +cranial cavity. Further back the alisphenoids meet one another for a +narrow area, and then diverge again, so that between each and the +rostrum of the basisphenoid there appears an opening (fig. 44, III, +VI) through which the oculomotor and abducens nerves leave the +cranium. Further back still each is united for a short space with the +basisphenoid, pterygoid and quadrate, and then becomes separated from +the quadrate by a large foramen, the =foramen ovale= (fig. 44, V), +through which the whole of the trigeminal nerve passes out. + +The dorsal portion of the parietal segment is formed by the _parietal_ +(fig. 45, 4), which though double in the embryo, early comes to form a +single bone. It extends over the posterior part of the cranial cavity, +and is continuous in front with the frontal, behind with the +supra-occipital, and laterally with the postfrontals, squamosals, +alisphenoids, pro-otics and epi-otics. It forms the inner boundary of +a large rounded vacuity on the roof of the skull, the =supratemporal +fossa=. + +The =frontal segment= is very imperfectly ossified, there being no +certain representatives of either the ventral member, the presphenoid, +or the lateral members, the orbitosphenoids. On the dorsal side there +is, however, a large development of membrane bones. There is a large +_frontal_ (fig. 45, 3), unpaired, except in the embryo, united behind +with the parietal and postfrontal, and drawn out in front into a long +process which is overlapped by the prefrontals and posterior part of +the nasals. The frontal ends off freely below, owing to the +orbitosphenoidal region being unossified, it forms a considerable part +of the roof of the cranial cavity, but takes no part in the formation +of the wall. + +Each _prefrontal_ (fig. 45, 14) forms part of the inner wall of the +orbit and sends ventralwards a process which meets the palatine. + +The _postfrontals_ (fig. 44, 6) are small bones lying at the sides of +the posterior part of the frontal. Each is united with a number of +bones, on its inner side with the frontal and parietal, behind with +the squamosal, and ventrally with the alisphenoid. It also unites by +means of a strong descending process with an upgrowth from the jugal, +and thus forms a =postorbital bar= separating the orbit from the +lateral temporal fossa. The postfrontal forms also part of the outer +boundary of the supratemporal fossa. + + +2. THE SENSE CAPSULES. + +Skeletal capsules occur in connection with each of the three special +sense organs of sight, of hearing and of smell. + +The =Auditory capsules= and associated bones. + +Three bones, the =epi-otic=, =opisthotic= and =pro-otic=, together +form the auditory or =periotic= capsule of each side. They are wedged +in between the lateral portions of the occipital and parietal segments +and complete the cranial wall in this region. Their relations to the +surrounding structures are very complicated, and many points can be +made out only in sections of the skull passing right through the +periotic capsule. The relative position of the three bones is, +however, well seen in a median longitudinal section. The =opisthotic= +early becomes united with the exoccipital, while the =epi-otic= +similarly becomes united with the supra-occipital, the =pro-otic= +(fig. 45, 7),--seen in longitudinal section to be pierced by the +prominent =trigeminal foramen=--alone remaining distinct throughout +life. The three bones together surround the essential organ of hearing +which communicates laterally with the deep tympanic cavity by the +=fenestra ovalis=. + +The =tympanic cavity=, leading to the exterior by the =external +auditory meatus= (fig. 44, 16), is well seen in a side-view of the +skull; it is bounded on its inner side by the periotic bones, +posteriorly in part by the exoccipital, and elsewhere mainly by the +quadrate. A large number of canals and passages open into it. On its +inner side opening ventro-anteriorly is the =fenestra ovalis=, opening +ventro-posteriorly the =internal auditory meatus= (fig. 45, VIII), +while dorsally there is a wide opening which forms a communication +through the roof of the brain-case with the tympanic cavity of the +other side. On its posterior wall is the prominent foramen through +which the facial nerve passes on its way to its final exit from the +skull through the exoccipital, this foramen is bounded by the +quadrate, squamosal, and exoccipital. + +The opening of the fenestra ovalis is in the fresh skull occupied by +the expanded end of the auditory ossicle, the =columella=, whose outer +end articulates by a concave facet with a trifid =extra-columellar= +cartilage which reaches the tympanic membrane. The lower process of +this extra-columella passes into a cartilaginous rod which lies in a +canal in the quadrate and is during life continuous with Meckel's +cartilage within the articular bone of the mandible. + +The columella and extra-columella are together homologous with the +chain of mammalian auditory ossicles. + + * * * * * + +The =Optic capsules= and associated bones. + +Two pairs of bones are associated with the optic capsules, viz. the +_lachrymals_ and the _supra-orbitals_. The _lachrymal_ (fig. 44, 3) is +a fairly large flattened bone lying wedged in between the maxillae, +nasal, jugal, and prefrontal. It forms a considerable part of the +anterior boundary of the orbit, and is pierced by two foramina. On the +orbital edge is a large hole leading into a cavity within the bone +which lodges the naso-lachrymal sac, and communicates with the narial +passage by a wide second foramen near the anterior end of the bone. +The _supra-orbital_ is a very small loose bone lying in the eyelid +close to the junction of the frontal and prefrontal. + + * * * * * + +The =Olfactory capsules= and associated bones. + +Two pairs of membrane bones, the _vomers_ and _nasals_, are developed +in association with the olfactory organ, but the =mesethmoid= is not +ossified. + +The _vomers_ form a pair of delicate bones, each consisting of a +vertical plate (fig. 45, 15), which with its fellow separates the two +narial passages, and of a horizontal plate which forms much of their +roof. The vomers articulate with one another and with the pterygoids, +palatines, and maxillae. + +The _nasals_ (fig. 45, 2) are very long narrow bones extending along +the middle line from the frontal almost to the anterior nares. They +are continuous laterally with the premaxillae, maxillae, lachrymals and +prefrontals. They form the roof of the narial passages. + +3. THE UPPER JAW AND SUSPENSORIAL APPARATUS. + +These are enormously developed in the Crocodile and are firmly united +to the cranium. It will be most convenient to begin by describing the +bones at the anterior end of the jaw and to work back thence towards +the brain-case. The most anterior bones are the _premaxillae_. The +_premaxillae_ (figs. 44 and 45, 1) are small bones, each bearing five +pairs of teeth, set in separate sockets in their alveolar borders. +They constitute almost the whole of the boundary of the =anterior +nares=, which are confluent with one another and form a large +semicircular opening in the roof of the skull, leading into the wide +narial passage. They are also partially separated from one another in +the ventral middle line, by the small =anterior palatine vacuity= +(fig. 43, A, 8). They form the anterior part of the broad =palate=. +The alveolar border on each side between certain of the teeth is +marked by pits which receive the points of the teeth of the other jaw. +The first pair of these pits in the premaxillae are often so deep as to +be converted into perforations. Pits of the same character occur +between the maxillary and mandibular teeth. + +[Illustration FIG. 45. LONGITUDINAL SECTION THROUGH THE SKULL OF AN +ALLIGATOR (_Caiman latirostris_). × 1/3. (Brit. Mus.) + + 1. premaxilla. + 2. nasal. + 3. frontal. + 4. parietal. + 5. supra-occipital. + 6. epi-otic. + 7. pro-otic. + immediately in front of the figure 7 is the prominent foramen + for the trigeminal nerve. + 8. opisthotic. + 9. basi-occipital. + 10. quadrate. + 11. pterygoid. + 12. basisphenoid. + 13. alisphenoid. + 14. prefrontal. + 15. vomer. + 16. maxilla. + 17. palatine. + 18. dentary. + 19. splenial. + 20. angular. + 21. supra-angular. + 22. articular. + 23. coronoid. + 24. exoccipital. + 25. squamosal. + 26. jugal. + 27. external mandibular foramen. + 28. internal mandibular foramen. + VIII. internal auditory meatus. + XII. hypoglossal foramen.] + +The _maxillae_ (figs. 43, A, 2 and 44, 2) are a pair of very large +bones and bear the remaining teeth of the upper jaw, set in sockets +along their alveolar borders. On the dorsal side each maxillae is +continuous with the premaxillae, nasal, lachrymal, and jugal, while +ventrally it meets its fellow in a long straight suture and forms the +greater part of the long bony palate. The maxillae are separated in the +middle line posteriorly by processes from the palatines, while further +back they meet the transpalatines. The internal or nasal surface, like +that of the premaxillae, is excavated by a deep longitudinal groove, +the =narial passage=. In a ventral view of the skull a number of small +openings (fig. 43, A, 21) are seen close to the alveolar border, these +are the openings of small vascular canals which lead into the +=alveolar sinus=, a passage traversing the maxillae, and transmitting +the superior maxillary branch of the trigeminal nerve and certain +blood-vessels. This alveolar sinus opens posteriorly by the more +external of the two large holes in the maxillae, which lie close to the +anterior edge of the posterior palatine vacuity, to be described +immediately. The more internal of these holes, on the other hand, +leads into a cavity lodging the nasal sac. Behind the maxillae the +completeness of the palate is broken up by the large oval =posterior +palatine vacuities= (fig. 43, A, 7); these are separated from one +another in the middle line by the palatines, and are bounded elsewhere +by the maxillae, transpalatines, and pterygoids. + +The _palatines_ (fig. 43, A, 3) are long and rather narrow bones +interposed between the maxillae in front and pterygoids behind. They +meet one another in a long suture and form much of the posterior part +of the palate, while the whole length of their dorsal surface +contributes to the floor of the narial passage. The dorsal surface of +each bone is also drawn out on its outer side into a prominent ridge +which forms much of the side and roof of the narial passage, being in +contact with the vomer and pterygoid, and at one point by means of a +short ascending process with the descending process of the +prefrontal. + +The _pterygoids_ (figs. 43, A, 4, and 45, 11) are a pair of large +bones, each consisting of a median more or less vertical part, which +becomes ankylosed to its fellow in the middle line early in life, and +of a wide horizontal part which meets the transpalatine. They +completely surround the posterior nares (fig. 43, A, 5) and their +median portions form the whole boundary of the posterior part of the +narial passage, and assist the palatines and vomers in bounding the +middle part. The horizontal parts form the posterior part of the +secondary palate, while the dorsal surface of each looks into the +=pterygoid fossa=, a large cavity lying below the quadrate and +quadratojugal at the side of the skull. The lateral margin adjoining +the transpalatine is in the fresh skull terminated by a plate of +cartilage against which the mandible plays. Dorsally the pterygoid +articulates with the basisphenoid, quadrate, and alisphenoid. + +The _transpalatines_ (fig. 44, 11) connect the pterygoids with the +jugals and maxillae, articulating with each of the three bones by a +long pointed process. The jugal process meets also a down-growth from +the postfrontal. + +The _jugals_ or _malars_ (fig. 44, 5) are long somewhat flattened +bones which are united to the lachrymals and maxillae in front, while +passing backwards each is united behind to the _quadratojugal_ (fig. +44, 12), the two forming the =infratemporal arcade= which constitutes +the external boundary of the orbit and lateral temporal fossa. The +jugal is united below to the transpalatine, and the two bones together +form an outgrowth, which meeting that from the postfrontal forms the +=postorbital bar=, and separates the orbit from the lateral temporal +fossa. The quadratojugals are small bones and are united behind with +the quadrates. + +The =quadrate= (figs. 43, A, 13 and 44, 8) of each side is a large +somewhat flattened bone firmly fixed in among the other bones of the +skull. It is terminated posteriorly by an elongated slightly convex +surface, coated with cartilage in the fresh skull, by which the +mandible articulates with the cranium. The dorsal surface of the +quadrate is flat behind, further forwards it becomes much roughened +and articulates with the exoccipital and squamosal; further forwards +still it becomes marked by a deep groove which forms the floor of the +external auditory meatus and part of the tympanic cavity. The anterior +boundary of the quadrate is extremely irregular, it is united dorsally +with the postfrontal, pro-otic, and squamosal, and more ventrally with +the alisphenoid. The smooth ventral surface looks into the pterygoid +fossa. In front the quadrate forms the posterior boundary of the +supratemporal fossa and foramen ovale, and is continuous with the +alisphenoid, while it sends down a thin plate meeting the pterygoid +and basisphenoid. On the inner side of the dorsal surface of the +quadrate near the condyle, is a small foramen which leads into a tube +communicating with the tympanic cavity, by a foramen lying in front of +and ventral to that for the exit of the facial nerve. By this tube air +can pass from the tympanic cavity into the articular bone of the +mandible. + +The _squamosal_ (fig. 44, 7) meets the quadrate and exoccipital below, +and forms part of the roof of the external auditory meatus, while +above it forms part of the roof of the skull and has a pitted +structure like that of the other bones of the roof. It is continuous +with the postfrontal in front, forming with it the =supratemporal +arcade= which constitutes the outer boundary of the supratemporal +fossa. It meets also the parietal on its inner side, forming the +=post-temporal bar=, the posterior boundary of the supratemporal +fossa. + +It may be useful to recapitulate the large vacuities in the surface of +the Crocodile's cranium. + +_Dorsal surface._ + +1. =The Supratemporal fossae=. Each is bounded internally by the +parietal, behind by the =post-temporal bar= formed by the parietal and +squamosal, and externally by the =supratemporal arcade= formed by the +squamosal and postfrontal. The postfrontal meets the parietal in front +and forms the anterior boundary of the supratemporal fossa. + +2. The =Lateral temporal= or =infratemporal fossae=. These lie below +and to the outer side of the supratemporal fossae. Each is bounded +dorso-internally by the supratemporal arcade; and behind by a +continuation of the post-temporal bar formed by the quadrate and +quadratojugal. The external boundary is the =infratemporal arcade= +formed of the quadratojugal and jugal, while in front the fossa is +separated from the orbit by the =postorbital bar= formed by the +junction of outgrowths from the postfrontal and jugal. + +3. The =Orbits=. Each is bounded behind by the postorbital bar, +externally by the jugal forming a continuation of the infratemporal +arcade, in front by the lachrymal, and internally by the frontal and +prefrontal. + +4. The =Anterior nares=. These form an unpaired opening bounded by the +premaxillae. + +_Posterior surface._ + +5. The =Foramen magnum=. The exoccipitals form the chief part of its +boundary, but part of the ventral boundary is formed by the +basi-occipital. + +6. The =Pterygoid fossae=. These form a pair of large cavities at the +sides of the occipital region of the skull. The dorsal boundary is +formed by the quadrate and quadratojugal, the ventral by the +pterygoid, the internal chiefly by the quadrate, pterygoid, +alisphenoid, and basisphenoid. The transpalatine forms a small part of +the external boundary which is incomplete. + +_Ventral surface._ + +7. The =Posterior nares=. These form a median unpaired opening (fig. +43, A, 5) bounded by the pterygoids. + +8. The =Posterior palatine vacuities=. Each is bounded by the maxillae +in front, the maxillae and transpalatine externally, the transpalatine +and pterygoid behind, and the palatine on the inner side (fig. 43, A, +7). + +9. The =Anterior palatine vacuity=. This is unpaired and is bounded by +the premaxillae (fig. 43, A, 8). + + +(_b_) THE LOWER JAW OR MANDIBLE. + +The mandible is a strong compact bony structure formed of two halves +or =rami=, which are suturally united at the symphysis in the middle +line in front. Each ramus is formed of six separate bones. + +The most anterior and largest of these is the _dentary_ (figs. 44, 20, +and 45, 18), which forms the symphysis, and greater part of the +anterior half of the jaw, and bears along the outer part of its dorsal +border a number of sockets or =alveoli= in which the teeth are placed. +Lying along the inner side of the dentary is a large splint-like bone, +the _splenial_ (fig. 45, 19), which does not extend so far forwards as +the symphysis, and is separated from the dentary posteriorly by a +large cavity. Forming the lower part of all the posterior half of the +jaw is the large _angular_ (figs. 44, 22, and 45, 20), which underlies +the posterior part of the dentary in front and sends a long process +below that bone to the splenial. On the inner side of the jaw there is +an oval vacuity, the =internal mandibular foramen= (fig. 45, 28), +between the angular and the splenial; through this pass blood-vessels +and branches of the inferior dental nerve. Lying dorsal to the angular +is another large bone, the _supra-angular_ (figs. 44, 18, and 45, 21). +It extends back as far as the posterior end of the jaw and forwards +for some distance dorsal to the dentary and splenial. It forms part of +the posterior margin of a large vacuity, the =external mandibular +foramen=, which is bordered above and in front by the dentary and +below by the angular; it gives passage to the cutaneous branch of the +inferior dental nerve. The concave surface for articulation with the +mandible and much of the posterior end of the jaw is formed by a short +but solid bone, the =articular= (fig. 45, 22), which in young skulls +rather readily becomes detached. The remaining mandibular bone is the +_coronoid_ (fig. 45, 23), a very small bone of irregular shape +attached to the angular below, and to the supra-angular and splenial +above. + +(_c_) THE HYOID. + +The hyoid of the Crocodile consists of a wide flattened plate of +cartilage, the =basilingual plate= or =body of the hyoid=, and a pair +of =cornua=. + +The =basilingual plate= (fig. 53, 1) is rounded anteriorly and marked +by a deep notch posteriorly. The =cornua= (fig. 53, 3), which are +attached at a pair of notches near the middle of the outer border of +the basilingual plate, are partly ossified, but their expanded ends +are formed of cartilage. They pass at first backwards and then upwards +and inwards. They are homologous with part of the first branchial +arches of Selachians. + +The columella and extra-columella have been already described (p. +251). + +C. THE RIBS AND STERNUM. + +=Thoracic ribs.= + +The Crocodile has ten pairs of =thoracic ribs=, all except the last +one or two of which consist of three parts,--a vertebral rib, an +intermediate rib and a sternal rib. + +Of the =vertebral ribs= the third may be taken as a type, it consists +of a curved bony rod which articulates proximally with the transverse +process of the vertebra by two facets. The terminal one of these, the +=capitulum= or =head=, articulates with a notch on the side of the +transverse process; the other, the =tuberculum=, which lies on the +dorsal surface a short distance behind the head, articulates with the +end of the transverse process. From near the distal end an imperfectly +ossified uncinate =process= (see p. 190) projects backwards. + +The =intermediate ribs= are short and imperfectly ossified; they are +united with the =sternal ribs= (fig. 46, 3), which are large, +flattened, likewise imperfectly ossified structures, and articulate at +their distal ends with a pair of long divergent =xiphisternal horns= +(fig. 46, 5), which arise from the posterior end of the sternum +proper. The last pair of sternal ribs are attached to the preceding +pair, not to the xiphisternal horns. + +The first and second vertebral ribs differ from the others in the fact +that the tuberculum forms a fairly long outstanding process. + + +=Cervical ribs.= + +Movable ribs are attached to all the cervical as well as to the +thoracic vertebrae. Those borne by the atlas and axis are long, narrow +structures attached by a fairly broad base, and tapering gradually. +The ribs borne by the third to seventh cervical vertebrae are shaped +like a =T= with a double base, one limb of which, corresponding to the +tuberculum (fig. 41, 7), articulates with a short transverse process +arising from the neural arch, while the other, corresponding to the +capitulum, articulates with a surface on the centrum. The ribs +attached to the eighth and ninth cervical vertebrae are intermediate +in character between the =T=-shaped ribs and the ordinary thoracic ribs. +The anterior limb of the =T= is shortened, the posterior one is drawn +out, forming the shaft of the rib. The distal portion of the rib of +the ninth cervical vertebra is unossified. + +The =Sacral ribs= have been described in connection with the sacral +vertebrae. + + +THE STERNUM. + +The =sternum= of Crocodiles is a very simple structure, consisting of +a plate of cartilage (fig. 46, 2) lying immediately dorsal to the +interclavicle, and drawn out posteriorly into a pair of long +=xiphisternal horns= (fig. 46, 5). + + +THE ABDOMINAL SPLINT RIBS. + +Lying superficially to the recti muscles of the ventral body-wall, +behind the sternal ribs, are seven or eight series of slender curved +bones, the _abdominal ribs_ (fig. 46, 4). Each series consists of four +or more bones, arranged in a =V=-like form with the angle of the =V= +directed forwards. They show a considerable amount of variability in +number and character. They are really membrane bones, and are in no +way homologous with true ribs, but correspond rather with the more +posterior of the bones constituting the plastron of Chelonia. + +[Illustration FIG. 46. STERNUM AND ASSOCIATED MEMBRANE BONES OF A +CROCODILE (_C. palustris_) × 1/3. (Brit. Mus.) + +The last pair of abdominal ribs which are united with the epipubes by +a plate of cartilage have been omitted. + + 1. interclavicle. + 2. sternum. + 3. sternal rib. + 4. abdominal splint rib. + 5. xiphisternal horn.] + + +2. THE APPENDICULAR SKELETON. + +This includes the skeleton of the two pairs of limbs and their +respective girdles. + +[Illustration: FIG. 47. LEFT HALF OF THE PECTORAL GIRDLE OF AN +ALLIGATOR (_Caiman latirostris_) × 2/3. (Brit. Mus.) + + 1. scapula. + 2. coracoid. + 3. interclavicle. + 4. glenoid cavity.] + + +THE PECTORAL GIRDLE. + +The pectoral girdle of the Crocodile is less complete than is that of +most reptiles. It consists of a dorsal bone, the =scapula=, and a +ventral bone, the =coracoid=, with a median unpaired element, the +_interclavicle_; but there is no separate representative either of the +clavicle or precoracoid. + +The =scapula= (fig. 47, 1) is a large bone, flattened and expanded +above where it is terminated by an unossified margin, the +=suprascapula=, and thickened below where it meets the coracoid. The +scapula forms about half the =glenoid cavity= (fig. 47, 4) for +articulation with the humerus, and has the lower part of its anterior +border drawn out into a roughened ridge. + +The =coracoid= (fig. 47, 2) is a flattened bone, much expanded at +either end; it bears on its upper posterior border a flattened surface +which forms half the glenoid cavity, and is firmly united to the +scapula at its dorsal end. Its ventral end meets the sternum. + +The _interclavicle_ (figs. 46, 1, and 47, 3) is a long narrow +blade-shaped bone lying along the ventral side of the sternum; about a +third of its length projects beyond the sternum in front. + + +THE ANTERIOR LIMB. + +This is as usual divisible into three portions, the upper arm, +fore-arm and manus. + +The =upper arm= or =brachium= contains one bone, the =humerus.= + +The =humerus= (fig. 48, A, 1) is a fairly long stout bone, +considerably expanded at either end. The proximal end or head is +evenly rounded and is formed by an epiphysis ossifying from a centre +different from that forming the shaft. It articulates with the glenoid +cavity. The shaft bears on the flexor surface, at some little distance +behind the head, a prominent rounded protuberance, the =deltoid +ridge.= The distal end or trochlea is also formed by an epiphysis and +is partially divided by a groove into two convex surfaces; it +articulates with the two bones of the fore-arm, the radius and ulna. + +[Illustration FIG. 48. A, RIGHT ANTERIOR, AND B, RIGHT POSTERIOR LIMB +OF A YOUNG ALLIGATOR (_Caiman latirostris_). (Brit. Mus.) + +A × 1/2. B × about 1/3.] + + 1. humerus. + 2. radius. + 3. ulna. + 4. radiale. + 5. ulnare. + 6. pisiform. + 7. patch of cartilage representing carpalia 1 and 2; between it + and the radiale should be another flattened patch, the centrale. + 8. carpalia 3, 4, and 5 (fused). + 9. first metacarpal. + 10. proximal phalanx of second digit. + 11. second phalanx of fifth digit. + 12. femur. + 13. tibia. + 14. fibula. + 15. tibiale, intermedium and centrale (fused). + 16. fibulare. + 17. tarsalia 1, 2, and 3 (fused). + 18. tarsalia 4 and 5 (fused). + 19. first metatarsal. + 20. ungual phalanx of second digit. + 21. fifth metatarsal. + +The =radius= and =ulna= are nearly equal in size and each consists of +a long shaft terminated at either end by an epiphysis. + +The =radius= (fig. 48, A, 2) or pre-axial bone is slightly the smaller +of the two. It has a straight cylindrical shaft and is slightly and +nearly evenly expanded at either end. The proximal end which +articulates with the humerus is flat or slightly concave, the distal +end which articulates with the carpus is slightly convex. + +The =ulna= (fig. 48, A, 3) or postaxial bone is a curved bone rather +larger than the radius. Its proximal end is large and convex, but is +not drawn out into an olecranon process. + +The =Manus= consists of the =carpus= or =wrist=, and the =hand=. + +The =Carpus=. This differs considerably from the more primitive type +met with in the Turtle. It consists of six elements arranged in a +proximal row of three and a distal row of two, with one intervening. +The bones of the proximal row are the radiale, the ulnare, and the +pisiform. The =radiale= (fig. 48, A, 4) is the largest bone of the +carpus: it is a somewhat hour-glass shaped bone, with its ends formed +by flattened epiphyses. It articulates by its proximal end with the +whole of the radius, and partly also with the ulna, and by its distal +end with the centrale. + +The =ulnare= (fig. 48, A, 5) is a smaller bone, also somewhat +hour-glass shaped; it articulates proximally with the pisiform and +radiale, not quite reaching the ulna. The third bone of the proximal +row is the =pisiform= (fig. 48, A, 6), an irregular bone, articulating +with the ulna, radiale, and fifth metacarpal. The =centrale= is a +flattened cartilaginous element applied to the distal surface of the +radiale. + +The distal row of carpals consists of two small structures. The first +of these forms a small cartilaginous patch, which is wedged in between +the first and second metacarpals, the centrale and the bone +representing carpalia 3, 4 and 5; this cartilaginous patch represents +=carpalia 1 and 2= (fig. 48, A, 7). The bone representing =carpalia 3, +4 and 5= is a good deal larger, rounded, and well-ossified; it +articulates with the ulnare, the pisiform, and the third, fourth, and +fifth metacarpals. + +The =hand=. Each of the five digits consists of an elongated +metacarpal, terminated at each end by an epiphysis, and of a varying +number of phalanges. The terminal phalanx of each digit has an +epiphysis only at its proximal end, the others have them at both ends. + +The first digit, or =pollex=, is the stoutest, and has two phalanges, +the second has three, the third four, the fourth three, and the fifth +two. The terminal phalanx of each of the first three digits is pointed +and sheathed in a horny claw; and is also marked by a pair of +prominent lateral grooves. + +THE PELVIC GIRDLE. + +The pelvic girdle of the Crocodile consists of four parts, a dorsal +element, the =ilium=, an anterior ventral element, the =pubis=, a +posterior ventral element, the =ischium=, and an accessory anterior +ventral element, the =epipubis=. All except the epipubis take part in +the formation of the =acetabulum=, which is perforated by a prominent +hole. + +The =ilium= (fig. 49, 1) is a thick strong bone, firmly united on its +inner side with the two sacral ribs. Its dorsal border is rounded, its +ventral border bears posteriorly two irregular surfaces, completed by +epiphyses, which are united respectively with the ischium and pubis. + +The =ischium= (fig. 49, 2)--the largest bone of the pelvis, is +somewhat contracted in the middle and expanded at either end. Its +proximal end, which is formed by an epiphysis, bears two surfaces, one +of which is united to the ilium, while the other forms part of the +acetabulum. The anterior border is also drawn out dorsally into a +strong process, which is terminated by a convex epiphysis, and is +united to the pubis. The ventral end of the ischium forms a flattened +blade, meeting its fellow in a median symphysis. + +The =pubis= (fig. 49, 3) is much smaller than either the ilium or +ischium; it forms a small patch of unossified cartilage, interposed +between the anterior parts of the ilium and ischium. + +[Illustration FIG. 49. PELVIS AND SACRUM OF AN ALLIGATOR (_Caiman +latirostris_) × 1/2. (Brit. Mus.) + + 1. ilium. + 2. ischium. + 3. true pubis. + 4. epipubis (so-called pubis). + 5. acetabular foramen. + 6. neural spines of sacral vertebrae. + 7. symphysis ischii. + 8. process bearing prezygapophysis.] + +The =epipubis= (fig. 49, 4) is a large bone with a thickened proximal +end, which is loosely articulated to the ischium, and a flattened +expanded distal end, which is united with its fellow, and with the +last pair of abdominal ribs by a large plate of cartilage. This bone +is generally described as the pubis. + + +THE POSTERIOR LIMB. + +This is as usual divisible into three portions, the =thigh=, the +=crus= or =shin=, and the =pes=. + +The =thigh= is formed by the =femur= (fig. 48, B, 12), a moderately +long stout bone, not unlike the humerus; it articulates with the +acetabulum by a fairly prominent rounded =head=. The distal end +articulating with the tibia and fibula is also expanded, and is +partially divided into equal parts by anterior and posterior grooves. +The flexor surface bears a fairly prominent trochanteric ridge. Each +end of the femur is formed by an epiphysis. + +The =crus= or =shin= includes two bones, the =tibia= and =fibula=. +Both are well developed, but the tibia is considerably the larger of +the two. + +The =tibia= (fig. 48, B, 13) is a strong bone with a flattened +expanded proximal end articulating with almost the whole of the end of +the femur, and a similarly expanded distal end articulating with a +bone representing the fused astragalus and centrale. + +The =fibula= (fig. 48, B, 14) is flattened proximally, and articulates +with only quite a small part of the femur, while distally it is more +expanded, and articulates with the fibulare (calcaneum) and with a +facet on the side of the fused astragalus and centrale. + +The =Pes= consists of the =tarsus= or =ankle=, and the =foot=. + +The =Tarsus=. This, like the carpus, is much reduced and modified from +the primitive condition. It consists of only four bones, arranged in +two rows of two each. The two bones of the proximal row are much +larger than are those of the distal row. The pre-axial of them (fig. +48, B, 15) representing the fused =astragalus= (tibiale and +intermedium) and =centrale=, articulates proximally with the tibia and +fibula, and distally with the first metatarsal, and a small bone +representing the first three tarsalia. The postaxial bone, the +=calcaneum= (fibulare) (fig. 48, B, 16), is drawn out into a +prominent posterior process forming a heel such as is almost unknown +elsewhere except in mammals. It articulates with the fibula, the +tibiale-centrale, and distally with a bone representing the fourth and +fifth tarsalia, and with the fifth metatarsal. + +The two bones forming the distal row of tarsals are both small and +rounded; one represents the first three tarsalia fused together, the +other tarsalia 4 and 5. + +The =Foot=. The =foot= has five digits, but the fifth is much reduced, +consisting only of a short metatarsal. The first four =metatarsals= +are all long bones, slightly expanded at each end, and terminated by +small epiphyses. The first digit has two phalanges, the second three, +the third four, and the fourth five. The terminal or =ungual phalanx= +in each instance is grooved and pointed, and in the case of the first +three digits bears a horny claw. The ungual phalanx progressively +decreases in size from the first to the fourth. The fifth digit +consists only of a small, somewhat square metatarsal (fig. 48, B, 21), +attached to the bone representing the fused fourth and fifth +tarsalia. + + +FOOTNOTES: + +[90] Free use has been made of L.C. Miall's _Studies in Comparative +Anatomy_, I., _The Skull in Crocodilia_, London, 1878. See also W.K. +Parker, _Tr. Z.S._, vol. XI. 1885, p. 263. + + + + +CHAPTER XVI. + +GENERAL ACCOUNT OF THE SKELETON IN REPTILES. + + +EXOSKELETON. + +The exoskeleton both epidermal and dermal is exceedingly well +developed in reptiles. + + +EPIDERMAL EXOSKELETON. + +This generally has the form of overlapping horny =scales= which invest +outgrowths of the dermis, and are found covering the whole body in +most Rhynchocephalia, Ophidia, and Lacertilia, and many Crocodilia. In +the Ophidia the ventral surface of the tail is commonly covered by a +double row of broad scales, while the ventral surface of the precaudal +part of the body is covered by a single row. In the burrowing snakes +(Typhlopidae) and some sea snakes (Hydrophidae) these broad scales do +not occur, the scales of the ventral surface being similar to those of +the dorsal. + +In the Chelonia with the exception of _Dermochelys_, _Trionyx_ and +their allies there is a well-developed system of horny shields having +a regular arrangement which has been described in the account of the +Turtle's skeleton[91]. + +The =rattle of the rattlesnake= is an epidermal structure formed of +several loosely articulated horny rings, produced by the modification +of the epidermal covering of the end of the tail, which instead of +being cast off when the rest of the outer skin is shed is retained +loosely interlocked with the adjoining ring or joint. New rings are +thus periodically added to the base of the rattle, and in old animals +the terminal ones wear away and are lost. + +Horny claws occur on the ends of some or all of the digits in most +living reptiles. + +Owen's Chameleon bears three epidermal horns, one arising from the +nasal and two from the frontal region. + +In the Chelonia, some of the Theromorpha such as _Udenodon_ and +_Dicynodon_, probably also in the Pterosauria and _Polyonax_ among the +Dinosaurs, the jaws are more or less cased in horny beaks. The horny +beaks of Chelonia are variable; sometimes they have cutting edges, +sometimes they are denticulated, sometimes they are adapted for +crushing. + + +DERMAL EXOSKELETON. + +Nearly all Crocodilia, many Dinosauria, some Rhynchocephalia and +Pythonomorpha, and some Lacertilia such as _Tiliqua_, _Scincus_ and +_Anguis_ have a dermal exoskeleton of bony scutes, developed below and +corresponding in shape to the epidermal scales. Sometimes as in +_Caiman sclerops_, _Jacare_ and _Teleosaurus_, the scutes completely +invest the body, being so arranged as to form a dorsal and a ventral +shield, and a continuous series of rings round the tail. In +_Crocodilus_ they are confined to the dorsal surface, and in +_Alligator_ to the dorsal and ventral surfaces. The scutes of some +extinct forms articulate with one another by a peg and socket +arrangement as in some Ganoid fish. + +The =carapace= of most Chelonia is a compound structure, being partly +endoskeletal and formed from the ribs and vertebrae, partly from +plates derived from the dermal exoskeleton. The common arrangement is +seen in fig. 36. All the surface plates are probably exoskeletal in +origin, but united with the ventral surfaces of the costal and neural +plates respectively are the expanded ribs and neural arches of the +vertebrae. + +The =plastron= in the common genus _Chelone_ (fig. 37) includes nine +plates of bone, one unpaired and four pairs; they will be referred to +in connection with the ribs and pectoral girdle. + +In the Leathery Turtle (_Dermochelys_) the carapace and plastron +differ completely from those of any other living form. The carapace +consists of a number of polygonal ossifications fitting closely +together and altogether distinct from the vertebrae and ribs. The +plastron is imperfectly ossified, and not united with the pelvis, and +the whole surface of both carapace and plastron is covered with a +tough leathery skin, without horny shields. + +Some of the extinct Dinosauria have an enormously developed dermal +exoskeleton. Thus in _Stegosaurus_ and _Omosaurus_ the dorsal surface +is provided with flattened plates or with spines reaching a length of +upwards of two feet. In _Polacanthus_ the posterior part of the body +is protected by a bony shield somewhat recalling that of the little +armadillo _Chlamydophorus_. No exoskeleton is known in Ichthyosauria, +Sauropterygia, Pterosauria, many Dinosauria and Theromorpha, and some +Lacertilia, such as _Chamaeleon_ and _Amphisbaena_. + + +TEETH. + +The teeth of reptiles are generally well developed, and in the great +majority of forms are simple conical structures, uniform in character, +generally somewhat recurved, and often with serrated edges. Another +common type of tooth is that with a laterally compressed triangular +crown provided with a double cutting edge which may or may not be +serrated. The teeth are mainly formed of dentine, with usually an +external layer of enamel, and often a coating of cement on the root. +Vasodentine is found below the dentine in _Iguanodon_. The teeth of +reptiles never have the enamel deeply infolded, nor do they have +double roots. + +Teeth may be present not only on the jaw-bones, but as in many +_Squamata_, also on the palatines, pterygoids or vomers. The method +by which they are attached to the bones varies much. Sometimes as in +_Iguana_ and some other lizards, they are pleurodont[1], sometimes +they are acrodont[92], as in the Rhynchocephalia, Pythonomorpha, +Ophidia and some Lacertilia such as _Agama_. Again they may be set in +a continuous groove as in the Ichthyosauria and young Crocodilia. +Finally the teeth may be _thecodont_ or placed in distinct sockets as +in the Theromorpha, Sauropterygia, adult Crocodilia, Sauropoda and +Theropoda. In _Iguanodon_ the teeth are set in shallow sockets in a +groove one side of which is higher than the other; the method of +attachment thus shows points of resemblance to the thecodont +condition, the pleurodont condition, and that met with in the +Ichthyosauria. + +In _Ichthyosaurus_ the teeth are marked by a number of vertical +furrows, and it is from a furrow of this nature greatly enlarged and +converted into a tube that the channel down which flows the poison of +venomous snakes is derived. + +In most reptiles the dentition is more or less homodont. The only +reptiles in which a definite heterodont dentition is known are the +extinct Theromorpha, and in them the teeth vary greatly. Thus +_Udenodon_ is toothless, the jaws having been probably cased in a +horny beak. In _Dicynodon_ the jaws are likewise toothless with the +exception of a pair of permanently growing tusks borne by the maxillae. +_Dicynodon_ is the only known reptile whose teeth have permanently +growing pulps. In _Pariasaurus_ the teeth are uniform and very +numerous, and though placed in distinct sockets are ankylosed to the +jaw. In _Galesaurus_ and _Cynognathus_ three kinds of teeth can be +distinguished, slender conical incisor-like teeth, large canine-like +teeth, and cheek teeth with two or three cusps. The teeth in +_Galesaurus_ are confined to the jaws, in _Placodus_ and its allies, +however, large flat crushing teeth are attached to the palatines as +well as to the jaw-bones, and in _Pariasaurus_ the vomer, palatine and +pterygoid all bear teeth as well as the jaw bones. The upper jaw of +_Sphenodon_ and other Rhynchocephalia is provided with two parallel +rows of teeth, one borne on the maxillae and one on the palatines, the +mandibular teeth bite in a groove between these two rows. The bone of +the jaws in _Sphenodon_ is so hard that when the teeth get worn away, +it can act as a substitute. In the young _Sphenodon_ the vomers bear +teeth, as they do also in _Proterosaurus_. + +There is generally a continuous succession of teeth throughout life, +the new tooth coming up below, or partly at the side of the one in +use, and causing the absorption of part of its wall or base. In this +way the new tooth comes to lie in the pulp cavity of the old one. This +method of succession is well seen in the Crocodilia. + +[Illustration FIG. 50. PREPARATION OF PART OF THE RIGHT MANDIBULAR +RAMUS OF _Crocodilus palustris_ × 1/2. (Brit. Mus.) + + 1. tooth in use. + 2. fairly old germ of future + tooth. + 3. symphysial surface of the + mandible.] + +Teeth have been detected in embryos of _Trionyx_, but otherwise no +teeth are known to occur in Chelonia, or in _Pteranodon_ +(Pterosauria), while the anterior part of the jaw is edentulous in +_Iguanodon_, _Polyonax_ and some other Dinosaurs, and in +_Rhamphorhynchus_. + + +ENDOSKELETON. + + +VERTEBRAL COLUMN. + +The vertebral column is commonly divisible into the usual five +regions, but in the Ophidia, Ichthyosauria, and Amphisbaenidae among +Lacertilia, only into caudal and precaudal regions. In the Chelonia +there are no lumbar vertebrae. + +The form of the vertebral centra is very variable. A large proportion +of extinct reptiles,--several entire orders,--and the earlier and more +primitive forms in some of the other groups have amphicoelous +vertebrae. Vertebrae of this type occur in the Theromorpha, +Ichthyosauria, most Sauropterygia and Rhynchocephalia, and many +Dinosauria, also in some of the early Crocodilia such as _Belodon_, +_Teleosaurus_ and _Goniopholis_, and the Geckonidae among Lacertilia. + +The majority of living reptiles have procoelous vertebrae. Thus they +occur in the Lacertilia (excluding the Geckos), the Ophidia, and the +Crocodilia, also among extinct forms in the Pterosauria and many +Dinosauria. On the other hand some Dinosauria such as _Iguanodon_ have +opisthocoelous cervical vertebrae, while others have opisthocoelous +thoracic vertebrae. The vertebrae of the Ceratopsidae and some +Sauropterygia, the thoracic vertebrae of _Iguanodon_, and the sacral +vertebrae of Crocodilia have flat centra. The first caudal vertebra of +modern Crocodilia is biconvex, and in the Chelonia all types of +vertebral centra are found. The cervical vertebrae of _Sphenodon_ are +noticeable for the occurrence of a small pro-atlas, which may +represent the neural arch of a vertebra in front of the atlas. + +In most reptiles the vertebrae are fully ossified, but in some of the +more primitive forms the notochord persists in the centre of the +vertebra (i.e. intervertebrally), this is the case for instance in +many of the Theromorpha and Rhynchocephalia, and also in the Geckos. +In other reptiles it persists longest intravertebrally. + +The centrum of each of the caudal vertebrae of most Lacertilia is +traversed by an unossified septum along which it readily breaks. + +Chevron bones occur below the caudal vertebrae in Lacertilia, +Chelonia, Ichthyosauria, many Dinosauria, and _Sphenodon_, +articulating with quite the posterior part of the centrum which bears +them. In Lacertilia and Crocodilia (fig. 41, 3) the axis has a +well-marked odontoid process. The ventral portions of the +intervertebral discs are sometimes ossified, forming wedge-shaped +inter centra, as in Geckos, and the cervical vertebrae of _Sphenodon_. + +In snakes, Theropod Dinosaurs, and the iguanas among lizards, the +neural arches are provided with _zygosphenes_, and _zygantra_. + +The neural arches are usually firmly ankylosed to the centra, but in +the Crocodilia and some Chelonia, Sauropterygia, and Dinosauria, the +suture between the centrum and neural arch persists at any rate till +late in life. In the Ichthyosauria the neural arches were united to +the centra by cartilage only. + +The thoracic vertebrae of some of the Theromorpha (_Dimetrodon_) are +remarkable for the extraordinary development of the neural spine, and +those of Chelonia for the absence of transverse processes. + +In living reptiles the number of sacral vertebrae is nearly always +two, but in the Theromorpha, Dinosauria, and Pterosauria, as many as +five or six bones may be ankylosed together in the sacral region. In +Crocodiles the two halves of the pelvis sometimes articulate with +different vertebrae. The vertebrae of some of the great Sauropoda are +remarkably hollowed out, having a large vacuity on each side of the +centrum communicating with a series of internal cavities. The whole +structure of these vertebrae shows a combination of great strength +with lightness. + + +THE SKULL. + +The reptilian skull is well ossified and the bones are noticeable for +their density. The true cranium is often largely concealed by a +secondary or false roof of membrane bones, which is best seen in the +Ichthyosauria and some of the Chelonia. In other reptiles the false +roof is more or less broken up by vacuities exposing the true cranial +walls. The ethmoidal region is the only one in which much of the +primordial cartilaginous cranium remains. The lateral parts of the +sphenoidal region are also as a rule not well ossified. + +In some reptiles, such as most Lacertilia and Chelonia, the orbits are +separated only by the imperfect interorbital septum, while in others, +such as the Ophidia, Crocodilia and Amphisbaenidae, the cranial cavity +extends forwards between the orbits. + +In the occipital region all four bones are ossified. The great +majority of reptiles have a single convex occipital condyle, but some +of the Theromorpha such as _Cynognathus_ have two distinct condyles as +in mammals. Sometimes, as in Chelonia, Ophidia and Lacertilia, the +exoccipitals, as well as the basi-occipital, take part in the +formation of the single condyle; sometimes, as in Crocodiles, it is +formed by the basi-occipital alone, as in birds. The relations of the +bones to the foramen magnum vary considerably, in Chelonia the +basi-occipital generally takes no part in bounding it, and in the +Theromorpha, Crocodilia, and Ophidia, the supra-occipital is excluded. +The parietals are paired in Geckos and Chelonia alone among living +forms, and in the extinct Ichthyosauria and some Theromorpha; in all +other reptiles they are united. + +The frontals are paired in Ichthyosauria (fig. 32, 5), Chelonia, +Ophidia, _Sphenodon_ (fig. 52, B, 4) and some extinct crocodiles, such +as _Belodon_. They are completely fused in living Crocodilia and some +Lacertilia and Dinosauria. In the gigantic _Polyonax_ they are drawn +out into a pair of enormous horns, and the parietals and squamosals +are greatly expanded behind. + +An interparietal foramen occurs in the Theromorpha, the Ichthyosauria +(fig. 32, 10), _Sphenodon_, the Sauropterygia and most Lacertilia. +The posterior part of the skull is curiously modified in some +Chamaeleons, the parietals and supra-occipitals being drawn out into a +backwardly-projecting sagittal crest which unites with the two +prolongations from the squamosals. In other Chamaeleons (_C. bifidus_) +prolongations of the prefrontals and maxillae form large +forwardly-projecting bony processes. + +The roof of the skull is characterised by the development of +prefrontals and postfrontals, which lie respectively near the anterior +and posterior extremity of the orbit. In Theromorpha, Squamata, +Crocodilia, and some Dinosauria lachrymals are developed. There is a +ring of bones in the sclerotic in the Ichthyosauria (fig. 32, 15), the +Metriorhynchidae among Crocodiles and some Rhynchocephalia, +Dinosauria, and Pterosauria. + +The pro-otic lies in front of the exoccipital and together with the +opisthotic forms the hind border of the fenestra ovalis. In Chelonia +the opisthotic remains separate, in all other living reptiles it fuses +with the exoccipital. The epi-otic fuses with the supra-occipital. + +The parasphenoid, so important in Ichthyopsids, has very often +disappeared completely; it is present, however, in the Ichthyosauria, +the Plesiosauridae, and a number of Squamata, in many Ophidia its +anterior part forming the base of the interorbital septum. + +In the Plesiosauridae and most Lacertilia, but not in the +Amphisbaenidae, a slender bone, the epipterygoid, occurs uniting the +parietal or the anterior end of the pro-otic with the pterygoid. A +homologous arrangement occurs in the Ichthyosauria and some Chelonia. + +In most reptiles a transpalatine occurs, connecting the maxillae with +the pterygoid, but this is absent in the Chelonia, and some +Dinosauria, and in the Typhlopidae among snakes. + +The quadrate is always well developed, and except in the Squamata is +firmly fixed to the surrounding bones. The Chamaeleons also, among the +Squamata, have a fixed quadrate, and in them too the quadratojugal is +absent. Separate nasal bones do not occur in any living Chelonia. + +The vomers are generally paired as in Squamata, sometimes unpaired as +in Chelonia. + +[Illustration FIG. 51. DORSAL (TO THE LEFT) AND VENTRAL (TO THE RIGHT) +VIEWS OF THE SKULL OF THE COMMON SNAKE (_Tropidinotus natrix_). (After +PARKER.) + + 1. premaxillae (fused). + 2. anterior nares. + 3. nasal. + 4. prefrontal. + 5. frontal. + 6. parietal. + 7. maxillae. + 8. transpalatine. + 9. palatine. + 10. pterygoid. + 11. pro-otic. + 12. exoccipital. + 13. supra-occipital. + 14. opisthotic. + 15. epi-otic. + 16. quadrate. + 17. parasphenoid. + 18. basisphenoid. + 19. basi-occipital. + 20. occipital condyle. + 21. splenial. + 22. dentary. + 23. angular. + 24. articular. + 25. supra-angular. + 26. coronoid. + 27. vomer. + 28. squamosal. + + IX, X foramina for the ninth + and tenth cranial nerves.] + +The disposition of the bones of the jaws is subject to much +modification in the Ophidia in order to adapt them for swallowing very +large prey. The arrangements again differ greatly in the venomous and +non-venomous snakes. In the non-venomous snakes, such as _Python_ and +_Tropidonotus_, the palatine is large and is fixed to the pterygoid +which extends outwards (fig. 51, 10) so as to be united to the +quadrate, and is at the same time firmly connected by the +transpalatine with the maxillae. The quadrate is united to the +squamosal, which is loosely attached to the cranium. The premaxillae is +moderately developed and bears teeth, and the maxillae forms a long bar +loosely connected with the rest of the skull. The rami of the mandible +are united only by an extremely elastic ligament. It is as regards the +maxillae and premaxillae that the skulls of venomous and non-venomous +snakes differ most. In the rattlesnake (_Crotalus_) and other venomous +snakes the premaxillae is extremely small and toothless. The maxillae is +small and subcylindrical, and is movably articulated to the lachrymal, +which also is capable of a certain amount of motion on the frontal. +The maxillae is connected by means of the transpalatine with the +pterygoid, which in its turn is united to the quadrate. When the mouth +is shut the quadrate is directed backwards, and carrying back the +pterygoid and transpalatine pulls at the maxillae and causes its +palatal face, to which the poison teeth are attached, to lie back +along the roof of the mouth. When the mouth opens the distal end of +the quadrate is thrust forward, and this necessitates the pushing +forward of the pterygoid and transpalatine, causing the tooth-bearing +surface of the maxillae to look downwards and the tooth to come into +the position for striking. + +The Ophidian skull is also noticeable for the absence of the jugals +and quadratojugals. In poisonous snakes the place of the jugal is +taken by the zygomatic ligament which connects the quadrate and +maxillae. + +The extent to which the palate is closed in reptiles varies much. In +many reptiles, such as the Squamata and Ichthyosauria, the palate is +not complete, both palatines and pterygoids being widely separated in +the middle line. In others, such as the Crocodilia, Sauropterygia, and +Chelonia, there is a more or less complete bony palate. In many +Chelonia this is chiefly formed of the vomer, palatines, and +pterygoids, the posterior nares being mainly bounded by the palatines. +In living Crocodilia, however, outgrowths are formed from the +pterygoids and palatines which arch round and meet one another +ventrally, forming a secondary palate (fig. 43, A), which completely +shuts off the true sphenoidal floor of the skull, and causes the +posterior nares which are bounded by the pterygoids to open very far +back. Though this feature is common to all postsecondary crocodiles, +it is interesting to notice that it is not found in the earlier forms, +but that its gradual evolution can be traced. In the Triassic +_Belodon_, for instance, the posterior nares open far forwards, and +are not surrounded by either the palatines or pterygoids. In the +Jurassic crocodile, _Teleosaurus_, the posterior nares lie further +back, being surrounded by the palatines, but the pterygoids do not +meet them. Finally, in the Tertiary forms the arrangements are as in +living crocodiles. + +A short secondary hard palate is found also in the Theriodontia. The +palatines of _Ichthyosaurus_ are noticeable for their transverse +position, which recalls that in the Frog. + +The various =fossae= or =vacuities= in the false roof of the skull are +important, and their relations may best be understood by a description +of their mode of occurrence in _Sphenodon_, a form in which they are +very completely developed. + +[Illustration FIG. 52. SKULL OF HATTERIA. (_Sphenodon punctatus_). A, +lateral; B, dorsal; C, ventral; D, posterior. (After VON ZITTEL.) + + 1. premaxillae. + 2. nasal. + 3. prefrontal. + 4. frontal. + 5. postfrontal. + 6. parietal. + 7. squamosal. + 8. quadratojugal. + 9. quadrate. + 10. postorbital. + 11. jugal. + 12. maxillae. + 13. vomer. + 14. palatine. + 15. pterygoid. + 16. transpalatine. + 17. exoccipital. + 18. epipterygoid. + 19. basisphenoid. + 20. supratemporal fossa. + 21. infratemporal or lateral temporal + fossa. + 22. orbit. + 23. post-temporal fossa. + 24. foramen magnum. + 25. anterior nares. + 26. interparietal foramen. + 27. dentary. + 28. supra-angular. + 29. articular.] + +In _Sphenodon_, then, on the dorsal surface of the skull, are the +large =supratemporal fossae= (fig. 52, 20). Their inner margins are +separated from one another by the parietal walls of the cranium, while +externally each is bounded by a bony arch, the =supratemporal arcade=, +formed of the postfrontal, postorbital, and squamosal. Posteriorly the +boundary is formed by a =post-temporal bar=, formed by the parietal +and squamosal. Below the supratemporal arcade is another large +vacuity, the =infratemporal= or =lateral temporal fossa= (fig. 52, +21). This is bounded above by the supratemporal arcade, and is +separated from the orbit in front by the =postorbital bar=, formed by +the union of outgrowths from the jugals and postorbitals. Behind it is +bounded by a continuation of the post-temporal bar formed of the +squamosal and quadratojugal, and below by an =infratemporal arcade=, +which is chiefly composed of the quadratojugal and jugal. + +Below the post-temporal bar is a third vacuity, the =post-temporal +fossa= (fig. 52, D, 23), bounded above by the post-temporal bar and +below by the exoccipital and opisthotic. + +_Sphenodon_ and the Crocodilia are the only living reptiles with +complete supratemporal and infratemporal arcades, but they are both +present in the extinct Pterosauria and some Dinosauria. + +Supratemporal fossae, bounded below by supratemporal arcades, occur in +all reptiles except some Chelonia, the Ophidia, the Geckonidae among +Lacertilia, and the Pariasauria and others among Theromorpha; they are +specially large in _Nothosaurus_ among the Sauropterygia, _Dicynodon_ +among the Theromorpha, and many Crocodilia and Pterosauria. In some +Dinosaurs, such as _Ceratosaurus_, they are very small, while the +infratemporal fossae are correspondingly large. + +In _Elginia_[93] (Theromorpha) and some Chelonia, such as _Chelone_, +there are no fossae on the surface of the skull, a complete false roof +being developed; in other Chelonia, such as _Trionyx_, the true +cranium is freely visible, the only part of the false roof developed +being the infratemporal arcade. + +In many reptiles large =pre-orbital vacuities= occur; they are +specially large in the Pterosauria and in some of the Crocodilia and +Dinosauria (fig. 35, 3). In some Pterosauria they are confluent with +the orbits. + +The premaxillae are usually separate, but sometimes, as in some Ophidia +(fig. 51, 1), Chelonia, Lacertilia (Agamidae), and Dinosaurs +(Ceratopsia) they are united. In the Dinosaur _Hadrosaurus_ they are +exceedingly large and spatulate. In the Rhynchocephalian +_Hyperodapedon_ they are drawn out into a strongly curved beak. + +As regards the mandible, sometimes, as in most Rhynchocephalia, +Ophidia and Pythonomorpha, the rami have only a ligamental union; +sometimes, as in Crocodilia, the Rhynchosauridae and the majority of +Lacertilia, they are suturally united. In Chelonia (fig. 28, B, 12), +and apparently in Pterosauria, the two dentaries are completely fused +together. The sutures between the various bones of the lower jaw +usually persist, but in Ophidia those between the angular, +supra-angular, articular and coronoid are obliterated. There are +sometimes large vacuities in the mandible, as in Theromorpha, +Crocodilia, and some Dinosauria. In _Iguanodon_, _Polyonax_, +_Hypsilophodon_ and _Hadrosaurus_ among Dinosaurs the mandible has a +predentary or mento-meckelian bone which, in some cases at any rate, +was probably sheathed in a horny beak. + +The principal part of the auditory ossicular chain is formed by a +rod-like columella. The development of the hyoid apparatus varies, and +it often happens that the first branchial arch is better developed +than is the hyoid arch. In the Crocodilia and Chelonia there is a +large basilingual plate or body of the hyoid (fig. 53, 1); but while +in the Crocodilia the first branchial forms the only well-developed +arch, in the Chelonia the first and second branchials are both +strongly developed, and the hyoid is often fairly large. + + +THE RIBS. + +[Illustration FIG. 53. HYOIDS OF AN ALLIGATOR (_Caiman latirostris_) +(TO THE LEFT) AND OF A GREEN TURTLE (_Chelone midas_) (TO THE RIGHT) × +5/8. (Brit. Mus.) + +The cartilaginous portions are dotted. + + 1. basilingual plate or body of + the hyoid. + 2. hyoid arch. + 3. first branchial arch (anterior + cornu). + 4. second branchial arch (posterior + cornu).] + +Ribs are always present, and may be attached to any of the precaudal +vertebrae. In most reptiles the posterior cervical vertebrae bear +ribs, while the atlas and axis are ribless; in Crocodiles and Geckos, +however, ribs are borne even by the atlas and axis. On the other hand, +in the Chelonia none of the cervical vertebrae bear obvious ribs. In +the following groups the thoracic ribs have both capitula and +tubercula--Theromorpha, Ichthyosauria, Crocodilia, Dinosauria, +Pterosauria. In the other groups each rib articulates by a single +head, and the position of the facet is subject to a considerable +amount of variation, thus in the Squamata it lies on the centrum, and +in the Sauropterygia on the neural arch, while in the Chelonia the rib +articulates with the contiguous parts of two centra instead of +directly with one. + +In most reptiles a greater or smaller number of ribs are united +ventrally with a sternum; but in snakes a continuous series of similar +ribs, all articulating freely with the vertebral column, extends from +the third cervical vertebra to the end of the trunk. The number of +ribs connected with the sternum varies from three or four in Lizards +to eight or nine in Crocodiles. Those which reach the sternum are +nearly always divided into vertebral, sternal, and intermediate +portions, and as a rule only the vertebral portion is completely +ossified. In Crocodiles a number of sternal ribs are connected with a +cartilaginous arch, which is attached to the hind end of the sternum, +and represents the xiphisternum. The sacral ribs connecting the +vertebral column with the ilia are very distinct in Crocodiles; in +these animals and _Sphenodon_ the vertebral ribs have backwardly +projecting uncinate processes as in birds. + +In the curious arboreal lizard, _Draco volans_, the posterior ribs are +long and straight, and support a parachute-like expansion of the +integument used in its long flight-like leaps. In Chelonia the ribs +are generally combined with the carapace. + +In Ichthyosauria, Sauropterygia, Crocodilia and _Sphenodon_, abdominal +splint ribs occur; and probably all except the first of the paired +ossifications forming the plastron of Chelonia are of similar +character. Abdominal ribs have quite a different origin from true +ribs, for while true ribs are cartilage bones, abdominal ribs have no +cartilaginous precursors, but are simply the ossified tendons of the +rectus abdominalis muscle. + + +THE STERNUM. + +A sternum occurs in the following groups of reptiles: Rhynchocephalia, +nearly all Lacertilia, Pythonomorpha, Crocodilia, and Pterosauria, and +is generally more or less rhomboidal or shield-shaped. In Pterosauria +it is keeled and bears some resemblance to that of birds. It may have +been replaced by membrane bone. + +[Illustration FIG. 54. VENTRAL VIEW OF THE SHOULDER-GIRDLE OF STERNUM +OF A LIZARD (_Loemanctus longipes_) × 2. (After PARKER.) + + 1. interclavicle. + 2. clavicle. + 3. scapula. + 4. coracoid. + 5. precoracoidal process. + 6. glenoid cavity. + 7. sternum. + 8. xiphisternum. + 9. sternal rib.] + +The sternum is absent in Sauropterygia, Ichthyosauria, Chelonia, +Ophidia, and most of the snake-like Amphisbaenidae among Lacertilia; +while it is not well known in Theromorpha and Dinosauria. In the +Sauropod _Brontosaurus_, however, two rounded bones occur near the +base of the coracoids, and these probably represent ossified patches +in a sternum, which was mainly cartilaginous; similar structures occur +in _Iguanodon_. + +The sternum frequently remains wholly cartilaginous, especially in +Lacertilia; sometimes it becomes calcified, but true ossification does +not as a rule take place. + + +APPENDICULAR SKELETON. + + +THE PECTORAL GIRDLE. + +The pectoral girdle is well developed in all groups of reptiles except +the Ophidia, occurring even in the limbless Amphisbaenidae. It is very +solid in the Theromorpha. As a rule all three cartilage bones, +scapula, coracoid, and precoracoid are represented, and frequently +also the membrane bones,--clavicles, and interclavicle. + +The coracoids are generally flat expanded bones, which sometimes, as +in Sauropterygia and Ichthyosauria, meet in a ventral symphysis; +sometimes, as in Lacertilia, are united with the sides of the sternum. +In Chelonia neither the coracoids nor precoracoids meet one another, +but their free ends are connected by fibrocartilaginous bands. In +Lacertilia the coracoids are pierced by fenestrae. + +The precoracoid is generally represented, but the Theromorpha are the +only reptiles in which it is separately ossified; it forms a +well-marked process on the coracoid in Lacertilia (fig. 54, 5). It is +absent in Ichthyosauria, and Dinosauria, and probably in +Sauropterygia. In some Lacertilia and Chelonia the sternal ends of the +coracoids are unossified and form epicoracoids; in some Chelonia there +are also epiprecoracoids; but neither these nor the epicoracoids +overlap their fellows of the opposite side as they do in arciferous +Anura (see p. 185). In some Lacertilia with degenerate limbs the +pectoral girdle is also much reduced, in _Ophisaurus apus_ the ventral +borders of the coracoids are widely separated. + +A scapula is always present, and is generally expanded distally, but +in the Chelonia the distal end is cylindrical. In the Theromorpha it +has an acromial process with which the precoracoid articulates, and it +is very large in Dinosauria. In the Chelonia the scapula and +precoracoid are ossified continuously. Among the Pterosauria, +_Pteranodon_ has an unique pectoral girdle; the scapula and coracoid +are ankylosed and the scapula articulates with the neural spines of +several ankylosed vertebrae. + +Clavicles occur in some Theromorpha such as _Pariasaurus_, and also in +the Ichthyosauria, Sauropterygia, Rhynchocephalia, and most +Lacertilia. They are absent in the Pterosauria, the Chamaeleons among +Lacertilia, the Ophidia and the Crocodilia. They are wanting too in +the Chelonia, unless the first pair of ossifications in the plastron +are to be regarded as clavicles. In the Sauropterygia bones regarded +as the clavicles and interclavicle are generally well developed. The +unpaired ossification in the plastron of Chelonia is an interclavicle, +and a representative of the same bone occurs arising from the sternum +in Pterosauria. A well developed =T=-shaped interclavicle is found in +Ichthyosauria, Rhynchocephalia, Lacertilia, and some Theromorpha, such +as _Pariasaurus_. + + +THE LIMBS. + +In most reptiles there are two pairs of pentedactylate limbs provided +with claws, but in nearly all Ophidia and some Lacertilia +(_Amphisbaena_, _Lialis_, _Anguis_) the limbs have entirely +disappeared. In a few Ophidia such as _Python_ traces of the posterior +limbs occur, and in _Chirotes_ among the Amphisbaenidae there are +minute anterior limbs. The Lacertilians, _Chalcides_ (_Seps_) and +_Ophisaurus_ (_Bipes_, _Pseudopus_) have very small posterior limbs. + +The limbs are as a rule adapted for walking, but in Ichthyosauria, +Sauropterygia, Pythonomorpha and some Chelonia, they have the form of +swimming paddles, the relative size of the manus and pes being +increased, while that of the proximal and middle portions of the limbs +is reduced. This reduction is carried to its furthest extent in the +Ichthyosauria in which radius and ulna, tibia and fibula, have the +form of short polygonal bones similar to those constituting the manus +and pes. In the Pythonomorpha the reduction of the limb bones is not +quite so marked, in the Sauropterygia it is less, and still less in +the Chelonia. In the earlier Ichthyosauria too, the limb bones are not +so short as they are in the later forms. The Ichthyosaurian limb is +also remarkable, firstly for the fact that both humerus and femur are +terminated by concave articulating surfaces instead of by convex +condyles, and secondly for the great multiplication of the phalangeal +bones, each digit being sometimes composed of a series of over twenty. +Sometimes too the number of series is increased, either by the +bifurcation of some of the digits or by the development of marginal +bones. In the Sauropterygia the phalanges are likewise increased above +the normal but not so much as in Ichthyosauria. The humerus and femur +of Sauropterygia are noticeable for the enormous size of the terminal +epiphyses which form in each case by far the greater part of the bone. + + +THE ANTERIOR LIMB. + +The anterior limb is usually approximately equal in length to the +posterior, but in many Dinosauria it is considerably the shorter of +the two. The humerus is generally without distinct condyles, but they +are well developed in the Theromorpha, the Lacertilia and _Sphenodon_. + +In the Theromorpha, some Rhynchocephalia, and some Sauropterygia, such +as _Mesosaurus_, the humerus has an ent-epicondylar foramen; in +Lacertilia, Chelonia and some Dinosauria there is an ect-epicondylar +foramen or groove; _Sphenodon_ possesses both ent- and ect-epicondylar +foramina. The radius and ulna are always separate. In some Chelonia, +such as _Chelydra_, the carpus has a very simple arrangement, namely, +a proximal row of three bones, the radiale, intermedium and ulnare, +and a distal row of five carpalia, with one bone, the centrale, +between the two rows. Many reptiles have a carpus only slightly +different from this. Thus the carpus in _Sphenodon_ differs mainly in +having two centralia, that of most Lacertilia, in having the centrale +and intermedium fused. + +Crocodiles have a much reduced carpus with the radiale and ulnare +considerably elongated. The manus in Chamaeleons is curiously +modified, having the first three digits arranged in one group and +turned inwards, and the fourth and fifth in another group turned +outwards; carpalia 3 and 4 are united. + +In the Pterosauria the anterior limbs form wings, the phalanges of the +fifth digit being very greatly elongated to support the wing membrane. +The first digit is vestigial and the second, third, and fourth are +clawed. + + +THE PELVIC GIRDLE. + +The pelvic girdle is well developed in all reptiles which have +posterior limbs, but is absent or quite vestigial in Ophidia and those +Lacertilia which have no posterior limbs. The ilium and ischium agree +in their general characters throughout all the various groups of +reptiles, but that is not the case with the pubis. + +In many reptiles such as Chelonia, Ichthyosauria and Lacertilia the +ilia are small, more or less cylindrical bones either directed +backwards, or vertically placed as in the Chamaeleons. In the +Crocodilia they are larger and more expanded, while in Dinosauria and +Pterosauria they are greatly elongated both in front of, and behind, +the acetabulum. The ischia are generally strongly developed somewhat +square bones meeting in a ventral symphysis. In Dinosauria the ischium +(fig. 35, 9) is a much elongated and backwardly-directed bone, +bearing a forwardly projecting obturator process. In Pterosauria the +ischium is fused with the ilium, and in both pterosaurs and crocodiles +the ilium and ischium are the only bones taking part in the formation +of the acetabulum. In most Lacertilia there is an unpaired structure, +the _hypo-ischium_ or _os cloacae_ projecting back from the symphysis +ischii, which is usually separated from the symphysis pubis by a large +space, the _foramen cordiforme_. In some Lacertilia and Chelonia there +is a cartilaginous bar dividing the foramen cordiforme into two +obturator foramina; in many Chelonia this bar is ossified. Among +_Ophidia_, _Python_, _Tortrix_, _Typhlops_ and their allies have a +structure representing a vestigial ischio-pubis: but in most Ophidia +there is no trace of the pelvis. In some Theromorpha all the bones of +the pelvis are completely fused, forming an os innominatum as in +mammals; the pubes and ischia are so completely fused that sometimes +as in _Pariasaurus_ even the obturator foramina are closed. + +Concerning the reptilian pubis there are considerable difficulties. +Sometimes there is only a single pubic structure present, sometimes +there are two. The reptilian pubis is best understood by comparing the +arrangements met with in the various other groups with that in the +Orthopod Dinosaurs such as _Iguanodon_. In _Iguanodon_ the pubis +consists of two portions, viz. of a moderately broad pre-pubis +directed downwards and forwards, and of a narrow greatly elongated +post-pubis directed backwards parallel to the ischium. The pubis is +united to both ilium and ischium, the acetabulum has a large +unossified space, and neither pre-pubes nor post-pubes meet in ventral +symphyses. The arrangement bears a great resemblance to that of Ratite +birds. In Lacertilia, Chelonia, Rhynchocephalia and Ichthyosauria +together with Theropod and Sauropod Dinosaurs the pubis corresponds to +the pre-pubis of _Iguanodon_ and is a more or less cylindrical bone +expanded at both ends, meeting its fellow in a ventral symphysis. In +Chelonia and Lacertilia the pubis bears a lateral process which is +homologous with the post-pubis of Iguanodon. In Lacertilia and +sometimes in Chelonia there is a cartilaginous epipubis attached to +the anterior border of the pubic symphysis; this is well developed in +the Chamaeleons and Geckos. In Crocodilia there is, forming the +anterior and ventral portion of the acetabulum, a patch of cartilage +(fig. 49, 3) which is probably the true pubis homologous with that of +lizards and with the pre-pubis of _Iguanodon_. The large bone +generally called the pubis in Crocodiles is probably an epipubis. + + +THE POSTERIOR LIMB. + +The posterior limb is entirely absent in some Lacertilia and in most +Ophidia, though traces occur in _Python_, _Tortrix_ and _Typhlops_. In +the Ichthyosauria, Sauropterygia and Pythonomorpha the posterior limbs +form swimming paddles and have been already referred to. + +The arrangement of the proximal and middle segments of the limb is +fairly constant in all reptiles with limbs adapted for walking, and +the tibia and fibula are always separate. The pes is however subject +to a considerable amount of variation, especially as regards the +tarsus. In some Chelonia the tarsus like the carpus has an extremely +simple arrangement, consisting of a proximal row of three bones, the +tibiale, intermedium and fibulare, a centrale, and a distal row of +five tarsalia. In most living reptiles, however, the tibiale and +intermedium are as in mammals united, forming the astragalus. In +Crocodiles (fig. 48, B, 15) the centrale is also united with the +tibiale while the distal tarsalia are very slightly developed. The +calcaneum in Crocodiles is drawn out into a long process forming a +heel in a manner almost unique among Sauropsida. In _Sphenodon_ and +Lacertilia the tibia and fibula articulate with a single large bone +representing the whole proximal row of tarsalia. + +The pes is generally pentedactylate, but in some Crocodiles the fifth +digit is vestigial (fig. 48, B), and in some Dinosauria (fig. 35) +there are only three digits. The North American Dinosaurs present a +continuous series ranging from a pentedactylate plantigrade form like +_Morosaurus_, to such a form as _Hallopus_ with a highly digitigrade +and specialised pes reduced to three functional digits, and a +vestigial fifth metatarsal. The second, third and fourth metatarsals +in this form are nearly two-thirds as long as the femur, and the +calcaneum is drawn out into a heel much as it is in most mammals. + +In Lacertilia, Orthopoda and many Chelonia, the ankle joint comes to +lie between the proximal and distal row of tarsals as in birds. + + +FOOTNOTES: + +[91] See pp. 214 and 215. + +[92] These terms are defined on p. 199. + +[93] E.T. Newton, _Phil. Trans._ vol. CLXXXIV, B, p. 431 (1893). + + + + +CHAPTER XVII. + +CLASS. AVES[94]. + + +Birds form a large and extremely homogeneous class of the vertebrata, +and are readily distinguished from all other animals by the possession +of an epidermal exoskeleton having the form of feathers. Feathers +differ from hairs in the fact that they grow from papillae formed of +both the horny and the Malpighian layer of the epidermis, which +papillae at first project from the surface, and only subsequently +become imbedded in pits of the dermis. A dermal exoskeleton does not +occur in birds. + +The endoskeleton is characterised by its lightness, the large bones +being generally hollow; but the pneumaticity does not vary in +proportion to the power of flight. The cervical part of the vertebral +column is very long and flexible, while the post-cervical portion is +generally very rigid, owing to the fusion of many of the vertebrae, +especially in the lumbar and sacral regions. The vertebrae are +generally without epiphyses to their centra. The cervical vertebrae in +living forms have saddle-shaped articulating surfaces, and many of +them bear ribs. The thoracic ribs in almost all birds have large +uncinate processes. The sternum is very large, and the ribs are always +attached to its sides, not as in many reptiles to anybackwardly +projecting process or processes. The sternum ossifies from two or more +centres. + +The skull is extremely light, and its component bones show a great +tendency to fuse together completely. The facial part of the skull is +prolonged into a beak, chiefly formed of the premaxillae; this beak is +in all modern birds devoid of teeth, and is coated externally with a +horny epidermal sheath. The quadrate is large and freely movable. The +supratemporal arcade[1] is imperfect, while the infratemporal +arcade[95] is complete. There are no postorbital or postfrontal bones. +Neither parotic processes nor an interparietal foramen occur. There +are commonly large pre-orbital vacuities. The palatines and pterygoids +never form a secondary bony palate as in Crocodiles. Part of the floor +of the skull is formed by a wide _basitemporal_ (paired in the embryo) +which is continued in front as a long slender _rostrum_; these +structures have replaced the parasphenoid of Ichthyopsids. Cartilage +or bone is always developed in the sclerotic. The first branchial arch +is well developed, the hyoid arch but slightly. The coracoids are +large, and the clavicles are nearly always united forming the +_furcula_. There is no separate interclavicle and hardly any trace of +a precoracoid. + +The anterior limbs form wings, and the manus is in the adult always +much modified, never having more than three digits. The three bones of +the pelvis are, except in Archaeornithes, always ankylosed together in +the adult, and the ilium is greatly prolonged in front of the +acetabulum, which is perforated. The ilia are not connected with the +sacrum by ossified sacral ribs. The pubes and ischia are directed +backwards parallel to one another, and except in a very few forms +never meet their fellows in ventral symphyses. The fibula is generally +much reduced. The proximal tarsal bones are always ankylosed to the +tibia, and the distal tarsals to the metatarsals, so that the ankle +joint is _intertarsal_. The first metatarsal is nearly always free. +The pes never has more than four digits in the adult. + +The class Aves is most conveniently divided into two subclasses: 1. +Archaeornithes. 2. Neornithes. + + +Subclass I. ARCHAEORNITHES. + +The only form referred to this subclass of extinct birds is +_Archaeopteryx_[96], the earliest known bird. In this animal the +skeleton does not seem to be pneumatic. The cervical and trunk +vertebrae are generally thought to be flat, certainly their +articulating surfaces are not saddle-shaped. There is no long compound +sacrum as in modern birds. The tail is longer than the whole body, the +caudal vertebrae are twenty in number, they gradually taper as traced +away from the trunk, and each bears a pair of feathers. The posterior +caudal vertebrae are not united together to form a _pygostyle_. The +upper jaw bears thirteen pairs of conical teeth, planted in distinct +sockets in the maxillae and premaxillae, but the mandible has only three +pairs. The presence of these teeth forms the most essential difference +between the skull of _Archaeopteryx_ and that of modern birds, and the +fact that they occur on the premaxillae renders it improbable that a +horny beak was present. There is a ring of ossifications in the +sclerotic. The ribs do not show uncinate processes, and articulate +with the vertebrae by single heads not divided into capitula and +tubercula. Abdominal ribs appear to have been present. The furcula is +large, and the scapula has a well developed acromion. The sternum is +unknown. The radius and ulna are approximately equal in size. In the +manus the first, second and third digits[97] are present, each +terminated by a claw. The second digit is considerably the longest, +while the third includes four phalanges. The three bones of the pelvis +probably remained distinct throughout life. The tarsals are ankylosed +respectively to the tibia and metatarsals as in other birds. The +metatarsals are ankylosed together, and the pes has four digits. + + +_Subclass II._ NEORNITHES. + +To this subclass may be referred all known birds except +_Archaeopteryx_. They all agree in having a short tail whose component +vertebrae are commonly ankylosed together forming a pygostyle. The +three metacarpals do not all remain distinct. The bones of the pelvis +are ankylosed together, and to a large though variable number of +vertebrae. There are three orders, the Ratitae, Odontolcae, and +Carinatae. + + +_Order_ 1. RATITAE. + +The Ratitae differ from _Archaeopteryx_ and the great majority of +Carinatae in being flightless. The bones are generally not pneumatic, +containing marrow instead of air, in the Ostrich however they are very +pneumatic. The tail is short and the posterior caudal vertebrae are +generally ankylosed together forming a pygostyle. The pectoral girdle +has comparatively a much smaller size than in Carinatae, clavicles are +small or absent, and the scapula and coracoid lie nearly in the same +straight line. The ilium and ischium do not as in Carinatae unite +posteriorly, and enclose a foramen except in very old Rheas and Emeus. +The quadrate articulates with the cranium by a single head. The vomers +unite and form a broad plate, separating the palatines, pterygoids and +basisphenoidal rostrum. + +The anterior limbs are greatly reduced in size or even absent, while +the posterior limbs are greatly developed and adapted for running. The +tibia and fibula are quite distinct. + +Many ornithologists agree that the various forms grouped together as +Ratitae are not all very closely allied to one another, that they +resemble one another mainly in having lost the power of flight, and do +not form a natural group. + +The Ratitae include the following groups:-- + +_Æpyornithes_[98], huge extinct birds from Madagascar. + +_Apteryges_, including the Apteryx of New Zealand. + +_Dinornithes_[99], the Moas, huge extinct birds from New Zealand, and +some of the neighbouring islands. + +_Megistanes_, including the Cassowaries (_Casuarius_) of Australia, +New Guinea, and some of the neighbouring islands; and the Emeus +(_Dromaeus_) of Australia. + +_Rheornithes_, including the Rheas of S. America. + +_Struthiornithes_, including the Ostriches (_Struthio_) now living in +Africa, and found fossil in N. India and Samos. + + +_Order_ 2. ODONTOLCAE. + +This order includes only an extinct N. American bird +_Hesperornis_[100]. The jaws are provided with a series of sharp teeth +placed in continuous grooves, but the premaxillae are toothless, and +were probably sheathed in a horny beak. The rami of the mandible are +not ankylosed together in front. The skeleton is not pneumatic. The +cervical vertebrae have saddle-shaped articulating surfaces as in +ordinary birds, and the thoracic vertebrae are not ankylosed together. +The tail is comparatively long, and formed of twelve vertebrae with +only slight indications of a pygostyle. The ribs have uncinate +processes. The anterior limb is quite vestigial, being reduced to a +slender humerus. The posterior limb is very powerful and adapted for +swimming. + + +_Order_ 3. CARINATAE. + +This order includes the vast majority of living birds. The cervical +vertebrae have saddle-shaped articulating surfaces (except in the +Ichthyornithiformes). The posterior caudal vertebrae are ankylosed +forming a pygostyle. The quadrate articulates with the cranium by a +double head. In all except the Tinamidae the vomers are narrow behind +and not interposed between the palatines, pterygoids and +basisphenoidal rostrum. The sternum has a median keel, and the +anterior limbs are in the great majority of cases adapted for flight. +Clavicles are well developed, and the scapula and coracoid are nearly +at right angles to one another. The various groups into which the +Carinatae are divisible are shown in the table on pp. 40-42. Their +special characters will not be dealt with. + +[Illustration FIG. 55. _Gallus bankiva_ var. _domesticus_. THE LEFT +HALF OF THE SKELETON. The skull, vertebral column, and sternum are +bisected in the median plane. (After Marshall and Hurst.) + +A, acetabulum. B, cerebral fossa. CB, cerebellar fossa. CL, clavicle. +CO, coracoid. CR, cervical rib. C 1 = one, first cervical vertebra. +FE, femur. HC, ventral end of clavicle. HU, humerus. HY, hyoid. IF, +ilio-sciatic foramen. IL, ilium. IS, ischium. L, lachrymal. MC 3, +postaxial metacarpal. MN, mandible. MS, xiphoid processes. MT, +tarso-metatarsus. MT 1, first metatarsal. N, nasal. OP, optic foramen. +P, premaxillae. PB, pubis. PL, palatine. PY, pygostyle. R, radius. RC, +radial carpal. S, keel of sternum. SC, scapula. T, tibio-tarsus. TH 4, +fourth thoracic vertebra. U, ulna. UC, ulnar carpal. UP, uncinate +process. Z, infra-orbital bar. 1, 2, 3, 4, first, second, third and +fourth digits of pes. 3, pre-axial, 4, middle, and 5, postaxial digit +of manus.] + + +FOOTNOTES: + +[94] M. Fürbringer, _Untersuchungen zur Morphologie und Systematik der +Vögel_, I. and II. Amsterdam, 1888. Cf. H. Gadow, _Nature_, XXXIX. +1888, pp. 150 and 177. + +T.H. Huxley, "On the classification of birds." _P.Z.S._, London, 1867. + +E. Selenka and H. Gadow, _Vögel_ in Bronn's _Classen und Ordnungen des +Thierreichs_ 1869-1890. + +[95] See p. 283. + +[96] R. Owen, _Phil. Trans._, vol. CLIII., p. 33; 1863. T.H. Huxley, +_P. R.S._, vol. XVI., p. 243; 1868. C. Vogt, _Rev. Scient._, ser. 2, +tom. 9, p. 241; 1879. C.H. Hurst, _Nat. Sci._, vol. III., p. 275; +1893; vol. VI., pp. 112, 180, 244; 1895. W.P. Pycraft, _Nat. Sci._, +vol. V., pp. 350 and 437; 1894; and vol. VIII., p. 261; 1896. + +[97] According to Hurst the fourth and fifth digits are also present. + +[98] See C.W. Andrews, _P.Z.S._, 1894, p. 108. + +[99] See T.J. Parker, _Tr. Zool. Soc. London_, vol. XIII., pt. 2, +1895, and F.W. Hutton, several papers in _Tr. N. Zealand Inst._, 1893 +and 1895. + +[100] See O.C. Marsh. _Odontornithes. A monograph of the extinct +toothed birds of N. America._ New Haven, 1880. + + + + +CHAPTER XVIII. + +THE SKELETON OF THE WILD DUCK (_Anas boschas_). + + +I. EXOSKELETON. + +The exoskeleton of the Duck and indeed of all birds is entirely +epidermal in origin. Its most important part consists of =feathers=, +but it includes also the following horny structures:-- + +(_a_) =scales=, which cover the toes and tarso-metatarsus; + +(_b_) =claws=, which are attached to the distal phalanges of the toes +and of the pollex; + +(_c_) the wide =beak=, which sheaths both upper and lower jaws, and +whose edges are raised into lamellae, which act as strainers. + + +FEATHERS. + +A well developed feather, such as one of the large quill feathers of +the wing or tail, consists of the following parts: A main stem, the +=scapus=, which forms the axis running along the whole length of the +feather, and is divided into (1) a proximal hollow cylindrical +portion, the =calamus= or =quill=, and (2) a distal solid portion, the +=rachis= or =shaft=, which is square in section, flexible and grooved +along its ventral surface, and bears a number of lateral processes, +the =barbs=. The =calamus= which is partly imbedded in a pit in the +dermis, bears two holes: one, the =inferior umbilicus=, is at its +proximal end, and into it enters a vascular outgrowth from the dermis; +the other, the =superior umbilicus=, lies on the ventral surface at +the junction of the calamus and scapus. + +The =barbs= are a series of narrow elastic plates, attached by their +bases to the rachis, and with their edges looking upwards and +downwards. The barbs are connected together by a number of smaller +processes, the =barbules=, which interlock with one another by means +of hooklets, and bear the same relation to the barbs that the barbs do +to the rachis. The barbs and barbules, together with the rachis, +constitute the =vexillum= or =vane= of the feather. Any feather having +the above type of structure is called a =penna= or a =contour +feather=, from the fact that it helps to produce the contour of the +body. + + +VARIETIES OF FEATHERS. + +1. =Pennae.= There are two kinds of pennae or contour feathers. + +(_a_) The =quills=. These form the large feathers of the wing and +tail. They are divided into two groups, the =remiges=, or wing quills, +and the =rectrices=, or tail quills. + +The =remiges=[101] include three sets of feathers, the =primaries= or +=metacarpo-digitals=, which are attached to the bones of the manus, +the =secondaries= or =cubitals=, which are attached to the ulna, and +the =humerals=, which are attached to the humerus. + +The =primaries= differ from all the other quill feathers in having the +posterior half of the vane much wider than the anterior half. They are +ten in number, and of these six, the =metacarpal= quills (fig. 57, +14), are attached to the second and third metacarpals, one, the +=ad-digital= (fig. 57, 15), to the phalanx of the third digit, two, +the =mid-digitals= (fig. 57, 16), to the first phalanx of the second +digit, and two, the =pre-digitals= (fig. 51, 17), to the second +phalanx of the second digit. One of the pre-digitals is very small, +and is called the =remicle= (fig. 57, 11). + +[Illustration FIG. 56. THE WING OF A WILD DUCK (_Anas boschas_). + +The upper figure shows the dorsal side of a right wing, the lower +figure the ventral side of a left wing. × 1/3. (Brit. Mus.) + + 1. scapulars. + 2. tectrices marginales. + 3. tectrices minores. + 4. bastard wing. + 5. tectrices majores. + 6. metacarpo-digitals or primaries. + 7. tectrices mediae. + 8. cubitals or secondaries. + 9. pennae humerales. + 10. pennae axillares.] + +In addition, a group of three quill feathers is attached to the first +digit, constituting the =bastard wing= or =ala spuria= (fig. 56, 4). + +The =secondaries= or =cubitals= (fig. 56, 8) form a group of seventeen +feathers, attached to the ulna; they are shorter than the primaries, +and do not have the posterior half of the vane much wider than the +anterior half. + +The =humerals= (figs. 56, 9 and 57, 12) form a group of eight small +feathers, of varying length, attached to the anterior half of the +humerus. + +[Illustration FIG. 57. WINGS OF A WILD DUCK WITH THE COVERTS REMOVED +(_Anas boschas_). × 1/3. + +A. Right wing seen from the dorsal side. B. Left wing disarticulated +and seen from the ventral side. (Brit. Mus.) + + 1. humerus. + 2. radius. + 3. ulna. + 4. radial carpal. + 5. ulnar carpal. + 6. first phalanx of first digit. + 7. second metacarpal. + 8. third metacarpal. + 9. first phalanx of second digit. + 10. second phalanx of second digit. + 11. remicle. + 12. pennae humerales. + 13. cubitals or secondaries. + 14. metacarpal quills. + 15. ad-digital. + 16. mid-digitals. + 17. pre-digital.] + +(b) The =tectrices= or =coverts= are short feathers, which cover over +the quills of the rectrices and remiges, and clothe the body +generally. Their barbules are less developed than is the case with the +quill feathers, so that the barbs separate readily from one another, +especially at the base of the vane. The nomenclature of the various +patches of coverts on the wings is seen in fig. 56. A small patch of +backwardly-directed feathers surrounding the external auditory opening +are known as the =auriculars=. + +2. The =filoplumes= are rudimentary feathers, consisting of a minute +stem and slightly developed vane. They are left in the skin after the +other feathers have been removed. + +3. The =plumulae=, or down feathers, have the stem very slightly +developed, while the barbs are soft and free from one another. They +are distributed all over the body, not only among the contour +feathers, but also over the spaces (_apteria_) which bear no contour +feathers. + +In the young bird the rudiments of the new feathers are formed at the +bases of the embryonic down feathers, and as they grow they push them +out from the skin. The embryonic down feathers however remain attached +to the apices of the new feathers till these have reached a length of +about an inch; they are then shed. + + +II. ENDOSKELETON. + +As compared with that of the Turtle or Crocodile, the endoskeleton of +the Duck is characterised by: + +1. The great lightness of the bones, many of which contain air +cavities. + +2. The tendency to become ankylosed together shown by many of the +bones. + +3. The modification of the anterior limbs and girdle for the purpose +of flight. + + +1. THE AXIAL SKELETON. + +This, as in other vertebrates, is divisible into-- + + A. The vertebral column. + + B. The skull. + + C. The ribs and sternum. + + +A. THE VERTEBRAL COLUMN. + +The vertebral column of the duck, like that of the great majority of +birds, presents a number of well-marked characteristics, contrasting +strongly with those of the generality of higher vertebrates. The centra +are always without epiphyses. The neck is exceedingly long, about as +long as all the rest of the vertebral column put together, and is +remarkable for its flexibility. The trunk portion of the vertebral +column on the other hand is characterised by extreme rigidity, and +the marked tendency shown by the component vertebrae to fuse together +into one almost continuous mass. The most rigid part of the vertebral +column is that to which the pelvis is united, as no less than seventeen +vertebrae take part in the union. The tail of the duck, like that of +all living birds, is very short, and the posterior caudal vertebrae are +united together, forming the =pygostyle=. The vertebral column may be +divided into cervical, thoracic, lumbar, sacral, and caudal regions, +but the boundaries between the several regions are ill-defined. + + +THE CERVICAL VERTEBRAE. + +All the vertebrae anterior to the first one that bears a rib meeting +the sternum are regarded as cervical vertebrae. There are therefore +sixteen cervical vertebrae, the last two of which bear well developed +ribs. All are freely movable on one another. + +As a typical cervical vertebrae, any one from the fifth to the ninth +may be taken. The vertebra is rather elongated, and is very lightly +and strongly made, its most characteristic feature being the shape of +the articulating surfaces of the centra, which are generally described +as saddle-shaped. The anterior surface is convex from above downward, +and concave from side to side, while the posterior and more prominent +surface is concave from above downwards and convex from side to side. +The neural arch is low, and is drawn out into a slight blade-like +=neural spine=. Its base is deeply notched on both sides posteriorly +for the exit of the spinal nerves. Above these notches it is drawn out +into two rather prominent diverging processes, which bear the +=postzygapophyses=,--two flattened surfaces which look downwards and +outwards. The =transverse processes= form irregular outgrowths from +the anterior two-thirds of the sides of the vertebra; each projects +for a short distance downwards and outwards, and is terminated +posteriorly by a short backwardly-projecting spine. The transverse +processes are shown by development to ossify from separate centres, +and are therefore to be regarded as cervical ribs, and each is +perforated at its base by a canal for the passage of the vertebral +artery. Above the anterior end of the vertebrarterial canal are a pair +of thickened outgrowths, which bear upwardly and inwardly directed +=prezygapophyses=. Each transverse process is perforated near its +middle by a prominent foramen through which passes a vein which is +connected with the jugular vein. + +The third and fourth cervical vertebrae resemble the succeeding ones +in most respects, but have small =hypapophyses=, and the neural spines +are less blade-like. The posterior cervical vertebrae (tenth to +sixteenth) differ somewhat from the middle ones. They are shorter and +more massive, the neural arch is much shorter, being deeply notched in +the middle line in front and behind. The transverse processes arise +from the anterior half of the vertebra only, and in the eleventh +vertebra each is drawn out below into a pair of rather prominent +downwardly and inwardly directed processes. In the twelfth vertebra +these processes have almost coalesced, and in the thirteenth vertebra +they have coalesced completely, forming a prominent =hypapophysis=. In +the succeeding vertebrae this hypapophysis rapidly decreases in size. + +The fifteenth and sixteenth cervical vertebrae resemble the succeeding +thoracic vertebrae, having short thick centra and prominent squarely +truncated neural spines; the sides of the neural arches are very +deeply notched. The fifteenth vertebra has a short transverse process, +perforated by a wide vertebrarterial foramen, but this foramen is +absent in the sixteenth. The transverse processes of the fifteenth +vertebra bear two facets for the articulation of the capitulum and +tuberculum of the rib. The sixteenth vertebra has its tubercular facet +on the transverse process, but the capitular facet is borne on the +centrum. + +The second or =axis= vertebra is small, and has the centrum drawn out +into a comparatively very large hypapophysis. The posterior +articulating surface of the centrum is saddle-shaped, the anterior +nearly flat: above it the centrum is prolonged into the prominent +=odontoid process=, which is shown by development to be the detached +centrum of the atlas. The neural arch is deeply notched in the middle +line in front, and at the sides behind. It is drawn out posteriorly +into a wide massive outgrowth, which overhangs the third vertebra and +bears the downwardly-directed postzygapophyses. The prezygapophyses +are situated at the sides of the anterior end of the neural arch, and +look directly outwards. The transverse processes are very slightly +developed, and are pierced by the vertebrarterial canals. + +The =atlas= vertebra is a very slight ring-like structure, thickened +ventrally and bearing in front a prominent concave cavity for +articulation with the occipital condyle of the skull. Posteriorly it +bears a more or less flattened surface for articulation with the +centrum of the axis. It surrounds a large cavity partially divided +into a larger dorsal portion, which is the neural canal, and a smaller +ventral portion which lodges the odontoid process. The sides of the +atlas are pierced by the vertebrarterial canals, above which there are +two slight backwardly-projecting outgrowths bearing the +postzygapophyses on their inner faces. + + +THE THORACIC VERTEBRAE. + +The thoracic region includes all the vertebrae bearing free ribs, +except the first two, viz. those whose ribs do not reach the sternum. +There are seven thoracic vertebrae. The first four have centra with +saddle-shaped articulating surfaces, but are more or less firmly +united together by their neural spines; the last two are completely +ankylosed by their centra to the lumbar vertebrae. + +Each of the first five vertebrae has a prominent, vertical, abruptly +terminated neural spine, and straight transverse processes. The +zygapophyses and articulating surfaces at the ends of the centra are +well developed. The third, fourth, fifth, and sixth vertebrae have +very prominent hypapophyses. The articular facets for the ribs are +well marked, those for the tubercula lying at the free ends of the +transverse processes, and those for the capitula at the sides of the +anterior ends of the centra. The sixth and seventh thoracic vertebrae +are firmly fused by their centra and neural arches to one another and +to the lumbar vertebrae behind, and by their transverse processes to +the ilia. The sixth has its centrum terminated in front by a +saddle-shaped articulating surface, and bears a pair of prominent +prezygapophyses. Its transverse processes and centrum bear facets for +the tubercula and capitula of the ribs respectively. In the seventh +vertebra the tubercular facet is wanting. + + +THE SACRUM. + +[Illustration FIG. 58. A, DORSAL AND B, VENTRAL VIEW OF THE PELVIS AND +SACRUM OF A DUCK (_Anas boschas_). + + 1. ilium. + 2. ischium. + 3. pubis. + 4. pectineal process. + 5. lumbar vertebrae. + 6. true sacral vertebrae.] + +The =sacrum= generally consists of seventeen vertebrae fused with one +another and with the ilia. Their number may be reckoned from the +number of foramina for the exit of spinal nerves. The two most +anterior of these vertebrae bear ribs and have been already described +with the other thoracic vertebrae. Their neural spines and those of +the four succeeding vertebrae are fused together, forming a continuous +crest of bone completely united laterally with the ilia. The +transverse processes of all these six vertebrae are well developed, +but those of the posterior two (fig. 58, B, 5) are much the stoutest. +The next three vertebrae have broad centra, but their transverse +processes are very slightly developed and have no ventral elements. +These seven vertebrae belong to the =lumbar= series. The remaining +eight vertebrae have well-developed transverse processes, which in the +case of the first three or four are divisible into dorsal and ventral +elements. All the dorsal elements are united to form a pair of +flattened plates, partially separated by a series of foramina from a +median plate formed by the united neural arches. Laterally they are +continuous with the ischia. The first two of this series of vertebrae +are shown by their relation to the nerves to be the true =sacrals= +(fig. 58, B, 6), the remaining six belonging to the =caudal= series. + +Behind them come the six free caudal vertebrae, succeeded by a +terminal piece, the =pygostyle=, formed of a number of vertebrae fused +together; this bears the rectrices or tail quills. + +[Illustration FIG. 59. SKULL OF A DUCK (_Anas boschas_). × 1. + +A. Dorsal view of the cranium. B. Palatal view of the mandible. C. The +Hyoid. + +For numbers see Fig. 60.] + + +B. THE SKULL. + +[Illustration FIG. 60. A. Ventral view of the cranium of a Duck (_Anas +boschas_). B. Cranium and mandible seen from the left side. × 1. + + 1. maxillae. + 2. premaxillae. + 3. anterior nares. + 4. nasal process of premaxillae (fig. 59). + 5. nasal. + 6. frontal (fig. 59). + 7. lachrymal. + 8. postfrontal process. + 9. parietal (fig. 59). + 10. jugal. + 11. quadratojugal. + 12. quadrate. + 13. condyle of mandible. + 14. posterior articular process. + 15. dentary at symphysis. } + 16. basi-hyal. } (fig. + 17. uro-hyal. } 59). + 18. basibranchial. } + 19. vomer. + 20. palatine. + 21. pterygoid. + 22. anterior palatine foramen. + 23. basitemporal. + 24. foramen leading into tympanic cavity. + 25. bristle inserted into posterior opening of carotid canal. + 26. bristle inserted into posterior opening of Eustachian canal. + 27. bristle emerging through anterior opening of carotid canal. + Close by is seen the bristle emerging through the anterior + opening of the Eustachian canal. + 28. fenestral recess. + 29. maxillo-palatine. + 30. lambdoidal crest. + 31. rostrum. + +I. II. IV. V. IX. X. nerve foramina.] + +The skull of the duck, like that of birds in general, is characterised +(1) by its lightness, (2) by the contrast between the bones of the +cranium proper and those forming the rest of the skull, for the bones +forming the cranium proper are closely fused together, the sutures +between them being nearly all completely obliterated in the adult, +while the bones forming the face are loosely connected with the +cranium proper; (3) by the prolongation of the face into a long +toothless beak; (4) by the size of the orbits, and their position +entirely in front of the cranium, so that they are separated from one +another only by a thin interorbital septum. + +For purposes of description the skull may be divided into + + (1) The cranial portion. + (2) The facial portion. + (3) The mandible. + (4) The hyoid. + + +(1) THE CRANIAL PORTION. + +This is a rounded box expanded dorsally and posteriorly, but tapering +antero-ventrally. In the young skull the divisional lines between the +several bones can be easily seen, but in the adult they are quite +obliterated. + +(_a_) The _dorsal surface_ is rounded, expanded in front and behind, +but encroached upon in the middle by the cavities of the orbits. There +is a prominent divisional line in front, separating it from the facial +part of the skull. It is formed mainly by the _frontal_ (fig. 59, A, +6) and _parietal_ bones, but the frontals diverge a little anteriorly +and enclose between them the ends of the _nasal processes_ (fig. 59, +A, 4) of the _premaxillae_. Just in front of the orbit the outer +margins of the frontals are either notched or pierced by a pair of +foramina. + +(_b_) At the _posterior end_ of the cranium the most prominent feature +is the large, almost circular =foramen magnum=, through which the +spinal cord and brain communicate; this in young birds is seen to be +bounded by four distinct bones, dorsally by the =supra-occipital=, +ventrally by the =basi-occipital=, and laterally by the +=exoccipitals=. + +The =basi-occipital= forms the main part of a prominent convex knob, +the =occipital condyle=, with which the atlas articulates. The +occipital condyle is slightly notched above, and the ventral surface +of the cranium is deeply pitted just in front of it; the exoccipitals +also contribute slightly to its formation. Slightly in front of and +ventral to the foramen magnum is a small foramen through which the +hypoglossal nerve leaves the cranial cavity. + +The =supra-occipital= is separated from the parietal by a suture line +along which run a pair of prominent ridges, the =lambdoidal crests= +(fig. 60, B, 30). There are often a pair of prominent vacuities in the +supra-occipital dorsal to the foramen magnum. The =epi-otics= and +=opisthotics= become completely fused with the bones of the occipital +segment at a very early stage. + +(_c_) The _ventral surface_ of the cranium is wide behind, where it is +formed by a broad transverse membrane bone, the _basitemporal_ (fig. +60, A, 23), the sides of which are fused with the auditory capsules. +Slightly in front of and an eighth of an inch external to the +hypoglossal foramen the cranial wall is pierced by a pair of foramina +through which the tenth or pneumogastric nerves leave (fig. 60, A, X). +At the sides of the basitemporal are a pair of depressions, the +=tympanic recesses=, in each of which are three holes. Straight lines +joining these holes would form an isosceles triangle with its apex +directed forwards. Of the two holes at the base of the triangle, the +one nearer the middle line and leading into the cranial cavity, is for +the exit of the ninth or glossopharyngeal nerve (fig. 60, A, IX), it +lies just in front of the pneumogastric foramen. The more external +leads into the tympanic cavity, while the more anterior at the apex of +the triangle is the =posterior opening of the carotid canal= (fig. 60, +A, 25), which traverses the base of the cranium, and during life +lodges the carotid artery. + +The anterior end of the basitemporal is pierced near the middle line +by a pair of holes, the =anterior openings of the Eustachian canals=; +while just in front of these and a little further removed from the +middle line are the anterior openings of the =carotid canals=. +Bristles passed in through the posterior openings of the carotid +canals will emerge here (fig. 60, A, 27). In front of the basitemporal +the base of the cranium is formed by the =rostrum= (fig. 60, A, 31), +or thickened basal portion of the interorbital septum; this bears two +prominent surfaces with which the pterygoids articulate. In some kinds +of duck these surfaces are borne by well-marked basi-pterygoid +processes. + +(_d_) _The side of the cranium._ At the base of the posterior end is +seen the deep =tympanic cavity=. The dorsal part of this is divided by +a vertical partition into two halves; of these the more anterior is +the larger, and forms a deep funnel-shaped cavity, the =posterior +opening of the Eustachian canal= (fig. 60, B, 26). A bristle passed +into this opening emerges through the anterior opening of the +Eustachian canal. The more posterior of the two is the =fenestral +recess= (fig. 60, B, 28), and is in its turn divided by a slender +horizontal bar into a dorsal hole, the =fenestra ovalis=, and a +ventral hole, the =fenestra rotunda=. During life the fenestra ovalis +lodges the proximal end of the =columellar= chain. Lying at the outer +side and slightly dorsal to the tympanic cavity is a deep depression, +the =lateral tympanic recess=, and immediately in front of this is the +articular surface for the quadrate. The tympanic cavity is bounded +below by the basitemporal, posteriorly by the exoccipital, and above +by the _squamosal_, a membrane bone, which roofs over a good deal of +the side of the cranium, and bears ventrally a prominent surface with +which the quadrate articulates. Just in front of this is a large round +hole, the =trigeminal foramen= (fig. 60, B, V), behind which the +squamosal is drawn out into a short process. + +In front of the squamosal there is a prominent forwardly-projecting +=postfrontal process= (fig. 60, 8), which ossifies from a different +centre from that forming the squamosal, but in the adult is completely +fused with it. + +The =orbit= forms a large more or less hemispherical cavity which +lodges the eyeball. It is separated from its fellow of the opposite +side by an imperfect partition, the =interorbital septum=. In the +young skull it is seen to be bounded above by the frontal, with which +the _lachrymal_ (fig. 60, 7) is fused anteriorly, forming a large +backwardly-projecting process; while behind it is bounded by +the =alisphenoid=. The interorbital septum is formed by the +ossification and coalescence of the =mesethmoid= in front, with the +=orbitosphenoid= behind, and the =rostrum= below. The boundary of the +orbit below is very imperfect, the zygomatic arch being incomplete. + +The interorbital septum is pierced by the very prominent =optic +foramen= (fig. 60, B, 2), just behind which are the two much smaller +foramina for the exit of the oculomotor and pathetic (fig. 60, B, IV) +nerves, the more anterior being that for the oculomotor. + +Above and slightly in front of the optic foramen is a median opening, +the =olfactory foramen.= This leads into the cranial cavity behind, +and in front is continued forwards as a groove between the +interorbital septum and the frontal. + + +(2) THE FACIAL PART OF THE SKULL. + +This includes the olfactory capsule and associated bones, and the +upper jaw. + +The bones associated with the olfactory capsules are the _nasals_ and +_vomer_. The _nasals_ (figs. 59 and 60, 5) lie on the dorsal surface +immediately in front of the cranium, and are separated from one +another by the nasal processes of the premaxillae. Each is completely +fused in the adult with the corresponding maxillae and premaxillae, the +three bones together forming the boundary of the =anterior nares.= The +_vomer_ (fig. 60, 19) is unpaired and forms a small median vertical +plate lying ventral to the anterior continuation of the interorbital +septum. + +The bones of the upper jaw consist on each side of two slender arcades +which in front converge and are attached to the large beak, while +behind they diverge but are united by the =quadrate=. + +The =inner arcade= is formed by the pterygoid and palatine. The +_pterygoid_ (fig. 60, 21) is a short flattened bone, which articulates +behind with the quadrate, and on its inner side with a large flattened +surface borne by the rostrum, in front it meets the palatine, or +sometimes ends freely with a long antero-dorsally directed point. + +The _palatine_ (fig. 60, 20) is a slender irregular bone flattened +dorso-ventrally at its anterior end where it articulates with the +beak, and laterally behind. It gives off at its posterior end a +process, which is sometimes united with the vomer, sometimes projects +forwards, and meets its fellow dorsal to the vomer. In the large space +between it and the vomer is the opening of the =posterior nares=. + +The _premaxillae_ (figs. 59 and 60, 2) are very large, and form nearly +a third of the big shovel-shaped beak. They constitute the inner, and +part of the front boundary of the anterior nares, and send back a pair +of _nasal processes_ which partially separate the nasals from one +another. + +The =outer arcade= forms the slender =suborbital bar=, and consists +mainly of two rod-like bones, which in the adult are completely fused +together. The posterior of these is the _quadratojugal_ (figs. 59 and +60, 11) which articulates with the quadrate, the anterior is the small +and slender _jugal_ or _malar_ (figs. 59 and 60, 10). The extreme +anterior part of the bar is formed by the _maxillae_. The main part of +the maxillae however lies anterior to the suborbital bar, and extends +forwards along the side of the premaxillae forming all the lateral part +of the beak (figs. 59 and 60, 1); it also sends inwards a plate, the +=maxillo-palatine= (fig. 60, A, 29), which completely fuses with its +fellow in the middle line, and forms the posterior boundary of the +anterior palatine foramen. The term =desmognathous= describes the +condition of the skull in which the maxillo-palatines fuse with one +another in the middle line in this way. + +The =quadrate= (fig. 60, 12), which unites the two arcades behind, is +a stout irregular four-cornered bone forming the =suspensorium=. It +articulates by its dorso-posterior corner with the squamosal, and by +its antero-internal corner with the pterygoid. The middle of its +ventral surface forms a hemispherical knob with which the mandible +articulates, while its dorso-anterior border is drawn out into a long +point which extends towards the interorbital septum. + + +(3) THE MANDIBLE. + +The =mandible= or lower jaw consists of two =rami= which are flattened +and fused together in the middle line in front, while behind they +diverge from one another and articulate with the quadrates. + +Each ramus is composed of five bones fused together, one being a +cartilage bone, and the other four membrane bones. The =articular= is +the only cartilage bone of the mandible, it bears the double condyle +(figs. 59 and 60, 13) or concave articular surface for the quadrate, +and is drawn out behind into a large hooked =posterior articular +process=. The articular is also drawn out into a prominent process on +each side of the articular surface for the quadrate, and is marked by +a deep pit opening posteriorly. The articular is continuous in front +with =Meckel's cartilage= which forms the original cartilaginous bar +of the lower jaw, and is ensheathed by the membrane bones. Of these +the _supra-angular_ forms the upper part of the mandible in front of +the articular, its dorsal surface is drawn out into a small =coronoid +process=, its outer surface also bearing a prominent process. The +_angular_ is a small bone which underlies the articular and +supra-angular on the inner side of the jaw. The _dentary_ (fig. 59, +15) forms the anterior half of each ramus, and is the largest bone of +the mandible; it is fused with its fellow at the symphysis in front, +and extends back below the supra-angular. The _splenial_ is a small +bone lying along the middle half of the inner side of each ramus of +the mandible. + + +(4) THE HYOID. + +With the hyoid apparatus is included the =columella=. This forms a +minute rod of bone, one end of which is expanded and fits into the +fenestra ovalis, while the other end, terminated by a triradiate piece +of cartilage, is attached to the tympanic membrane. The structure is +as a whole homologous with the auditory ossicles of mammals and the +hyomandibular of fish. + +The =hyoid= consists of a median unpaired portion, formed of two +pieces of bone, the =basi-hyal= (fig. 59, C, 16) in front, and the +=uro-hyal= (fig. 59, C, 17) behind, the two being placed end to end +and terminated anteriorly by an unpaired cartilaginous plate, the =os +entoglossum.= At the posterior end there come off a pair of long +=posterior cornua=, each of which consists of two pieces, a longer +=basibranchial= (fig. 59, C, 18), and a shorter =cerato-branchial=. +For the homology of these parts see p. 336. + + +THE RIBS AND STERNUM. + +The last two cervical vertebrae bear long movable ribs which +articulate by distinct capitular and tubercular processes, but do not +meet the sternum. The thoracic ribs are eight in number, and each is +divisible into a =vertebral= and a =sternal= portion. The first five +thoracic ribs are flattened curved bars of bone, which articulate by a +prominent =capitulum= with the centrum of the corresponding vertebra, +and by a =tuberculum= with the transverse process. Projecting +backwards from each is a large hooked =uncinate process.= The last +three ribs which are without uncinate processes, become progressively +more slender, and in the eighth the tubercular processes are lost. + +The sternal portions of the ribs are imperfectly ossified pieces, +short and comparatively thick in the case of the anterior ribs, longer +and more slender in the case of the posterior ribs. + + +THE STERNUM[1]. + +The =sternum= or breast bone is exceedingly large in the Duck, as in +all birds, and projects back far beyond the thorax over much of the +anterior part of the abdomen. It is an irregularly oblong plate of +bone, abruptly truncated behind, somewhat concave dorsally, and drawn +out ventrally into a prominent keel, the =carina=, which projects for +some distance forwards beyond the body of the sternum, and tapers off +gradually behind. The point where the carina joins the body of the +sternum is at the anterior end drawn out into a small process, the +=rostrum=[102]. Just dorsolateral to this are a pair of deep grooves, +the =coracoid grooves=, with which the coracoids articulate. + +The sides of the sternum are drawn out in front into a pair of short +blunt =costal processes;= and just behind these are a series of seven +surfaces with which the ends of the sternal ribs articulate. +Immediately behind these surfaces the sides are produced into a pair +of long backwardly-projecting =xiphoid processes= which nearly meet +processes from the posterior end of the sternum. + + +2. THE APPENDICULAR SKELETON. + +This consists of the skeleton of the anterior and posterior limbs and +of their respective girdles. + + +A. THE PECTORAL GIRDLE[103]. + +The pectoral girdle in almost all birds is strongly constructed and +firmly united to the sternum. It consists of three bones, a dorsal +element, the =scapula=, a posterior ventral element, the =coracoid=, +and an anterior ventral element, the _clavicle_. + +The =scapula= forms a long curved flattened bone expanded at its +anterior end, where it meets the coracoid, and lying across the ribs +at its tapering posterior end. It helps to form the imperfect =glenoid +cavity=, with which the humerus articulates. The =coracoid=, a shorter +but stouter bone than the scapula, has its upper end or =head= +thickened and bears on its posterior border an irregular surface, with +part of which the scapula articulates, while the rest forms part of +the glenoid cavity. The inner border of the coracoid adjoining the +articular facet for the scapula is produced into a strong process +which helps to complete the =foramen triosseum=, a space lying between +the adjoining ends of the scapula and coracoid, through which the +tendon of the second pectoral muscle passes. The lower part of the +coracoid, which is much flattened and expanded, and abruptly truncated +posteriorly, articulates with the coracoid groove of the sternum. The +_clavicle_ is a thickened curved membrane bone, which is fused with +its fellow in the middle line below, the two forming the _furcula_ or +merrythought. Its dorsal end is drawn out into a process which +articulates with the coracoid. + + +THE ANTERIOR LIMB OR WING. + +This consists of three parts, a proximal part, the upper arm or +=brachium=, a middle part, the fore-arm or =antibrachium=, and a +distal part, the =manus=. When extended for flight the parts lie +almost in the same straight line, but when at rest they are folded on +one another in the form of a Z, the brachium and manus pointing +backwards, and the antibrachium forwards. When extended for flight the +surfaces and borders of the wing correspond in position with those of +the primitive vertebrate limb[104], the pre-axial border being +directed forwards and the postaxial backwards, while the dorsal and +ventral surfaces look respectively upwards and downwards. But when the +wing is at rest, the humerus as it extends backwards becomes slightly +rotated, so that its dorsal surface looks more inwards than upwards, +while the dorsal surface of the antibrachium looks partially outwards +and upwards, and that of the manus mainly outwards. + +The =brachium= or =upper arm= contains only a single bone, the +=humerus= (fig. 57, 1). This is a large nearly straight bone expanded +at both ends. The proximal end is specially expanded, forming two +=tuberosities=, and a large convex =head= articulating with the +glenoid cavity. The =pre-axial tuberosity= is the smaller of the two, +but is continued by a prominent =deltoid ridge=, which extends for a +very short distance down the shaft. The =postaxial tuberosity= is the +larger, and below it there is a very deep pit, the =pneumatic +foramen=, which leads into an air cavity in the shaft of the bone. The +shaft is long and straight, and at the distal end of the bone is the +=trochlea= with two convex surfaces, one pre-axial with which the +radius articulates, the other postaxial for the ulna. + +The =fore-arm= or =antibrachium= consists of two bones, the =radius= +and =ulna=. These are of nearly equal length, and are separated from +one another by a considerable space except at their terminations. + +The =radius= (fig. 57, 2), the pre-axial and smaller bone, is straight +and fairly stout; its proximal end articulates with the humerus by a +slightly cupped surface, while its distal end, which articulates with +the carpus, is convex and somewhat expanded. + +The =ulna= (fig. 57, 3) is longer, stouter, and slightly curved. Its +proximal end is expanded, forming two surfaces which articulate with +the trochlea of the humerus; behind them it is drawn out into a short +blunt =olecranon process=. Its distal end is less expanded, and +articulates with the carpus and also with the radius. + +The =Manus=. This includes the carpus or wrist, and the hand. + +The =Carpus=. While in the embryo the carpus consists of five +distinct elements arranged in a proximal row of two and a distal row +of three, in the adult only the proximal bones can be clearly +distinguished, the distal ones having become completely ankylosed with +the metacarpals to form the =carpo-metacarpus=. + +The two distinct carpal bones are the radial carpal and the ulnar +carpal. The radial carpal (fig. 57, 4) is a small somewhat cubical +bone, wedged in between the manus and the radius and ulna. The ulnar +carpal (fig. 57, 5) is a somewhat larger, more irregular bone, lying +adjacent to the end of the ulna. It is deeply notched to receive the +carpo-metacarpus. + +=The hand.= In the adult bird the hand is in a much modified +condition; only the first three digits are represented, and the +metacarpals are all fused with one another and with the distal +carpalia to form the =carpo-metacarpus=. + +The most prominent part of the carpo-metacarpus is formed by the +=second metacarpal= (fig. 57, 7), a stout, straight bone expanded at +both ends. The =third metacarpal= (fig. 57, 8) is a more slender +curved bone fused at both ends with the second metacarpal. The =first +metacarpal= forms simply a small projection on the radial side of the +proximal end of the second metacarpal. + +The =phalanges=. The first digit or =pollex= includes two phalanges, +the distal one being very small and bearing a claw. + +The second digit includes three phalanges, the proximal one being +somewhat flattened. The third digit has a single small phalanx. + + +THE PELVIC GIRDLE. + +The bones constituting the pelvic girdle are not only as in other +higher vertebrates ankylosed together forming the innominate bones, +but are also ankylosed with a series of some seventeen sacral and +pseudosacral vertebrae. The =acetabulum= (fig. 61, 5) with which the +head of the femur articulates is incompletely ossified. + +The =ilium= (figs. 58 and 61, 1) is the largest bone of the pelvis. It +forms a long flattened plate extending for a considerable distance +both in front of and behind the acetabulum, and is fused along its +whole length with the transverse processes and neural spines of the +sacral and pseudosacral vertebrae. It forms more than half the +acetabulum, above and behind which it is produced to form a process, +the =antitrochanter= (fig 61, 8), with which the great trochanter of +the femur articulates. + +[Illustration FIG. 61. LATERAL VIEW OF THE PELVIS AND SACRUM OF A DUCK +(_Anas boschas_) × 2/3. + + 1. ilium. + 2. ischium. + 3. pubis. + 4. pectineal process. + 5. acetabulum. + 6. ilio-sciatic foramen. + 7. fused vertebrae. + 8. antitrochanter.] + +The =ischium= (figs. 58 and 61, 2) is a flattened bone which forms +about one-third of the acetabulum, and lies ventral to the posterior +part of the ilium. Its anterior portion is separated from the ilium by +the large oval =ilio-sciatic foramen= (fig. 61, 6), while behind this +the two bones are completely fused. + +The =pubis= (figs. 58 and 61, 3) is a very long slender bar of bone +which forms only a very small part of the acetabulum and runs back +parallel to the ventral surface of the ischium with which it is +loosely connected at its posterior end. For the greater part of their +length the two bones are separated by the long narrow =obturator +foramen=. Behind the ischium the pubis is produced into a long curved +downwardly-projecting process, and in front of the acetabulum it bears +a short blunt =pectineal= or =pre-pubic process= (fig. 61, 4) probably +homologous with the pre-pubis of Orthopod Dinosaurs. The remainder of +the pubis is homologous with the post-pubis of Orthopod Dinosaurs. + + +THE POSTERIOR LIMB. + +The leg of the bird is somewhat differently constructed from that of +other vertebrates owing to the fact that there is no free tarsus, the +proximal tarsals having fused with the tibia, and the distal with the +metatarsals. + +The =thigh= consists of a single bone, the femur. The =femur= is a +comparatively short bone with a straight shaft and expanded ends. The +proximal end bears on its inner side a rounded =head=, which +articulates with the acetabulum. On its outer side is an irregular +outgrowth, the =great trochanter=, while between the two is the +surface which meets the antitrochanter of the ilium. The posterior end +also is expanded and marked by a wide groove which lodges the +=patella=. On each side of the groove is a strong =condylar ridge= for +articulation with the tibia. The external condyle is deeply grooved +behind for articulation with the fibula. + +The =crus= or =shin= consists of two separate bones, (1) the +=tibio-tarsus=, formed by the fusion of the tibia with the proximal +row of tarsals, and (2) the =fibula=. + +The =tibio-tarsus= is a thick straight bone nearly twice as long as +the femur. Both ends of the bone are considerably expanded. The +proximal end bears two slight depressions which articulate with the +condyles of the femur, and a third depression which partly lodges the +patella. The proximal end of the anterior or extensor surface is drawn +out into a very prominent =cnemial crest= which bends over towards the +postaxial side of the bone; a slight ridge is continued from it all +the way down the shaft. The proximal part of the shaft of the +tibio-tarsus bears a roughened ridge with which the fibula is closely +connected. The distal end is expanded and rotated outwardly and forms +a prominent pulley-like surface which articulates with the +tarso-metatarsus. + +The =fibula= is reduced to the proximal portion only, which is +expanded and articulates with a depression behind the external condyle +of the femur. The fibula further extends about a third of the way down +the shaft of the tibio-tarsus. The =patella= or =knee-cap= is a +sesamoid bone due to an ossification in the tendon of the extensor +muscles of the leg. + +The =ankle joint= lies between the proximal and distal tarsals which +as previously mentioned fuse respectively with the tibia and +metatarsus. + +The =Pes=. The pes includes four digits, and consists of the +tarso-metatarsus and the phalanges. The proximal tarsals which are +fused with the tibia also really belong to the pes. + +The =tarso-metatarsus= is a strong straight bone nearly as long as the +femur, and is formed by the fusion of the distal tarsals with the +second, third and fourth metatarsals. The proximal end of the bone is +expanded and bears two facets for articulation with the tibio-tarsus, +and near them on the posterior surface is a large roughened +projection. The lines of junction between the several metatarsals are +marked along the shaft by slight ridges. At the distal end of the bone +the three metatarsals diverge from one another and each bears a +prominent convex pulley-like surface. The =first metatarsal= is +reduced to the distal end, which tapers to a point proximally, and is +attached by ligaments near the distal end of the tarso-metatarsus. + +The =digits=. Four digits are present, each consisting of a metatarsal +(already described) and a certain number of phalanges, the terminal +one being in each case clawed. The first digit or =hallux= has two +phalanges, the second three, the third four, and the fourth five. + + +FOOTNOTES: + +[101] See R.S. Wray, _P.Z.S._, 1887, p. 343. + +[102] Often called the manubrium, but not homologous with the +manubrium of the mammalian sternum. + +[103] Cp. fig. 63. + +[104] See p. 28. + + + + +CHAPTER XIX. + +GENERAL ACCOUNT OF THE SKELETON IN BIRDS. + + +EXOSKELETON. + +The epidermal exoskeleton of birds is very greatly developed, feathers +constituting its most important part. + +Three kinds of feathers are found, viz. (_a_) _pennae_ including +quills and coverts, (_b_) down feathers or _plumulae_, and (_c_) +_filoplumes_ which are rudimentary feathers. The structure of the +different kinds of feathers is described on pp. 303-306. + +Sometimes a fourth class of feathers, the _semiplumae_, is recognised. +They have the stems of pennae, and the downy barbs and barbules of +plumulae. + +In most birds the pennae are not uniformly distributed over the whole +surface of the body, but are confined to certain tracts, the +=pterylae=; while the intervening spaces or =apteria= are either bare +or covered only with down feathers. In some birds, however, such as +the Ratitae and the Penguins, pennae are evenly distributed over the +whole body. + +In many birds the calamus or quill bears two vexilla or vanes, the +second of which, called the =aftershaft= or =hyporachis=, is generally +much the smaller, and is attached to the under surface of the main +vexillum. In the Moas, Emeu and Cassowary the two vexilla in the adult +bird are nearly equal in size; though in the nestling Emeu one is much +longer than the other. The aftershaft is very small in most Passeres +and gallinaceous birds, but is comparatively large in Parrots, Gulls, +Herons and most birds of prey. It is absent or extremely small in the +Ostrich, _Apteryx_, _Rhea_, Pigeons, Owls, Anseres, and others. + +The quill feathers include two groups, the =remiges= or wing quills, +and the =rectrices= or tail quills. In most birds the primary remiges, +or those which are attached to the bones of the manus, are ten or +eleven in number, and are set in grooves in the bones, being firmly +attached to them. In the Ostrich however the primaries are little +specialised in character and are as many as sixteen in number. They +are also less definitely attached to the bones; as their ends do not +lie in grooves in the bones, but project beyond them. + +The secondary quills or those attached to the ulna vary much in number +according to the length of the bone. The large dark quills in the +wings of Cassowaries are the secondaries. + +The wing of Penguins is very little differentiated. It is covered at +the margin by overlapping scales which gradually merge into scale-like +feathers at the proximal end. The wing of the Penguin has nothing +comparable to the remiges of other birds. + +In some birds, such as Herons (_Ardea_), there occur in places +plumulae of a peculiar kind, which grow persistently and whose summits +break off into fine powder as fast as they are formed. These feathers +are known as _powder-down_ feathers. They occur also in some Parrots +and are then scattered indiscriminately all over the body. + +Other exoskeletal structures besides feathers are commonly well +developed. Thus the extremities of the jaws are sheathed in horny +=beaks= whose form varies enormously according to the special mode of +life. + +In ducks and geese the beak with the exception of the anterior end is +soft, and its edges are raised into lamellae, while in the Mergansers +these lamellae become pointed processes supported by bony outgrowths. +These lamellae act as strainers. In Parrots and Hawks, on the other +hand, nearly the whole of the beak is hard. + +The toes and tarso-metatarsus are usually featherless and are covered +either with granular structures or with well-formed scales. The toes +are nearly always provided with =claws=, and these vary in correlation +with the character of the beak. Claws[105] also sometimes occur on the +manus. Thus _Archaeopteryx_ and some Ostriches and Rheas have claws on +all three digits. Most Ostriches and Rheas, and many Anseres and birds +of prey, have them on the first two digits, while the Secretary bird +(_Gypogeranus_) and many fowls, ducks, and birds of prey, especially +kestrels, have a claw only on the pollex. In the Cassowary, Emeu, +Apteryx and some Ostriches and Rheas only the second digit is clawed. + +Claws should not be confounded with =spurs=, which are conical horny +structures developed on bony outgrowths of the radial side of the +carpus, metacarpus, or metatarsus. They occur in a number of birds, +but are most commonly developed in gallinaceous birds, by which they +are used for fighting. A single spur occurs on the metacarpus in +_Megapodius_, in _Palamedea_, in _Parra jacana_ and in _Hoplopterus +spinosus_, the Spur-winged plover. The Derbian Screamer, _Chauna +derbiana_, has two metacarpal spurs, borne on the first and second +metacarpals. The Spur-winged goose, _Plectropterus gambensis_, has a +carpal spur borne on the radial carpal. Metatarsal spurs are quite +common. + +The male Solitaire (_Pezophaps_) has large bony excrescences on the +wrist which may, like spurs, have been sheathed in horn and used for +fighting. + +=Teeth= do not occur in any living birds, but conical teeth imbedded +in separate sockets are present in _Archaeopteryx_ and _Ichthyornis_, +while in _Hesperornis_ similar teeth occur implanted in continuous +grooves in the mandibles and maxillae, the premaxillae being toothless. + +Except that teeth are partly dermal in origin, a dermal exoskeleton is +quite unrepresented in birds. + + +ENDOSKELETON. + +Perhaps the most striking feature of the endoskeleton of birds is its +pneumaticity. In the embryo all the bones contain marrow, but as +growth proceeds this becomes replaced by air to a variable extent in +different forms. In all birds some part of the skeleton is pneumatic. +Many small birds and _Apteryx_ and Penguins among larger ones have air +only in the skull; in Pigeons air is present in all the bones except +the caudal vertebrae, the leg bones, and those of the antibrachium and +manus; in Hornbills every bone contains air. + +[Illustration FIG. 62. THIRD CERVICAL VERTEBRA OF AN OSTRICH +(_Struthio camelus_). × 1. A anterior, B posterior, C dorsal view (A +and B after MIVART). + + 1. neural spine. + 2. neural canal. + 3. prezygapophysis. + 4. postzygapophysis. + 5. posterior articular surface of centrum. + 6. anterior articular surface of centrum. + 7. vertebrarterial canal. + 8. hypapophysis.] + + +VERTEBRAL COLUMN. + +The vertebral column of birds is readily divisible into a very mobile +cervical region, and an extremely rigid post-cervical region. In most +birds the vertebral centra are without terminal epiphyses, but these +structures are found in Parrots. The cervical vertebrae are generally +large and vary in number from eight or nine to twenty-three in Swans. +Except in some extinct forms, such as _Ichthyornis_ and _Apatornis_, +in which they are biconcave, the centra are characterised by having +saddle-shaped articulating surfaces, which in front are concave from +side to side and slightly convex from above downwards, while +posteriorly they are convex from side to side and concave from above +downwards. The atlas is small and ring-like, and its centrum is fused +with the axis forming the odontoid process. Cervical ribs are often +well developed, and in some of the Ratitae they remain for a long time +distinct from the vertebrae. + +The thoracic vertebrae are distinguished from the cervical by the fact +that their true ribs are united to the sternum by means of sternal +ribs. This distinction, however, though convenient, is somewhat +arbitrary, as it has been shown that in the fowl and gannet, two pairs +of ribs which in the adult are free from the sternum, are connected +with it in the embryo. When, as in the Swans, the thoracic vertebrae +are not all fused together, they generally have saddle-shaped +articulating surfaces, but sometimes, as in the Penguins, Auks and +Plovers, the centra are convex in front and concave behind. The trunk +vertebrae generally have well-marked neural spines, while in the +Divers the anterior ones have peculiar bifurcating hypapophyses. + +The trunk vertebrae are not readily divisible into thoracic and +lumbar. There are two true sacral vertebrae, but as development +proceeds a number of other vertebrae become fused with the true +sacrals, the whole forming a large compound sacrum. These pseudosacral +vertebrae generally include the lumbar, and some of the thoracic and +caudal vertebrae. Sixteen to twenty vertebrae or even more may be +included in the compound sacrum, and sometimes the whole of the trunk +vertebrae are fused together. In _Archaeopteryx_ however but five +vertebrae take part in the formation of the sacrum. + +In _Archaeopteryx_ there are twenty long caudal vertebrae, of which +the last sixteen carry a pair of feathers apiece, but in all other +birds the tail is short and in the great majority of cases the +posterior vertebrae are fused together, forming the pygostyle. In the +Ratitae and Tinamidae a pygostyle is rarely or imperfectly developed. +In _Hesperornis_ there are twelve caudal vertebrae, six or seven of +which are united by their centra only, forming an imperfect pygostyle. + +The free caudal vertebrae are generally amphicoelous. + + +THE SKULL. + +The skull of all birds from _Archaeopteryx_ onwards is essentially +similar, differing from the skull of reptiles mainly in the extent to +which the cranium is arched, and its greater size in proportion to the +jaws. + +Most of the bones of the cranium are pneumatic, and all show a marked +tendency to fuse together, and have their outlines obliterated by the +disappearance of the sutures. The several bones remain longest +distinguishable in the Ratitae and to a less extent in the Penguins. +The orbits are very large and lie almost entirely in front of the +cranium; they are separated by an interorbital septum which is +sometimes, as in _Chauna_ and _Scythrops_, very complete, sometimes, +as in Hornbills and the Common Heron, very slightly developed. As a +general rule the sclerotic is cartilaginous. + +The anterior nares are almost always situated far back at the base of +the beak near the orbits, but in _Apteryx_ they are placed right at +its extremity. In _Phororhacos_ they are placed very high up on the +enormous beak and are not separated by any bony partition. + +The skull of Parrots has some peculiarities. In some Parrots the +lachrymal sends back a process which meets the postorbital process of +the frontal and completes the orbit. In most birds the upper beak is +immovably fixed, but in some it is attached to the cranium, only by +the nasals and by flexible processes of the premaxillae, so that by +this means a kind of elastic joint is established and the beak is able +to be moved on the cranium. In the Parrots and _Opisthocomus_ there is +a regular highly movable joint. + +In Cassowaries the fronto-nasal region of the skull is produced into +an enormous bony crest, and in Hornbills a somewhat similar structure +occurs. Although true teeth do not occur in any known bird except +_Archaeopteryx_, _Hesperornis_, and _Ichthyornis_, another extinct +bird, _Odontopteryx_, has the margins of both jaws provided with +forwardly-directed tooth-like serrations, formed of part of the actual +jawbone: a living hawk, _Harpagus_, too, has a deeply notched bill, to +which correspond serrations in the premaxillae. + +A basi-pterygoid process of the basisphenoid abuts against the +pterygoid in Ratitae and in Tinamous, plovers, fowls, pigeons, ducks +and geese among Carinatae, recalling the arrangement met with in many +reptiles. The squamosal is sometimes, as in the fowl, united with the +postorbital process of the frontal. In the Carinatae the quadrate +articulates with the cranium by a double convex surface, in the +Ratitae by a single one. The premaxillae are always comparatively large +bones, the maxillae on the contrary are small, but give rise to +important inwardly-projecting maxillo-palatine processes. + +The relations of the palatines, pterygoids, maxillae, and vomers vary +considerably, and on them Huxley has based a classification of +birds[106]. In the Ratitae and the Tinamous (Tinamidae), among +Carinatae the vomers unite and form a large broad bone, separating the +palatines and the pterygoids from the rostrum. Huxley uses the term +=Dromaeognathous= to describe this condition. In all other Carinatae +the vomers are narrow behind, and the palatines and pterygoids +converge posteriorly and articulate largely with the rostrum. Three +modifications of this condition are distinguished by Huxley, and +termed =Schizognathous=, =Ægithognathous=, and =Desmognathous=. + +In the =Schizognathae= the vomers coalesce and form a narrow elongated +bone, pointed in front, separating the maxillo-palatine processes of +the premaxillae. Waders, fowls, penguins, gulls, some falcons and +eagles, American vultures, some herons and many owls have the +Schizognathous arrangement. In pigeons and sandgrouse there is no +vomer, but the other bones have the Schizognathous arrangement. + +In the =Ægithognathae= the arrangement is the same as in the +Schizognathae, except that the vomers are truncated in front. +Passeres, swifts, woodpeckers, humming birds, rollers, hoopoes have +this arrangement. + +In the =Desmognathae= (fig. 60, A) the maxillo-palatine processes +approach one another in the middle line, and either unite with the +vomers, or unite with one another, hiding the vomers. Thus a more or +less complete bony roof is formed across the palate. The vomers in +Desmognathae are small or sometimes absent. Ducks, storks, most +herons, most birds of prey and owls, pelicans, cormorants, parrots, +and flamingoes are Desmognathous. + +The mandible, as in other Sauropsids, consists of a cartilage bone, +the articular, and a series of membrane bones, the dentary, splenial, +coronoid, angular, and supra-angular, developed round the unossified +Meckel's cartilage. The dentaries of the two rami are nearly always +fused together, but in _Ichthyornis_ and _Archaeopteryx_ the two rami +are but loosely united. There is often a fontanelle between the +dentary and the posterior bones, while the angle is sometimes, as in +the fowl, drawn out into a long curved process. + +The hyoid apparatus (fig. 59, C) consists of a median portion, and a +pair of cornua. The median portion is composed of three pieces placed +end to end, and called respectively the os entoglossum, the basi-hyal, +and the uro-hyal. The os entoglossum is shown by development to be +formed by the union of paired structures and is probably homologous +with the hyoid arch of fishes. The basi-hyal and the long cornua, each +of which is composed of two or three pieces placed end to end, are +homologous with the first branchial arch of fishes, while the uro-hyal +is probably homologous with the second branchial arch of fishes. In +Woodpeckers the cornua are enormously long, and curve over the skull, +extending as far forwards as the anterior nares. + + +RIBS AND STERNUM. + +Well-developed ribs are attached to the posterior cervical vertebrae +as well as to the thoracic vertebrae. The ribs generally have uncinate +processes and separate capitula and tubercula, but uncinate processes +are absent in _Chauna Palamedea_ and apparently in _Archaeopteryx_. + +The sternum (fig. 63) is greatly developed in all birds. In the +embryo[107] it is seen to be derived from the union of right and left +plates of cartilage, formed by the fusion of the ventral ends of the +ribs. In the Ratitae and a few Carinatae, such as _Stringops_, it is +flat, but in the great majority of birds it is keeled, though the +development of the keel varies greatly. It is large in the flightless +Penguins, which use their wings for swimming. Traces of an +interclavicle may occur in the embryo. + + +PECTORAL GIRDLE. + +[Illustration FIG. 63. SHOULDER-GIRDLE AND STERNUM OF + + A. BLACK VULTURE (_Vultur cinereus_) × 1/3. + B. PEACOCK (_Pavo cristatus_) × 3/8. + C. PELICAN (_Pelicanus conspicillatus_) × 1/3. (All Camb. Mus.) + + 1. carina of the sternum. + 2. coracoid. + 3. scapula. + 4. clavicle. + 5. costal process. + 6. surfaces for articulation with the sternal ribs. + 7. xiphoid processes. + 8. fontanelle.] + +The pectoral girdle is also strongly developed in all Carinatae, but +is much reduced in Ratitae. In some Moas the sternum has no facet for +the articulation of the coracoid, and the pectoral girdle appears to +have been entirely absent; it is extremely small also in _Apteryx_. +Clavicles are generally well developed in the Carinatae, and small +ones are found also in _Hesperornis_, and in Emeus and Cassowaries. In +the other living Ratitae and in _Stringops_ they are absent. In some +Parrots, Owls and Toucans they do not meet one another ventrally. +Clavicles are especially stout in some of the birds of prey. They do +not generally touch the sternum, but sometimes, as in the Pelican +(fig. 63, C), Adjutant and Frigate bird, they are fused with it. + +In all Ratitae the scapula and coracoid lie almost in the same +straight line with one another, in the Carinatae they are nearly at +right angles to one another. + + +ANTERIOR LIMB. + +In the wing of nearly all birds the ulna is thicker than the radius, +but in _Archaeopteryx_ the two bones are equal in size. In the wing of +_Archaeopteryx_ there are three long digits with distinct metacarpals. +In all other birds the digits are modified, the metacarpals being +commonly fused and the phalanges reduced in number. In _Palamedea_ and +some other birds the metacarpus bears a bony outgrowth, which when +sheathed in horn forms a spur. + +In most of the Ratitae and in the extinct Dodo (_Didus_) and Solitaire +(_Pezophaps_) the wing is very small, but the usual parts are +recognisable. In _Hesperornis_ apparently only the humerus is present; +in some Moas, in which the wing is imperfectly known, the presence of +the humerus is indicated by traces of a glenoid cavity. In most Moas +the wing is apparently completely absent. As compared with those in +other Ratitae, the wings of the Ostrich and Rhea are well developed. +In the Ostrich (fig. 64, B) and Rhea, as in nearly all Carinatae, the +manus has three digits, but in _Apteryx_ there is only a single digit, +the second. The Penguins (fig. 64, A) too among Carinatae have only +two digits, but in their case it is the pollex which is missing. In +the Ostrich the third digit has two phalanges, in all other living +birds it has only one phalanx. + + +PELVIC GIRDLE. + +[Illustration FIG. 64. BONES OF THE RIGHT WING OF + + A. A PENGUIN × 1/3. (Camb. Mus.) + B. OSTRICH (_Struthio camelus_) × 1/7. (Partly after PARKER.) + C. GANNET (_Sula alba_) × 1/3. (Camb. Mus.) + +In C the distal phalanges of the pollex and second digit have been +omitted. + + 1. humerus. + 2. radius. + 3. ulna. + 4. second metacarpal. + 5. third metacarpal. + 6. pollex. + 7. second digit. + 8. cuneiform. + 9. sesamoid bone.] + +Birds have a very large pelvis and its characters are constant +throughout almost the whole group. The ilium is very large, and is +united along its whole length with the sacral and pseudosacral +vertebrae. The ischium is broad and extends back parallel to the ilium +with which in most birds it fuses posteriorly, further forward the +ilio-sciatic foramen separates the two bones. In _Tinamus_, +_Hesperornis_, _Apteryx_ (fig. 65, B, 2), and _Struthio_, the ischia +are separate from the ilia along their whole length except at the +acetabulum; in _Phororhacos_, on the other hand, the two bones are +fused along almost their whole length. The bone usually called the +pubis in birds corresponds to the post-pubis of Dinosaurs and forms a +long slender rod (fig. 65, 3) lying parallel to the ischium. In many +birds the ischia and pubes are united at their distal ends. This is +the case in the Ostrich (fig. 65, D), in which the ilia and ischia are +widely separated. In many birds the pubis is drawn out in front into +the pectineal process, this is specially large in _Apteryx_ (fig. 65, +B, 5), and in the embryos of many birds. It is probably homologous +with the pre-pubis of Dinosaurs but in some birds is formed in part by +the ilium. The acetabulum in birds is always perforate. + +In _Rhea_ (fig. 65, C, 2) and probably in _Archaeopteryx_ a symphysis +ischii occurs, and in the ostrich alone among birds there is a +symphysis pubis. In _Archaeopteryx_ all three bones of the pelvis are +distinct, but they are imperfectly known. In Ichthyornis they are also +distinct, in all other known birds they are fused together to a +greater or less extent. + +[Illustration FIG. 65. PELVIC GIRDLE AND SACRUM OF + + A. CASSOWARY (_Casuarius galeatus_) × 1/8. + B. OWEN'S APTERYX (_A. oweni_) × 1/2. + C. BROAD BILLED RHEA (_R. macrorhyncha_) × 1/6. + D. OSTRICH (_Struthio camelus_) × 1/10. (All Camb. Mus.) + + 1. ilium. + 2. ischium. + 3. pubis. + 4. acetabulum. + 5. pectineal process.] + + +POSTERIOR LIMB. + +The tibia is always well developed and has a very strong cnemial +crest. The proximal tarsals are fused with its distal end, the whole +forming a compound bone, the tibio-tarsus. There is frequently an +oblique bar of bone crossing the anterior face of the tibio-tarsus at +the distal end, just above the articular surface of the +tarso-metatarsus, this is absent in Ostriches and _Æpyornis_. The +fibula though in the embryo and in _Archaeopteryx_ equal in length to +the tibia, is in the adult of other birds always imperfect, its +proximal end is often fused with the tibia, and its distal end is +commonly atrophied. In the Penguins however the distal end is +complete. The distal tarsals fuse with the second, third and fourth +metatarsals, forming a compound bone, the tarso-metatarsus. The first +metatarsal is nearly always free but occasionally as in _Phaëthon_ it +is fused with the others. No adult bird has more than four digits in +the pes. In the Penguins the metatarsals are separate, and in many +birds larger or smaller gaps exist between the fused metatarsals. In +most birds the third metatarsal is curved so as not to lie in the same +plane as the others, but in the Penguins they all three lie in the +same plane. The metatarsals are clearly separated in _Archaeopteryx_. +In Gallinaceous birds the tarso-metatarsus bears a bony outgrowth +which is sheathed in horn and forms a spur. + +In most birds the first four toes are present while the fifth is +always absent. The first toe commonly has two phalanges, the second +three, the third four, and the fourth five. In Swifts the third and +fourth toes have only three phalanges. Many birds, such as all Ratitae +except _Apteryx_, have only three toes, the hallux being absent; in +the Ostrich the second toe is also gone with the exception of a small +metatarsal, so that the foot retains only the third and fourth digits, +the third being much the larger of the two and bearing a claw, while +the fourth is clawless. + +In the Swifts, Cormorants, and Penguins, all four toes are directed +forwards. In most birds the hallux is directed backwards, and the +other toes forwards. In the Owls the fourth toe can be directed +backwards as well as the hallux, while in Parrots, Cuckoos, +Woodpeckers, and Toucans the fourth toe is permanently reversed. In +Trogons the second toe is reversed in addition to the hallux, but not +the fourth. + + +FOOTNOTES: + +[105] W.K. Parker, _Phil. Trans._ vol. 179, p. 385, 1888; and _Ibis_, +1888, p. 124. + +[106] See T.H. Huxley, "On the Classification of Birds," _P.Z.S._ +1867. + +[107] B. Lindsay, _P.Z.S._ 1885, p. 684. + + + + +CHAPTER XX. + +CLASS MAMMALIA. + + +The skeleton of the members of this class, the highest of the +vertebrata, has the following characteristics:-- + +Some part of the integument at some period of life is always provided +with hairs; these are epidermal structures arising from short papillae +of the Malpighian layer of the epidermis, which at once grow inwards +and become imbedded in pits of the dermis. Sometimes scales or spines +occur, and epidermal exoskeletal structures in the form of hoofs, +nails, claws and horns are also characteristic. As regards the +endoskeleton, the vertebral centra have terminal epiphyses except in +the Ornithodelphia and some Sirenia. In the skull the cranial region +is greatly developed as compared with that in lower vertebrates, and +whereas in many reptiles the true cranium is largely concealed by a +false roof, in mammals the only relic of this secondary roof is found +in the zygomatic arch, and postorbital bar. In the adult all the bones +except the mandible, hyoid, and auditory ossicles are firmly united +together. The basisphenoid is well ossified, and there is no +parasphenoid. The pro-otic ossifies, and unites with the epi-otic and +opisthotic before they coalesce with any other bones. + +The skull articulates with the vertebral column by means of two convex +occipital condyles formed mainly by the exoccipitals, and the mandible +articulates with the squamosal without the intervention of the +quadrate. The latter is much reduced, and is converted into the +tympanic ring, while the hyomandibular of fish is represented by the +auditory ossicles[108]. + +The teeth are always attached to the maxillae, premaxillae and +mandibles, never to any of the other bones. They are nearly always +implanted in distinct sockets, and are hardly ever ankylosed to the +bone. The teeth of mammals are generally markedly heterodont, four +forms, incisors, canines, premolars, and molars, being commonly +distinguishable. Some mammals are _monophyodont_, having only a single +set of teeth, but the great majority are _diphyodont_, having two +sets, a deciduous or milk dentition, and a permanent dentition. + +The _incisors_, the front teeth, are simple, one-rooted, adapted for +cutting, and are nearly always borne by the premaxillae. Next come the +_canines_, one on each side in each jaw. They are generally large +teeth adapted for tearing or holding, and get their name from the fact +that they are largely developed in the dog. The remaining teeth form +the grinding series, the more posterior of them being the _molars_, +which are not preceded by milk teeth[109]. Between the molars and the +canines are the _premolars_, which do as a rule have milk or deciduous +predecessors, though very frequently the first of them is without a +milk predecessor. + +In describing the dentition of any mammal, for the sake of brevity a +formula is generally made use of. Thus, the typical mammalian +dentition is expressed by the formula + + _i_ 3/3 _c_ 1/1 _pm_ 4/4 _m_ 3/3 = 11/11, + +giving twenty-two teeth on each side, or forty-four altogether[110]. +The incisors are represented by _i_, the canines by _c_, the premolars +by _p_ or _pm_, and the molars by _m_. The numbers above the lines +represent the teeth in the upper jaw, those below the lines the teeth +in the lower jaw. The milk dentition is expressed by a similar formula +with _d_ (deciduous) prefixed to the letter expressing the nature of +the tooth. + +The following terms are of frequent use as characterising certain +forms of the grinding surfaces of teeth, and it will be well to define +them at once. + +_Bunodont_ is a term applied to teeth with broad crowns raised into +rounded tubercles, e.g. the grinding teeth of Pigs and Hippopotami; + +_Bilophodont_ to teeth marked by a simple pair of transverse ridges, +with or without a third ridge running along the outer border of the +tooth at right angles to the other two, e.g. the grinding teeth of +_Lophiodon_, Kangaroo, Manatee, Tapir, _Dinotherium_; + +_Selenodont_ to teeth marked by crescentic ridges running from the +anterior towards the posterior end of the tooth, e.g. the grinding +teeth of the Ox and Sheep. + +Teeth whose crowns are low so that their whole structure is visible +from the grinding surface are called _brachydont_, while those with +higher crowns, in which the bases of the infoldings of enamel are +invisible from the grinding surface are said to be _hypsodont_. +Bunodont teeth are brachydont, the teeth of the Horse and Ox are +hypsodont. + +Passing now to the appendicular skeleton--the shoulder girdle differs +markedly from that of Sauropsids in the fact that the coracoid, except +in the Ornithodelphia, is greatly reduced, generally forming only a +small process on the scapula. In the pelvis the pubes meet in a +ventral symphysis, except in some Insectivora and Chiroptera. In many +mammals a fourth pelvic element, the _acetabular bone_, is +distinguishable. The ankle joint is _cruro-tarsal_, or situated +between the proximal tarsal bones and the tibia and fibula. Carpalia 4 +and 5 are united forming the _unciform_; and the ulnar sesamoid bone +or _pisiform_ is generally well developed. In the proximal row of +tarsal elements there are only two bones, the calcaneum and +astragalus. Of these the calcaneum is the fibulare, and the +astragalus is generally regarded as the tibiale and intermedium +fused[111]. + + +_Subclass I._ ORNITHODELPHIA OR PROTOTHERIA. + +This subclass contains only a single order, the Monotremata, and the +following characteristics are equally applicable to the subclass and +to the order. The vertebral centra have no epiphyses, and the odontoid +process remains for a long time free from the centrum of the second +vertebra. With the exception of the atlas of _Echidna_ the cervical +vertebrae are without zygapophyses. The cranial walls are smooth and +rounded, and the sutures between the several bones early become +completely obliterated as in birds. The mandible is a very slight +structure, with no ascending ramus, and with the coronoid process (see +p. 398) and angle rudimentary. The auditory ossicles show a low state +of development. The tubercula of the ribs articulate with the sides of +the centra of the thoracic vertebrae, not with the transverse +processes. Some of the cervical ribs remain for a long time separate +from the vertebrae. Well ossified sternal ribs occur. No true teeth +are present in the adult. The young _Ornithorhynchus_ has functional +molar teeth, but in the adult their place is taken by horny plates. In +the Echidnidae neither teeth nor horny plates occur. + +The coracoid (fig. 66, 3) is complete and well developed, and +articulates with the sternum. A precoracoid (epicoracoid) occurs in +front of the coracoid, and there is a large interclavicle (fig. 66, +6). The ridge on the scapula, corresponding to the spine of other +mammals, is situated on the anterior border instead of in the middle +of the outer surface. Epipubic bones are present. In the Echidnidae, +but not in _Ornithorhynchus_[112], the central portion of the +acetabulum is unossified as in birds. The humerus has a prominent +deltoid crest; its ends are much expanded, and the distal end is +pierced by an ent-epicondylar foramen. The fibula has a broad proximal +process resembling an olecranon. The limbs and their girdles bear a +striking resemblance to those of some Theromorphous reptiles. + +[Illustration FIG. 66. VENTRAL VIEW OF THE SHOULDER-GIRDLE AND STERNUM +OF A DUCKBILL (_Ornithorhynchus paradoxus_) × 3/4 (after PARKER). + + 1 and 2. scapula. + 3. coracoid. + 4. precoracoid (epicoracoid). + 5. glenoid cavity. + 6. interclavicle. + 7. clavicle. + 8. presternum. + 9. third segment of mesosternum. + 10. sternal rib. + 11. intermediate rib. + 12. vertebral rib.] + +The order Monotremata includes only two living families, the +Echidnidae and Ornithorhynchidae. + + +MESOZOIC MAMMALIA[113]. + +It will be well here to briefly refer to certain mammals of small +size, the remains of which have been found in deposits of Mesozoic +age. In the great majority of cases they are known only by the lower +jaw, or sometimes only by isolated teeth. A large number of them are +commonly grouped together as the Multituberculata, and are sometimes, +partly owing to the resemblance of their teeth to those of +_Ornithorhynchus_, placed with the Prototheria, sometimes between the +Prototheria and the Metatheria. They are characterised by having a +single pair of large incisors in the lower jaw, and one large with one +or two smaller incisors in each premaxillae. The lower canines are very +small or altogether wanting. The incisors are separated by a diastema +from the grinding teeth, which are sometimes (_Tritylodon_) +characterised by the possession of longitudinal rows of little +tubercles separated by grooves, sometimes by having the premolars +provided with high cutting edges, whose surfaces are obliquely +grooved. Some of the Mesozoic mammals found associated with the +Multituberculata, have however a dentition of an altogether different +type, with at least three lower incisors, well developed canines and +premolars, and numerous molars with peculiar three-cusped or +tritubercular grinding surfaces. These mammals, one of the best known +of which is _Phascolotherium_, are commonly separated from the +Multituberculata, and are divided by Osborn into two groups, one +allied to the Marsupials, and one to the Insectivores. The group +showing Marsupial affinities is further subdivided into carnivorous, +omnivorous, and herbivorous subgroups. The members of both groups +commonly have four premolars, and six to eight molars in each +mandibular ramus. + + +_Subclass II._ DIDELPHIA OR METATHERIA. + +This subclass, like the previous one, contains only a single order, +viz. the Marsupialia[114]; but the forms referable to it are far more +numerous than in the case of the Monotremata. + +The integument is always furry, and the teeth are always +differentiated into incisors, canines, premolars and molars. Except in +_Phascolomys_, the number of incisors in the upper and lower jaws is +never equal, and the number in the upper usually exceeds that in the +lower jaw. There is no such regular succession and displacement of +teeth as in most mammals. Sometimes the anterior teeth are diphyodont, +and as a general rule the tooth commonly regarded as the last premolar +is preceded by a milk tooth. The majority of the permanent teeth of +most Marsupials are regarded as belonging to the milk series for two +reasons, (1) they are developed from the more superficial tissues of +the jaws, (2) a second set, the permanent teeth, begin to develop as +outgrowths from them, but afterwards become aborted[115]. + +The odontoid process at an early stage becomes fused with the centrum +of the second cervical vertebra, and the number of thoraco-lumbar +vertebrae is always nineteen. The skull has several characteristic +features. The tympanic bone remains permanently distinct, and the +anterior boundary of the tympanic cavity is formed by the alisphenoid. +The carotid canal perforates the basisphenoid, and the lachrymal canal +opens either outside the orbit or at its margin. There are generally +large vacuities in the palate. The angle of the mandible is (except in +_Tarsipes_) more or less inflected; and as a rule the jugal furnishes +part of the articular surface for the mandible. There is no +precoracoid (epicoracoid) or interclavicle, and the coracoid is +reduced to form a mere process of the scapula, not coming near the +sternum. + +Epipubic, or so-called marsupial bones[116], nearly always occur, and +a fourth pelvic element, the acetabular bone, is frequently developed. +The fibula is always complete at its distal end, sometimes it is fused +with the tibia, but often it is not only free but is capable of a +rotatory movement on the tibia. This is the case in the families +Phascolomyidae, Didelphyidae, and Phalangeridae. + +The Marsupialia can be subdivided into two main groups, according to +the character of the teeth:-- + + +1. POLYPROTODONTIA. + +In this group the incisors are small, subequal and numerous, not less +than 4/3. The canines are larger than the incisors, and the molars +have sharp cusps. The members of this group are all more or less +carnivorous or insectivorous. The group includes the families +Didelphyidae, Dasyuridae, Peramelidae, and Notoryctidae[117]. + + +2. DIPROTODONTIA. + +In this group the incisors do not exceed 3/3, and are usually 3/1, +occasionally 1/1. The first upper and lower incisors are large and +cutting. The lower canines are always small or absent, and so in most +cases are the upper canines. The molars have bluntly tuberculated, or +transversely ridged crowns. The group includes the families +Phascolomyidae, Phalangeridae, Macropodidae, and Epanorthidae. + + +_Subclass III._ MONODELPHIA OR EUTHERIA. + +This great group includes all the Mammalia except the orders +Monotremata and Marsupialia. Coming to their general characteristics-- +as in the Didelphia the odontoid process and cervical ribs early +become fused with the centra which bear them, while the coracoid is +reduced so as to form a mere process on the scapula, and there is no +precoracoid (epicoracoid), such as is found in Ornithodelphia. +Clavicles may be present or absent; when fully developed they +articulate with the sternum, usually directly, but occasionally, as in +some Rodents and Insectivores, through the remains of the sternal end +of the precoracoid. There is never any interclavicle in the adult, +though sometimes traces of it occur during development. In the pelvis +the acetabula are imperforate; and well-developed epipubic bones are +never found in the adult, though traces of them occur in some +Carnivores and foetal Ungulates. + + +_Order 1._ EDENTATA[118]. + +Teeth are not, as the name of the order seems to imply, always +wanting; and sometimes they are very numerous. They are, however, +always imperfect, and, with very few exceptions, are homodont and +monophyodont. They have persistent pulps, and so grow indefinitely and +are never rooted. In all living forms they are without enamel, +consisting merely of dentine and cement, and are never found in the +front part of the mouth in the situation occupied by the incisors of +other mammals. These characters derived from the teeth are the only +ones common to the various members of the order, which includes the +living sloths, ant-eaters, armadillos, pangolins and aard-varks, +together with various extinct forms, chiefly found in beds of late +tertiary age in both North and South America, the best known being the +Megatheridae and Glyptodonts. + + +_Order 2._ SIRENIA[119]. + +The skeleton of these animals has a general fish-like form, in +correlation with their purely aquatic habits. The fore limbs have the +form of paddles, but the number of phalanges is not increased beyond +the normal. There are no external traces of hind limbs. + +The whole skeleton and especially the skull and ribs is remarkably +massive and heavy. The dentition varies; in the two living genera +_Manatus_ and _Halicore_, incisor and molar teeth are present, in one +extinct genus, _Rhytina_, teeth are entirely absent, while in another, +_Halitherium_, the dentition is more decidedly heterodont than in +living forms. In the two living genera the dentition is monophyodont, +but in _Halitherium_ the anterior grinding teeth are preceded by milk +teeth. The tongue and anterior part of the palate and lower jaw are +covered with roughened horny plates. The skull is noticeable for the +size and backward position of the anterior nares, also for the absence +or small size of the nasal bones. There is no union of certain of the +vertebrae to form a sacrum, and in living forms the centra are not +terminated by well-formed epiphyses[120]. + +The cervical vertebrae are much compressed, but they are never +ankylosed together. In _Manatus_ there are only six cervical +vertebrae. The caudal vertebrae have well-developed chevron bones. The +humerus is distinctly articulated to the radius and ulna, and these +two bones are about equally developed, and are often fused together. +There are no clavicles, and the pelvis is vestigial, consisting of a +pair of somewhat cylindrical bones suspended at some distance from the +vertebral column. In living forms there is no trace of a posterior +limb, but in _Halitherium_ there is a vestigial femur connected with +each half of the pelvis. + + +_Order 3._ CETACEA[121]. + +In these mammals the general form is more fish-like than is the case +even in the Sirenia. The skin is generally almost completely naked, +but hairs are sometimes present in the neighbourhood of the mouth, +especially in the foetus. In some Odontoceti vestiges of dermal +ossicles have been described, and in _Zeuglodon_ the back was probably +protected by dermal plates. The anterior limbs have the form of +flattened paddles, showing no trace of nails, the posterior limb bones +are quite vestigial or absent, and there is never any external sign of +the limb. Teeth are always present at some period of the life history, +but in the whalebone whales they are only present during foetal life, +their place in the adult animal being taken by horny plates of baleen. +In all living forms the teeth are simple and uniform structures +without enamel; they have single roots, and the alveoli in which they +are imbedded are often incompletely separated from one another. As in +some forms traces of a replacing dentition have been described, it has +been concluded that the functional teeth of Cetacea belong to the milk +dentition. + +The texture of the bones is spongy. The cervical vertebrae are very +short, and though originally seven in number, are in many forms +completely fused, forming one solid mass (fig. 67). The odontoid +process of the axis is short and blunt, or may be completely wanting. +The lumbar and caudal vertebrae are large and numerous, and as +zygapophyses are absent, are very freely movable on one another; +zygapophyses are also absent from the posterior thoracic vertebrae. +The lumbar vertebrae are sometimes more numerous than the thoracic. +The epiphyses are very distinct, and do not unite with the centra till +the animal is quite adult. None of the vertebrae are united to form a +sacrum, but the caudal vertebrae have large chevron bones. + +[Illustration FIG. 67. CERVICAL VERTEBRAE OF A CA'ING WHALE +(_Globicephalus melas_) × 1/4. (Camb. Mus.) + + 1. centrum of seventh cervical vertebra. + 2. neural arch of seventh cervical vertebra. + 3. transverse process of atlas. + 4. foramen for exit of first spinal nerve. + 5. transverse process of axis. + 6. fused neural spines of atlas and axis.] + +The skull is peculiarly modified; the bones forming the occipital +segment show a specially strong development, and the cranial cavity is +short, high, and almost spherical. The supra-occipital is very large +and rises up to meet the frontals, thus with the interparietal +completely separating the parietals from one another. + +The frontals are expanded, forming large bony plates, which roof over +the orbits. The zygomatic process of the squamosal is extremely large +and extends forwards to meet the supra-orbital process of the frontal; +the zygomatic process of the jugal is on the contrary very slender. +The face is drawn out into a long rostrum, formed of the maxillae and +premaxillae surrounding the vomer and the mesethmoid cartilage. The +maxillae are specially large, and extend backwards so as to partially +overlap the frontals. The nasals are always small, and the anterior +nares open upwards between the cranium and rostrum. The periotics are +loosely connected with the other bones of the skull and the tympanics +are commonly large and dense. The mandible has hardly any coronoid +process, and the condyles are at its posterior end. + +There are no clavicles, but the scapula and humerus are well +developed. The humerus moves freely in the glenoid cavity, but all the +other articulations of the anterior limb are imperfect; the various +bones have flattened ends, and are connected with one another by +fibrous tissue, which allows of hardly any movement. Frequently the +carpus is imperfectly ossified. + +The number of digits in the manus is generally five, sometimes four, +and when there are four digits it is the third and not the first that +is suppressed. The number of phalanges in the second and third digits +almost always exceeds that which is normal in mammals, and the +phalanges are also remarkable for having epiphyses at both ends. The +pelvis is represented by two small bones which lie suspended +horizontally at some distance below the vertebral column; in some +cases vestiges of the skeleton of the hind limb are attached to them. + +The Cetacea are divided into three suborders. + + +_Suborder_ (1). ARCHAEOCETI. + +The members of this group are extinct; they differ from all living +Cetacea in having the dentition heterodont and in the fact that the +back was probably protected by dermal plates. The skull is elongated +and depressed, and the brain cavity is very small. The temporal fossae +are large, and there is a strong sagittal crest. The nasals and +premaxillae are a good deal larger than they are in living Cetacea, and +the anterior nares are usually far forward. The cervical vertebrae are +not fused with one another, and the lumbar vertebrae are unusually +elongated. + +The limbs are very imperfectly known, but while the humerus is much +longer than in modern Cetaceans, it is nevertheless flattened +distally, indicating that the limb was paddle-like, and that there was +scarcely any free movement between the fore-arm and upper arm. + +The best known genus is _Zeuglodon_, which is found in beds of Eocene +age in various parts of Europe, and in Alabama. + + +_Suborder_ (2). MYSTACOCETI OR BALAENOIDEA. + +These are the Whalebone Whales or True Whales. + +Calcified teeth representing the milk dentition occur in the foetus, +but the teeth are never functional, and always disappear before the +close of foetal life. There is a definite though small olfactory +fossa. The palate is provided with plates of baleen or whalebone. The +skull is symmetrical, and is extremely large in proportion to the +body. The nasals are moderately well developed, and the maxillae do not +overlap the orbital processes of the frontals. The lachrymals are +small and distinct from the jugals. The tympanics are ankylosed to the +periotics, and the rami of the mandible do not meet in a true +symphysis. The ribs articulate only with the transverse processes, and +the capitula are absent or imperfectly developed. Only one pair of +ribs meets the sternum, which is composed of a single piece. + +The group includes among others the Right whale (_Balaena_), +Humpbacked whale (_Megaptera_), and Rorqual (_Balaenoptera_). + + +_Suborder_ (3). ODONTOCETI. + +Teeth always exist after birth and baleen is never present. The teeth +are generally numerous, but are sometimes few and deciduous; the +dentition is homodont (except in _Squalodon_). The dorsal surface of +the skull is somewhat asymmetrical, there is no trace of an olfactory +fossa, the nasals are quite rudimentary, and the hind ends of the +maxillae cover part of the frontals; in all these respects the skull +differs from that of the Mystacoceti. The lachrymal may either be +united to the jugal or may be large and distinct. The tympanic is not +ankylosed to the periotic. The rami of the mandible are nearly +straight and become united in a long symphysis. Some of the ribs have +well developed capitula articulating with the vertebral centra. The +sternum is almost always composed of several pieces as in other +mammals, and several pairs of ribs are connected with it. There are +always five digits to the manus, though the first and fifth are +usually very little developed. + +The suborder includes the Sperm Whale (_Physeter_), Narwhal +(_Monodon_), Dolphin (_Delphinus_), Porpoise (_Phocoena_), and many +other living forms as well as the extinct _Squalodon_ which differs +from the other members of the suborder in its heterodont dentition. + +_Order 4._ UNGULATA. + +This order includes a great and somewhat heterogeneous group of +animals, a large proportion of which are extinct. They all (except +certain extinct forms) agree in having the ends of the digits either +encased in hoofs or provided with broad flat nails. The teeth are +markedly heterodont and diphyodont, and the molars have broad crowns +with tuberculated or ridged surfaces. Clavicles are never present in +the adult except in a few generalised extinct forms such as +_Typotherium_, and it is only recently that vestigial clavicles have +been discovered in the embryo[122]. The scaphoid and lunar are always +distinct. + +The order Ungulata may be subdivided into two main groups, Ungulata +vera and Subungulata. + + +_Section I._ UNGULATA VERA[123]. + +The cervical vertebrae except the atlas are generally opisthocoelous. +The feet are never _plantigrade_[124]. In all the living and the great +majority of the extinct forms the digits do not exceed four, the first +being suppressed. In the carpus the os magnum articulates freely with +the scaphoid, and is separated from the cuneiform by the lunar and +unciform. In the tarsus the cuboid articulates with the astragalus as +well as with the calcaneum, and the proximal surface of the astragalus +is marked by a pulley-like groove. All the bones of the carpus and +tarsus strongly interlock. These characters with regard to the carpus +and tarsus do not hold in _Macrauchenia_ and its allies. The humerus +never has an ent-epicondylar foramen. + +The group is divided into two very distinct suborders:-- + + +_Suborder_ (1). ARTIODACTYLA. + +The Artiodactyla have a number of well marked characters, one of the +most obvious being the fact that many of the most characteristic forms +have large paired outgrowths on the frontal bones. These may be (1) +solid deciduous bony _antlers_, or (2) more or less hollow bony +outgrowths which are sheathed with permanently growing horn. + +The premolar and molar teeth are usually dissimilar, the premolars +being one-lobed and the molars two-lobed; the last lower molar of both +the milk and permanent dentitions is almost always three-lobed. + +The grinding surfaces of the molar teeth have a tendency to assume one +of two forms. In the Pigs and their allies the crowns are bunodont[1], +while in the more highly specialised Ruminants the crowns are +selenodont[125]. The nasals are not expanded posteriorly, and there is +no alisphenoid canal[126]. The thoraco-lumbar vertebrae are always +nineteen. The symphysis of the ischia and pubes is very elongated, and +the femur has no third trochanter. The limbs never have more than four +digits, and are symmetrical about a line drawn between the third and +fourth digits; the digits, on the other hand, are never symmetrical in +themselves. The astragalus has pulley-like surfaces both proximally +and distally, and articulates with the navicular and cuboid by two +nearly equal facets. The calcaneum articulates with the lower end of +the fibula when that bone is fully developed. + +In the Artiodactyla are included the following living groups:-- + +_a._ Suina. Pigs and Hippopotami. + +_b._ Tylopoda. Camels and Llamas. + +_c._ Tragulina. Chevrotains. + +_d._ Ruminantia or Pecora. Deer, giraffes, oxen, sheep and antelopes. + + +_Suborder_ (2). PERISSODACTYLA[127]. + +In this group there are never any bony outgrowths from the frontals. +The grinding teeth form a continuous series, the posterior premolars +resembling the molars in complexity, and the last lower molar +generally has no third lobe. The cervical vertebrae with the +exception of the atlas almost always have markedly opisthocoelous +centra, but in _Macrauchenia_ they are flat. The nasals are expanded +posteriorly, and an alisphenoid canal is present. The thoraco-lumbar +vertebrae are never less than twenty-two in number and are usually +twenty-three. The femur has a third trochanter (except in +_Chalicotherium_). The third digit of the manus and pes is symmetrical +in itself, and larger than the others, and in some cases the other +digits are quite vestigial. The number of the digits of the pes is +always odd. The astragalus is abruptly truncated distally, and the +facet by which it articulates with the cuboid, is much smaller than +that by which it articulates with the navicular. The calcaneum does +not articulate with the fibula, except in _Macrauchenia_. The group +includes many extinct forms, and the living families of the Tapirs, +Horses and Asses, and Rhinoceroses. + + +_Section II._ SUBUNGULATA. + +In this group is placed a heterogeneous collection of animals, the +great majority of which are extinct. There is really no characteristic +which is common to them all, and which serves to distinguish them as a +group from the Ungulata vera. But the most distinctive character +common to the greatest number of them is to be found in the carpus, +whose bones in most cases retain their primitive relation to one +another, the os magnum articulating with the lunar and sometimes just +meeting the cuneiform, but in living forms at any rate not +articulating with the scaphoid. The feet frequently have five +functional digits, and may be plantigrade. The proximal surface of the +astragalus is generally flattened instead of being pulley-like as in +Ungulata vera. + + +_Suborder_ (1). TOXODONTIA. + +This suborder includes some very aberrant extinct South American +ungulates, which have characters recalling the Proboscidea, both +groups of Ungulata vera, and the Rodentia. The limbs are +subplantigrade or digitigrade, and the digits are three, rarely five, +in number, the third being most developed. The carpus resembles that +of the Ungulata vera, in that the bones interlock and the magnum +articulates with the scaphoid. In the tarsus, however, the bones do +not interlock. The astragalus has a pulley-like proximal surface +(except in _Astrapotherium_, in which it is flat), and articulates +only with the navicular, not meeting the cuboid. The calcaneum has a +large facet for articulation with the fibula, as in Artiodactyla. +There is no alisphenoid canal, and the orbit is confluent with the +temporal fossa. Some of the forms (e.g. _Nesodon_) referred to this +group have the typical mammalian series of forty-four teeth, but in +others the canines are undeveloped. In _Toxodon_ all the cheek teeth +have persistent pulps, while in _Nesodon_ and _Astrapotherium_ they +are rooted. A clavicle is sometimes present (_Typotherium_), and the +femur sometimes has a third trochanter (_Typotherium_ and +_Astrapotherium_), sometimes is without one (_Toxodon_). + +The remains of these curious Ungulates have been found in beds of late +Tertiary age in South America. + + +_Suborder_ (2). CONDYLARTHRA[128]. + +This group includes some comparatively small extinct ungulates, which +are best known from the Lower Eocene of Wyoming, though their remains +have also been found in deposits of similar age in France and +Switzerland. Their characters are little specialised, and they show +relationship on the one hand to the Ungulata vera and on the other to +the Hyracoidea. They also have characters allying them to the +Carnivora. They generally have the typical mammalian series of +forty-four teeth, the molars being brachydont and generally bunodont. +The premolars are more simple than the molars. The limbs are +plantigrade, and have five digits with rather pointed ungual +phalanges. The os magnum, as in living Subungulates, articulates with +the lunar, not reaching the scaphoid. The astragalus has an elongated +neck, a pulley-like proximal and a convex distal articular surface, +and does not articulate with the cuboid. The humerus has an +ent-epicondylar foramen, and the femur has a third trochanter. The +best known genus is _Phenacodus_; it is perhaps the most primitive +ungulate whose skeleton is thoroughly well known, and is of special +interest from the fact that it is regarded as the lowest stage in the +evolutionary series of the horse. Its remains are found in the Lower +Eocene of Wyoming. + + +_Suborder_ (3). HYRACOIDEA[129]. + +This group of animals is very isolated, having no very close allies, +either living or extinct. The digits are provided with flat nails, +except the second digit of the pes, which is clawed. Canine teeth are +absent, and the dental formula is usually given as _i_ 1/2, _c_ 0/0, +_pm_ 4/4, _m_ 3/3. The upper incisors are long and curved, and have +persistent pulps as in Rodents; their terminations are, however, +pointed, not chisel-shaped, as in Rodents. The lower incisors have +pectinated edges. The grinding teeth have a pattern much like that in +_Rhinoceros_. In the skull (fig. 83) the postorbital processes of the +frontal and jugal almost or quite meet. The jugal forms part of the +glenoid cavity for articulation with the mandible, and also extends +forwards so as to meet the lachrymal. There is an alisphenoid canal. +There are as many as twenty-one or twenty-two thoracic vertebrae, and +the number of thoraco-lumbar vertebrae reaches twenty-eight or thirty. +There are no clavicles, and the scapula has no acromion; the coracoid +process is, however, well developed. The ulna is complete. In the +manus the second, third and fourth digits are approximately equal in +size, the fifth is smaller, and the first is vestigial. The femur has +a slight ridge representing the third trochanter. The fibula is +complete, but is generally fused with the tibia proximally. There is a +complicated articulation between the tibia and astragalus, which has a +pulley-like proximal surface. In the pes the three middle digits are +well developed, but there is no trace of a hallux, and the fifth digit +is represented only by a vestigial metatarsal. + +The only representatives of the suborder are some small animals +belonging to the genus _Procavia_ (_Hyrax_), which is found in Africa +and Syria; some of the species are by many authors placed in a +distinct genus _Dendrohyrax_. + + +_Suborder_ (4). AMBLYPODA[130]. + +This suborder includes a number of primitive extinct Ungulates, many +of which are of great size. Their most distinguishing characteristics +are afforded by the extremities. In the carpus the bones interlock a +little more than is the case in most Subungulata, and the corner of +the os magnum reaches the scaphoid, while the lunar articulates +partially with both magnum and unciform, instead of only with the +magnum. In the tarsus the cuboid articulates with both the calcaneum +and the astragalus, which is remarkably flat. The manus and pes are +short, nearly or quite plantigrade, and have the full number of +digits. The cranial cavity is singularly small. Canine teeth are +present in both jaws, and the grinding teeth have short crowns, marked +by =V=-shaped ridges. The pelvis is large, the ilia are placed +vertically, and the ischia do not take part in the ventral symphysis. + +The best known animals belonging to this suborder are the +Uintatheriidae (Dinocerata)[131], found in the Upper Eocene of +Wyoming. They are as large as elephants, and are characterised by the +long narrow skull drawn out into three pairs of rounded protuberances, +by the strong occipital crest, and by the very large upper canines. + + +_Suborder_ (5). PROBOSCIDEA. + +This suborder includes the largest of land mammals, the Elephants, and +certain of their extinct allies. The limbs are strong, and are +vertically placed; the proximal segment is the longest, and the manus +and pes are pentedactylate and subplantigrade. The digits are all +enclosed in a common integument, and each is provided with a broad +hoof. The vertebral centra are much flattened and compressed, +especially in the cervical region. The number of thoracic vertebrae is +very great, reaching twenty. The skull (figs. 96 and 97) is extremely +large, this being due to the great development of air cells, which +takes place in nearly all the bones of the adult skull. In the young +skull there are hardly any air cells, and the growth of the cranial +cavity does not by any means keep pace with the growth of the skull in +general. The supra-occipital is very large, and forms a considerable +part of the roof of the skull. The nasals and jugals are short, and +the premaxillae very large. The rami of the mandible meet in a long +symphysis, and the ascending portion is very high. Canine teeth are +absent, and the incisors have the form of ever-growing tusks composed +mainly of dentine; in living forms they are present in the upper jaw +only. The grinding teeth are large, and in living forms have a very +complex structure and mode of succession. In some of the extinct +forms, such as _Mastodon_ and especially _Dinotherium_, the teeth are +much more simple. In every case the teeth have the same general +structure, consisting of a series of ridges of dentine, coated with +enamel. In the more specialised forms the valleys between the ridges +are filled up with cement. The acromion of the scapula has a recurved +process, similar to that often found in rodents. Clavicles are absent. +The radius and ulna are not ankylosed, but are incapable of any +rotatory movement. All the bones of the extremities are very short and +thick; the scaphoid articulates regularly with the trapezoid and the +lunar with the magnum. The ilia are vertically placed, and are very +much expanded; the ischia and pubes are small, and form a short +symphysis. The femur has no third trochanter, and the tibia and fibula +are distinct. The fibula articulates with the calcaneum, and the +astragalus is very flat. + + * * * * * + +Here brief reference may be made to the TILLODONTIA[132], a group of +extinct mammals found in the Eocene beds of both Europe and North +America. They seem to connect together the Ungulata, Rodentia, and +Carnivora. + +The skull resembles that of bears, but the grinding teeth are of +Ungulate type, while the second incisors resemble those of rodents, +and have persistent pulps. The femur has a third trochanter, and the +feet resemble those of bears in being plantigrade and having pointed +ungual phalanges, differing, however, in having the scaphoid and lunar +distinct. + + +_Order 5._ RODENTIA. + +The Rodents form a very large and well-defined group of mammals easily +distinguishable by their peculiar dentition. Canines are absent, and +the incisors are very large and curved, growing from persistent pulps. +They are rectangular in section and are much more thickly coated with +enamel on their anterior face than elsewhere; consequently, as they +wear down they acquire and retain a chisel-shaped (scalpriform) edge. +There is never more than one pair of incisors in the mandible, and +except in the Hares and Rabbits, there is similarly only a single pair +in the upper jaw. These animals are, too, the only rodents which have +well developed deciduous incisors. There is always a long diastema +separating the incisors from the grinding teeth. The grinding teeth, +which are arranged in a continuous series, vary in number from two to +six in the upper jaw, and from two to five in the lower jaw. The +number of premolars is always below the normal, often they are +altogether wanting, but generally they are 1/1. Sometimes the grinding +teeth form roots, sometimes they grow persistently. + +The premaxillae are always large, and the orbits always communicate +freely with the temporal fossae. The condyle of the mandible is +elongated from before backwards, and owing to the absence of a +postglenoid process to the squamosal, a backward and forward motion of +the jaw can take place. The zygomatic arch is complete, but the jugal +is short and only forms the middle of it. The palate is small, being +sometimes, as in the hares, narrowed from before backwards, sometimes +as in the mole-rats (Bathyerginae) narrowed transversely. + +The thoraco-lumbar vertebrae are usually nineteen in number. Clavicles +are generally present, and the acromion of the scapula is commonly +very long. The feet are as a rule plantigrade, and provided with five +clawed digits. + +There are two main groups of Rodentia; the Duplicidentata, or Hares +and Rabbits, which have two pairs of upper incisors, whose enamel +extends round to the posterior surface; and the Simplicidentata, in +which there is only a single pair of upper incisors, whose enamel is +confined to the anterior surface. This group includes all the Rodents +except the Hares and Rabbits. + + +_Order 6._ CARNIVORA. + +The living Carnivora form a natural and well-marked group, but as is +the case with so many other groups of animals, when their extinct +allies are included, it becomes impossible to readily define them. + +The manus and pes never have less than four well-developed digits, and +these are nearly always provided with more or less pointed nails, +generally with definite claws. The hallux and pollex are never +opposable. The dentition is diphyodont and markedly heterodont. The +teeth are always rooted, except in the case of the canines of the +Walrus. The incisors are generally 3/3, and are comparatively small, +while the canines are large, pointed, and slightly recurved. The cheek +teeth are variable, and are generally more or less compressed and +pointed; sometimes their crowns are flattened and tuberculated, but +they are never divided into lobes by deep infoldings of enamel. The +squamosal is drawn out into a postglenoid process, and the mandible +has a large coronoid process. The condyle of the mandible is +transversely elongated, and the glenoid fossa is very deep; in +consequence of this arrangement the mandible can perform an up and +down movement only, any rotatory or back and fore movement being +impossible. The jugal is large, and the zygomatic arch generally +strong, while the orbit and temporal fossa are in most cases +completely confluent. The scapula has a large spine. The clavicle is +never complete and is often absent, this forming an important +distinction between the skeleton of a Carnivore and of any Insectivore +except _Potamogale_. The humerus often has an ent-epicondylar foramen, +and the radius and ulna, tibia and fibula are always separate. The +manus is often capable of the movements of pronation and supination, +and the scaphoid, lunar and centrale are in living forms always united +together. + +The order Carnivora includes three suborders. + + +_Suborder_ (1). CREODONTA[133]. + +This suborder contains a number of extinct Carnivora, which present +very generalised characters. + +The cranial cavity is very small; and the fourth upper premolar and +first lower molar are not differentiated as carnassial teeth[134], as +they are in modern Carnivora. The Creodonta also differ from modern +Carnivora in the fact that the scaphoid and lunar are usually +separate, and that the femur has a third trochanter. The feet are +plantigrade. + +They resemble the Condylarthra, another very generalised group, in +having an ent-epicondylar foramen. + +They occurred throughout the Tertiary period in both Europe and North +America, and have also been found in India. One of the best known +genera is _Hyaenodon_. + + +_Suborder_ (2). CARNIVORA VERA or FISSIPEDIA. + +The skeleton is mainly adapted for a terrestrial mode of life, and the +hind limbs have the normal mammalian position. In almost every case +the number of incisors is 3/3. Each jaw always has one specially +modified _carnassial_ or sectorial tooth which bites like a scissors +blade against a corresponding tooth in the other jaw. In front of it +the teeth are always more or less pointed, while behind it they are +more or less broadened and tuberculated. In the manus the first digit, +and in the pes the first and fifth digits are never longer than the +rest, and the digits of both limbs are almost invariably clawed. Some +forms are plantigrade, some digitigrade, some subplantigrade. The +group includes all the ordinary terrestrial Carnivora, and is divided +into three sections:-- + +Æluroidea[135], including the cats, civets, hyaenas, and allied forms. + +Cynoidea, including the dog tribe. + +Arctoidea, including the bears, raccoons, weasels, and allied forms. + + +_Suborder_ (3). PINNIPEDIA[136]. + +In this suborder the limbs are greatly modified and adapted for a more +or less purely aquatic life, the proximal and middle segments of the +limbs are shortened, while the distal segment, especially in the leg, +is much elongated and expanded. There are always five well-developed +digits to each limb, and in the pes the first and fifth digits are +generally larger than the others. The digits generally bear straight +nails instead of claws, but even nails are sometimes absent. There is +no carnassial tooth, and the teeth in other ways differ considerably +from those of Carnivora vera. The incisors are always fewer than 3/3; +while the cheek teeth generally consist of four premolars and one +molar, all of very uniform character, being compressed with conical +crowns, and never more than two roots. + +The suborder includes three families--Otariidae (Eared Seals), +Trichechidae (Walrus), and Phocidae (Seals). + + +_Order 7._ INSECTIVORA[137]. + +This order contains a large number of small generally terrestrial +mammals. The limbs are plantigrade or subplantigrade, and are +generally pentedactylate. All the digits are armed with claws, and the +pollex and hallux are not opposable. The teeth are diphyodont, +heterodont, and rooted. The cheek teeth have tuberculated crowns, and +there are never less than two pairs of incisors in the mandible; often +the incisors, canines, and premolars are not clearly differentiated +from one another, and special carnassial teeth are never found. The +cranial cavity is small, and the facial part of the skull is generally +much developed; often the zygomatic arch is incomplete. Clavicles are +well developed (except in _Potamogale_), and the humerus generally has +an ent-epicondylar foramen. The femur frequently has a ridge +representing the third trochanter. There are two suborders: + + +_Suborder_ (1). DERMOPTERA. + +This suborder includes only a very aberrant arboreal genus +_Galeopithecus_, remarkable for its greatly elongated limb bones, and +peculiar dentition. The incisors of the lower jaw are deeply +pectinated or divided by several vertical fissures, the canines and +outer upper incisors have two roots. Ossified inter centra occur in +the thoraco-lumbar region of the vertebral column. + + +_Suborder_ (2). INSECTIVORA VERA. + +This suborder includes all the ordinary Insectivora, such as moles, +shrews and hedgehogs. The upper and lower incisors are conical, not +pectinated. + + +_Order 8._ CHIROPTERA[138]. + +This order is perhaps the best marked and most easily defined of all +the orders of mammals. The anterior limbs form true wings and the +whole skeleton is modified in relation to flight. + +The anterior limbs are vastly larger than the posterior; for all the +bones except the carpals are much elongated, and this applies +specially to the phalanges of all the digits except the pollex. + +The pollex is clawed and so is sometimes the second digit; the other +digits of the manus are without nails or claws. The teeth are +divisible into the four usual types and the series never exceeds _i_ +2/3 _c_ 1/1 _pm_ 3/3 _m_ 3/3 × 2, total 38. The milk teeth are quite +unlike the permanent teeth. The orbit is not divided by bone from the +temporal fossa. The vertebral column is short, and in old animals the +trunk vertebrae have a tendency to become partially fused together. +The cervical vertebrae are remarkably wide, and the development of +spinous processes is everywhere slight. The presternum has a prominent +keel for the attachment of the pectoral muscles. The clavicles are +very long and strong, and the scapula has a long spine and coracoid +process. The ulna is vestigial, consisting only of a proximal end +ankylosed to the radius. All the carpals of the proximal row--the +scaphoid, lunar and cuneiform--are united, forming a single bone. The +pelvis is very weak and narrow, and only in the Rhinolophidae do the +pubes meet in a symphysis. The anterior caudal vertebrae are +frequently united to the ischia. The fibula is generally vestigial, +and the knee joint is directed backwards instead of forwards. The pes +has five slender clawed digits, and the calcaneum is often drawn out +into a spur which helps to support the membrane connecting the hind +limbs with the tail. + +There are two suborders of Chiroptera: + +1. The MEGACHIROPTERA or Flying foxes, which almost always have smooth +crowns to the molar teeth, and the second digit of the manus clawed. + +2. The MICROCHIROPTERA including all the ordinary bats which have +cusped molar teeth, and the second digit of the manus clawless. + + +_Order 9._ PRIMATES. + +The dentition is diphyodont and heterodont, the incisors generally +number 2/2, and the molars, except in the Hapalidae (Marmosets), are +3/3. The cheek teeth are adapted for grinding, and the molars are more +complex than the premolars. A process from the jugal meets the +postorbital process of the frontal completing the postorbital bar. + +The clavicle is well developed, and the radius and ulna are never +united. The scaphoid and lunar of the carpus, and commonly also the +centrale, remain distinct from one another. As a rule both manus and +pes have five digits, but the pollex may be vestigial. The pollex is +opposable to the other digits, and so is the hallux except in Man; the +digits are almost always provided with flat nails. The humerus has no +ent-epicondylar foramen and the femur has no third trochanter. + +The order Primates is divisible into two suborders: + + +_Suborder_ (1). LEMUROIDEA. + +The skull has the orbit communicating freely with the temporal fossa +beneath the postorbital bar (except in _Tarsius_). The lachrymal +foramen is external to the margin of the orbit. Both pollex and hallux +are well developed. In the pes the second digit is terminated by a +long pointed claw, and so is also the third in _Tarsius_. The lumbar +region of the vertebral column is long, sometimes including as many as +nine vertebrae. Besides the Lemurs the group includes the aberrant +_Tarsius_ and _Chiromys_. + + +_Suborder_ (2). ANTHROPOIDEA. + +The skull has the orbit almost completely shut off from the temporal +fossa, and the lachrymal foramen is situated within the orbit. The +pollex is sometimes vestigial or absent. The second digit of the pes +has a flattened nail except in the Hapalidae, in which all the digits +of the pes except the hallux are clawed. + +The Anthropoidea are divided into five families: + +1. Hapalidae or Marmosets. + +2. Cebidae or American Monkeys. + +3. Cercopithecidae or Old World Monkeys. + +4. Simiidae or Anthropoid Apes. + +5. Hominidae or Men. + + +FOOTNOTES: + +[108] This is Gadow's view; according to Huxley the quadrate forms the +malleus; according to Baur it forms the zygomatic process of the +squamosal, and according to Broom the interarticular mandibular +cartilage. + +[109] According to Leche, _Morphol. Jahrb._ XIX. p. 502, the molar +teeth belong morphologically to the first series, i.e. they are milk +teeth without vertical successors. + +[110] The researches of Bateson, _P.Z.S._ 1892, p. 102, have shown +that cases of individual variation in the number of teeth are common. + +[111] Baur, however, suggests (_Anat. Anz._ vol. IV. 1889), that a +tibial sesamoid found in _Procavia_, many rodents, edentates and +_Ornithorhynchus_ is a vestigial tibiale, and that the astragalus is +the intermedium. + +[112] This perforation of the acetabulum in _Echidna_ is a secondary +character occurring late in development, and consequently is not of +phylogenetic importance. + +[113] See R. Owen, "Monograph of the Fossil Mammalia of the Mesozoic +Formation," _Pal. Soc. Mon._ 1871. + +H.F. Osborn, "Structure and Affinities of Mesozoic Mammals," _J. of +Philad. Acad._ 1888, vol. IX. + +O.C. Marsh, "Jurassic Mammals," _Amer. J. Sci._ 1878 _et seq._ + +[114] See Oldfield Thomas, _Brit. Mus. Cat. of Marsupialia and +Monotremata_ (1888). + +[115] W. Kükenthal, _Anat. Anz._ VI. p. 364, 1891. C. Röse, _Anat. +Anz._ VII. p. 639. + +[116] These bones however have no connection with the marsupium, being +nearly equally developed in both male and female. They are simply +sesamoid bones forming ossifications in the inner tendon of the +external oblique muscle, and are developed as supports for the +abdominal wall. Very similar structures have been independently +developed in various Amphibians, Reptiles and monodelphian Mammals. +See W. Leche, _Biol. Fören._ III. p. 120. + +[117] See H. Gadow, _P.Z.S._ 1892, p. 361. + +[118] See W.H. Flower, "On the Mutual Affinities of the Animals +composing the order Edentata," _P.Z.S._ 1882, p. 358. For the fossil +Edentates of N. America see E. Cope, _Amer. Natural._ 1889; for those +of S. America see various papers by F. Ameghino, H. Burmeister and R. +Owen. Also T.H. Huxley, "On the Osteology of Glyptodon," _Phil. +Trans._ 1865. + +[119] See J.F. Brandt, _Symbolae Sirenologicae_, St Petersburg, 1846, +1861, 1868. + +[120] Epiphyses are fully developed in _Halitherium_, and traces occur +in _Manatus_. + +[121] See P.J. van Beneden and P. Gervais, _Ostéographie des Cétacés_, +1869-80. + +[122] H. Wincza, _Morphol. Jahrb._ XVI., p. 647. + +[123] See M. Pavlow, "Études sur l'histoire paléontologique des +Ongulés." _Bull. Soc. Moscou_, 1887--1890. + +[124] In a _plantigrade_ animal the whole of the foot is placed on the +ground in walking. A _digitigrade_ animal places only its toes on the +ground. An intermediate condition is distinguished by the term +_subplantigrade_. + +[125] See p. 345. + +[126] See p. 401. + +[127] See E.D. Cope, "The Perissodactyla," _Amer. Natural._, 1887. + +[128] See E.D. Cope, "The Condylarthra," _Amer. Natural._, 1884, and +"Synopsis of the Vertebrates of the Puerco series," _Tr. Amer. Phil. +Soc._, 1888. O.C. Marsh, "A new order of extinct Eocene Mammals +(Mesodactyla)," _Amer. J. Sci._, 1892. + +[129] See O. Thomas, "On the species of Hyracoidea," _P.Z.S._, 1892, +p. 50. + +[130] See E.D. Cope, "The Amblypoda," _Amer. Natural._, 1884 and 1885. + +[131] See O.C. Marsh, "The Dinocerata," _U.S. Geol. Survey_, 1884, +vol. X. + +[132] See O.C. Marsh, _Amer. J. Sci._, 1875 and 1876. + +[133] E.D. Cope, "The Creodonta," _Amer. Natural._, 1884. W.B. Scott, +"Revision of the N. American Creodonta," _P. Ac. Philad._, 1892. + +[134] See next paragraph. + +[135] St G. Mivart, _The Cat_, London, 1881. + +[136] St G. Mivart, _P.Z.S._, 1885. + +[137] St G. Mivart, "On the Osteology of Insectivora," _J. Anat. +Physiol. norm. path._, 1867 and 1868, and _P.Z.S._, 1871. G.E. Dobson, +_Monograph of the Insectivora_, London, 1882--90. + +[138] See G.E. Dobson, _Brit. Mus. Catalogue of Chiroptera_, 1878. See +also other papers by the same author and by Oldfield Thomas. + + + + +CHAPTER XXI. + +THE SKELETON OF THE DOG[139] (_Canis familiaris_). + + +I. EXOSKELETON. + +The exoskeleton of the dog includes three sets of structures: 1. +hairs, 2. claws, 3. teeth. =Hairs= and =claws= are epidermal +exoskeletal structures, while =teeth= are partly of dermal, and partly +of epidermal origin. + +1. =Hairs= are delicate epidermal structures which grow imbedded in +little pits or follicles in the dermis. Specially large hairs forming +the =vibrissae= or =whiskers= grow attached to the upper lip. + +2. =Claws= are horny epidermal sheaths, one of which fits on to the +pointed distal phalanx of each digit. They are sharply curved +structures, and being in the dog non-retractile, their points are +commonly much blunted by friction with the ground. The claws of the +pollex, and of the hallux when it is present, however do not meet the +ground, and therefore remain comparatively sharp. + +3. =Teeth=[140]. Although as regards their mode of origin, teeth are +purely exoskeletal or tegumentary structures, they become so +intimately connected with the skull that they appear to belong to the +endoskeleton. + +Each tooth, as has been already described, consists of three distinct +tissues, dentine and cement of dermal origin, and enamel of epidermal +origin. + +[Illustration FIG. 68. DENTITION OF A DOG (_Canis familiaris_) × 1/2. +(Camb. Mus.) + + _i_ 2. second incisor. + _c._ canine. + _pm_ 1, _pm_ 4. first and fourth premolars. + _m_ 1. first molar.] + +The teeth of the dog (fig. 68) form a regular series arranged along +the margins of both upper and lower jaws, and imbedded in pits or +=alveoli= of the maxillae, premaxillae, and mandibles. They are all +fixed in the bone by tapering roots, and none of them grow from +persistent pulps. + +They are divisible into four distinct groups, the =incisors=, +=canines=, =premolars= and =molars=. There are three incisors, one +canine and four premolars on each side of each jaw. But while there +are three molars on each side of the lower jaw, the last is wanting in +the upper jaw. The dentition of the dog may then be represented by the +formula + + _i_ 3/3 _c_ 1/1 _pm_ 4/4 _m_ 2/3 × 2 = 42. + +In each jaw there is one large specially modified tooth called the +=carnassial=, the teeth in front of this are more or less pointed and +compressed, while those behind it are more or less flattened and +tuberculated. + +=Teeth of the upper jaw.= + +The first and second =incisors= are small teeth with long conical +roots and somewhat chisel-shaped crowns. Surrounding the base of the +crown there is a rather prominent ridge, terminated laterally by a +pair of small cusps. This ridge, the =cingulum=, serves to protect the +edge of the gums from injury by the hard parts of food. The third +incisor is a good deal like the others but larger, and has the +cingulum well developed though not terminated by lateral cusps. All +the incisors are borne by the premaxillae, the remaining teeth by the +maxillae. + +The =canine= is a large pointed tooth, slightly recurved and with a +long tapering root. + +The =premolars= are four in number, and in all the cingulum is fairly +well seen. The first is a very small tooth with a single tapering +root, the second and third are larger and have two roots, while the +fourth, the =carnassial=, is much the largest and has three roots. +Each of the second, third and fourth premolars has a stout blade, the +middle portion of which is drawn out into a prominent cone; the +posterior part of the fourth premolar forms a compressed ridge, and +at the antero-internal edge of the tooth there is a small inner +tubercle. + +The two =molar= teeth are of very unequal size. The first, which has +two anterior roots and one posterior, is wider than it is long, its +outer portion being produced into two prominent cusps, while its inner +portion is depressed. The second molar is a small tooth resembling the +first in its general appearance, but with much smaller outer cusps. + +=Teeth of the lower jaw.= + +The three =incisors= of the lower jaw have much the same character as +the first two of the upper jaw; while the =canine= is identical in +character with that of the upper jaw. + +The four =premolars= gradually increase in size from the first to the +last, but none are very large. The first premolar is a single-rooted +tooth resembling that of the upper jaw; the second, third and fourth +are two-rooted, like the second and third of the upper jaw, which they +closely resemble in other respects. + +The first =molar= forms the =carnassial= (fig. 84, V), and with the +exception of the canine, is much the largest tooth of the lower jaw; +it is a two-rooted tooth, with a long compressed bilobed blade, and a +posterior tuberculated talon or heel. The second molar is much +smaller, though likewise two-rooted, while the third molar is very +small and has only a single root. All the teeth except the molars are +preceded in the young animal by temporary =milk teeth=. These milk +teeth, though smaller, are very similar to the permanent teeth by +which they are ultimately replaced. + + +II. ENDOSKELETON. + +1. THE AXIAL SKELETON. + +This includes the vertebral column, the skull, and the ribs and +sternum. + + +A. THE VERTEBRAL COLUMN. + +This consists of a series of about forty vertebrae arranged in +succession so that their centra form a continuous rod, and their +neural arches a continuous tube, surrounding a cavity, the =neural +canal=. + +The vertebrae may be readily divided into five groups:-- + +1. The =cervical= or neck vertebrae. + +2. The =thoracic= or chest vertebrae which bear ribs. + +3. The =lumbar= vertebrae which are large and ribless. + +4. The =sacral= vertebrae which are fused with one another and united +with the pelvis. + +5. The =caudal= or tail vertebrae which are small. + +Except in the sacral region the vertebrae are movably articulated to +one another, while their centra are separated from one another by +cartilaginous =intervertebral discs=. + + +GENERAL CHARACTERS OF A VERTEBRA. + +Take as a type the =fourth lumbar vertebra=. It may be compared to a +short tube whose inner surface is smooth and regular, and whose outer +surface is thickened and drawn out in a variety of ways. The basal +part of the vertebra is the =centrum= or body which forms the +thickened floor of the neural canal. Its two ends are slightly convex +and are formed by the =epiphyses=, two thin plates of bone which are +at first altogether distinct from the main part of the centrum, but +fuse with it as the animal grows older; its sides are drawn out into a +pair of strong =transverse processes=, which project forwards, +outwards, and slightly downwards. The =neural arch= forms the sides +and roof of the neural canal, and at each end just above the centrum +bears a pair of =intervertebral notches= for the passage of the spinal +nerves, the posterior notches being considerably deeper than the +anterior. The neural arch is drawn out into a series of processes. +Arising from the centre of the dorsal surface is a prominent median +=neural spine= or =spinous process=, which projects upwards and +slightly forwards; its anterior edge is vertical, while its posterior +edge slopes gradually. At the two ends of the neural arch arise the +two pairs of =zygapophyses= or articulating surfaces, which +interlock with those of the adjacent vertebrae. The anterior or +=prezygapophyses= look inwards, and are large and concave; they are +borne upon a pair of large blunt outgrowths of the neural arch, the +=metapophyses=. The posterior or =postzygapophyses= are slightly +convex and look outwards and downwards; they are borne upon backwardly +projecting outgrowths of the neural arch. Lastly there are a pair of +minute projections arising from the posterior end of the neural arch, +below the postzygapophyses. These are the =anapophyses=. In young +individuals the development of all the processes of the various +vertebrae is less marked, and the epiphyses are obviously distinct. + +[Illustration FIG. 69. A, ATLAS AND B, AXIS VERTEBRA OF A DOG (_Canis +familiaris_) (after VON ZITTEL). + + 1. transverse process of atlas. + 2. vertebrarterial canal. + 3. foramen for exit of spinal nerve. + 4. neural spine. + 5. odontoid process. + 6. anterior articulating surface of centrum. + 7. centrum. + 8. transverse process of axis. + 9. postzygapophysis.] + + +THE CERVICAL VERTEBRAE. + +These are seven in number, as in almost all mammals. They are +characterised by the fact that they have small ribs fused with them, +forming transverse processes perforated by canals through which the +vertebral arteries run. + +The first, or =atlas= vertebra (fig. 69, A), differs much from all the +others; it is drawn out into a pair of wide wing-like transverse +processes (fig. 69, A, 1), and forms a ring surrounding a large +cavity. This cavity is during life divided into two parts by a +transverse ligament; the upper cavity is the true neural canal, while +the lower lodges the =odontoid process= of the second vertebra, which +is the detached centrum of the atlas. The neural arch is broad and +regular; it has no spinous process, and is perforated in front by a +pair of foramina for the passage of the first spinal nerves. The +mid-ventral portion of the atlas is rather thick, and bears a minute +backwardly-projecting hypapophysis. The bases of the broad transverse +processes are perforated by the =vertebrarterial canals= (fig. 69, A, +2). The atlas bears at each end a pair of large articulating surfaces; +those at the anterior end articulate with the condyles of the skull, +and are very deeply concave; those at the posterior end for +articulation with the axis, are nearly as large, but are flattened. +The atlas ossifies from three centres, one forming the mid-ventral +portion, the others the two halves of the remainder. + +The second, or =axis= vertebra (fig. 69, B), also differs much from +the other cervicals. The long and broad centrum has a very flat dorsal +surface, and is produced in front into the conical =odontoid process= +(fig. 69, B, 5), and bears a pair of very large convex outwardly +directed surfaces for articulation with the atlas. At its posterior +end it is drawn out into a pair of small backwardly-directed spines, +the transverse processes; these are perforated at their bases by the +vertebrarterial canals. The neural arch is deeply notched in front and +behind for the passage of the spinal nerves, and is drawn out above +into a very long compressed neural spine (fig. 69, B, 4), which +projects a long way forwards, and behind becomes bifid and thickened, +bearing a pair of flat downwardly directed postzygapophyses. In the +young animal the odontoid process is readily seen to ossify from a +centre anterior to that forming the anterior epiphysis of the axis. + +The remaining five cervical vertebrae, the third to the seventh +inclusive, have rather flattened wide centra, obliquely truncated at +either end. The neural spine progressively increases in size as the +vertebrae are followed back. The transverse processes vary +considerably; those of the third are divided into a thicker +backwardly-, and a more slender forwardly-projecting portion; those of +the fourth and fifth mainly extend downwards, and that of the sixth is +divided into a horizontal portion and a downwardly-projecting +=inferior lamella=. All the cervical vertebrae except the seventh have +the bases of the transverse processes perforated by the +vertebrarterial canals. The prezygapophyses in each case look upwards +and slightly inwards, while the postzygapophyses look downwards and +slightly outwards. + + +THE THORACIC VERTEBRAE. + +The =thoracic vertebrae= are twelve or thirteen in number, and all +bear movably articulated ribs. As a group they are characterised by +their comparative shortness, and in the case of the first eight or +nine by the great length of the backwardly-sloping neural spine. The +posterior thoracic vertebrae approach in character the succeeding +lumbar vertebrae. + +As type of the anterior thoracic vertebrae, take any one between the +second and sixth inclusive. The centrum is short, and has its +terminations vertically truncated. At the top of the centrum, at both +anterior and posterior ends on each side, is a demi-facet (fig. 70, A, +4), which, together with that on the adjacent vertebra, forms an +articulating surface for the capitulum of the rib. The neural arch is +small and deeply notched behind for the passage of the spinal nerve. +It is drawn out above into a very long neural spine (fig. 70, A, 1), +whose base extends back over the succeeding vertebra and bears the +downwardly-directed postzygapophyses (fig. 70, A, 6). The summit of +the neural arch is deeply notched in front, and on each side of the +notch are the prezygapophyses, which look almost vertically upwards. +The transverse processes are short and blunt, and are flattened below +(fig. 70, A, 3) for the articulation of the tubercula of the ribs. + +[Illustration FIG. 70. A, SECOND THORACIC, AND B, SECOND LUMBAR +VERTEBRA OF A DOG (_Canis familiaris_) SEEN FROM THE RIGHT SIDE (after +VON ZITTEL). + + 1. neural spine. + 2. centrum. + 3. transverse process bearing in A the facet for articulation with + the tuberculum of the rib. + 4. facet for articulation with the capitulum of the rib. + 5. metapophysis. + 6. postzygapophysis.] + +The posterior three or four thoracic vertebrae differ much from the +others. The centra are longer, the neural spines short and not +directed backwards, the articular facets for the heads of the ribs are +confined to the anterior end of the centrum of each vertebra, not +overlapping on to the preceding vertebra. The transverse processes are +small and irregular, and metapophyses and anapophyses are developed. +The prezygapophyses also look more inwards, and the postzygapophyses +more outwards than in the more typical thoracic vertebrae. + + +THE LUMBAR VERTEBRAE. + +The =lumbar vertebrae= are seven in number, and their general +characteristics have been already described. As a group they are +characterised by their large size, and the great development of the +transverse processes, metapophyses and neural spines. + + +THE SACRAL VERTEBRAE. + +Three vertebrae are commonly found fused together, forming the +=sacrum=; the divisions between the three being indicated by the +foramina for the exit of the spinal nerves. + +Of these three vertebrae, the first is much the largest, and is firmly +united to the ilium on each side by a structure formed by the +transverse processes and expanded ribs. In the adult this structure +forms one continuous mass, but in the young animal a ventral portion +formed by the rib is clearly distinguishable from a dorsal portion +formed by the transverse process. All three have low neural spines. +The anterior sacral vertebra bears a large pair of prezygapophyses, +while the posterior one bears a small pair of postzygapophyses. + + +THE CAUDAL VERTEBRAE. + +The =caudal vertebrae= are about nineteen in number. The earlier ones +have well-developed neural arches, transverse processes, and +zygapophyses, but as the vertebrae are followed back they gradually +lose all their processes, and the neural arch as well, becoming at +about the thirteenth from the end reduced to simple cylindrical +centra. + + +B. THE SKULL. + +The =skull= consists of the following three parts: (_a_) the cranium, +with which are included the skeletal supports of the various special +sense organs, and the bones of the face and upper jaw; (_b_) the lower +jaw or mandible, which is movably articulated to the cranium, and +(_c_) the hyoid. + + +(_a_) THE CRANIUM. + +The cranium is a compact bony box, forming the anterior expanded +portion of the axial skeleton. It has a longitudinal axis, the +=craniofacial= axis around which the various parts are arranged, and +this axis is a direct continuation of that of the vertebral column. +Similarly the cavity of the cranium is a direct continuation of the +spinal canal. The posterior part of the craniofacial axis, which has +relations only with the cranium, is called the =basicranial axis=. + +In the dog as in the other types previously described, the skull in +its earliest stages is cartilaginous, containing no bone. In the +adult, however, the cartilage is to a great extent replaced by bone, +and in addition to this cartilage bone, membrane bone is largely +developed, and intimately united with the cartilage bone to form one +complete whole. + +In the description of the dog's skeleton, as in those of the previous +types, the names of the membrane bones are printed in italics, while +those of the cartilage bones are printed in thick type. + +Most of the numerous foramina perforating the skull walls will be +described after the bones have been dealt with. + +For purposes of description the cranium may be further subdivided +into:-- + +1. The cranium proper or brain case. + +2. The sense capsules. + +3. The upper jaw. + + +1. THE CRANIUM PROPER OR BRAIN CASE. + +Taking the membrane and cartilage bones together, they are seen to be +more or less arranged in three segments, which however must not be +regarded as homologous with the segments forming the vertebral column. + +[Illustration FIG. 71. DIAGRAM OF THE RELATIONS OF THE PRINCIPAL BONES +IN THE MAMMALIAN SKULL (modified after FLOWER). + +Cartilage is dotted. Cartilage bones are marked by dots and dashes, +membrane bones are left white. + + 1. basi-occipital. + 2. exoccipital. + 3. supra-occipital. + 4. basisphenoid. + 5. alisphenoid. + 6. parietal. + 7. presphenoid. + 8. orbitosphenoid. + 9. frontal. + 10. periotic, immediately below which is the tympanic. + 11. lachrymal. + 12. ethmo-turbinal. + 13. maxillo-turbinal. + 14. nasal. + 15. mesethmoid. + 16. vomer. + 17. pterygoid. + 18. palatine. + 19. maxillae. + 20. premaxillae. + 21. squamosal. + 22. mandible. + 23. tympano-hyal. + 24. stylo-hyal. + 25. epi-hyal. + 26. basi-hyal. Between this and the epi-hyal is the cerato-hyal. + 27. thyro-hyal. + 28. jugal. + +Nerve exits are indicated by Roman numerals.] + +The =occipital segment= is the most posterior of the three, and +consists of four cartilage bones, which in the adult are commonly +completely fused together. They surround the great =foramen magnum= +(fig. 75, 2) through which the brain and spinal cord communicate. +Forming the lower margin of the foramen magnum is a large flat +unpaired bone, the =basi-occipital= (fig. 75, 5). Above this on each +side are the =exoccipitals=, whose sides are drawn out into a pair of +downwardly-directed =paroccipital processes=, which are applied to the +tympanic bullae[141]. The inner side of each exoccipital is converted +into the large rounded =occipital condyle= (fig. 72, 13) by which the +skull articulates with the atlas vertebra. The dorsal boundary of the +foramen magnum is formed by a large unpaired flat bone, the +=supra-occipital= (figs. 72 and 75, 1), which is continuous with a +small bone, the _interparietal_, prolonged forwards between the +parietal bones of the next segment. + +In old animals the interparietal forms the hind part of a prominent +ridge running along the mid-dorsal surface of the skull and called the +=sagittal crest=, while the junction line of the occipital and +parietal segments forms a prominent =occipital crest=. + +The plane in which the bones of the occipital segment lie is called +the occipital plane; the angle that it makes with the basicranial axis +varies much in different mammals. + +The =parietal segment= consists of both cartilage and membrane bones. +It is formed of five bones, which are in contact with those of the +occipital segment on the dorsal and ventral surfaces, while laterally +they are separated by the interposition of the auditory bones, and to +some extent of the squamosal. The =basisphenoid= (fig. 75, 6), an +unpaired bone forming the ventral member of this segment, is the +direct continuation of the basi-occipital. It tapers anteriorly, but +is rather deep vertically, its upper or dorsal surface bearing a +depression, the =sella turcica=, which lodges the pituitary body of +the brain. From the sides of the basisphenoid arise the =alisphenoids= +(fig. 75, 11) a pair of bones of irregular shape generally described +as wing-like; each gives off from its lower surface a =pterygoid +plate=, which is united in front with the palatine, and below with the +pterygoid. The alisphenoids are united above with a pair of large +nearly square bones, the _parietals_ (fig. 73, 2), which meet one +another in the mid-dorsal line. The line of junction is frequently +drawn out into a strong ridge, which forms the anterior part of the +=sagittal crest=. + +[Illustration FIG. 72. VERTICAL LONGITUDINAL SECTION TAKEN A LITTLE TO +THE LEFT OF THE MIDDLE LINE THROUGH THE SKULL OF A DOG (_Canis +familiaris_) × 3/5. (Camb. Mus.) + + 1. supra-occipital. + 2. interparietal. + 3. parietal. + 4. frontal. + 5. cribriform plate. + 6. nasal. + 7. mesethmoid. + 8. maxillae. + 9. vomer. + 10. ethmo-turbinal. + 11. maxillo-turbinal. + 12. premaxillae. + 13. occipital condyle. + 14. basi-occipital. + 15. tympanic bulla. + 16. basisphenoid. + 17. pterygoid. + 18. palatine. + 19. alisphenoid. + 20. internal auditory meatus. + 21. tentorium. + 22. foramen lacerum posterius. + 23. floccular fossa. + 24. coronoid process. + 25. condyle. + 26. angle. + 27. mandibular symphysis. + 28. inferior dental foramen. + 29. stylo-hyal. + 30. epi-hyal. + 31. cerato-hyal. + 32. basi-hyal. + 33. thyro-hyal. + XII. condylar foramen.] + +The =frontal segment=, which surrounds the anterior part of the brain, +is closely connected along almost its whole posterior border with the +parietal segment. + +Its base is formed by the =presphenoid= (fig. 75, 12), a very deep +unpaired bone, narrow and compressed ventrally, and with an irregular +dorsal surface. The presphenoid is continuous with a second pair of +wing-like bones, the =orbitosphenoids=. Each orbitosphenoid meets the +alisphenoid behind, but the relations of the parts in this region are +somewhat obscured by a number of large foramina piercing the bones, +and also by an irregular vacuity, the =foramen lacerum anterius= or +=sphenoidal fissure=, which lies between the orbitosphenoid and +alisphenoid, separating the lateral parts of the parietal and frontal +segments, in the same way as the space occupied by the auditory bones +separates the lateral parts of the occipital and parietal segments. +The orbitosphenoids pass obliquely forwards and upwards, and are +united above with a second pair of large membrane bones, the +_frontals_ (fig. 73, 3). The outer side of each frontal is drawn out +into a rather prominent rounded =postorbital process= (fig. 73, 10), +from which a ridge converges backwards to meet the sagittal crest. The +anterior part of the frontal is produced to form the long nasal +process, which is wedged in between the nasal and maxillae. + +[Illustration FIG. 73. DORSAL VIEW OF THE CRANIUM OF A DOG (_Canis +familiaris_) × 2/3. + + 1. supra-occipital. + 2. parietal. + 3. frontal. + 4. nasal. + 5. maxillae (facial portion). + 6. premaxillae. + 7. squamosal. + 8. jugal. + 10. postorbital process of frontal. + 11. infra-orbital foramen. + 12. anterior palatine foramen. + 13. lachrymal foramen. + _i_ 1. first incisor. + _c._ canine. + _pm_ 4. fourth premolar.] + +The cranial cavity is continuous in front with the =nasal= or +=olfactory cavities=, but the passage is partially closed by a screen +of bone, the =cribriform plate= (fig. 72, 5), which is placed +obliquely across the anterior end of the cranial cavity, and is +perforated by a number of holes through which the olfactory nerves +pass. The plane of the cribriform plate is called the =ethmoidal +plane=, and as was the case also with the occipital plane, the angle +that it makes with the basicranial axis varies much in different +mammals, and is of importance. The =olfactory fossa= in which lie the +olfactory lobes of the brain, is partially separated from the +=cerebral fossa=, or cavity occupied by the cerebral hemispheres, by +ridges on the orbitosphenoids and frontals. The presphenoid is +connected in front with a vertical plate formed partly of bone, partly +of unossified cartilage; this plate, the =mesethmoid= (fig. 72, 7), +separates the two olfactory cavities which lodge the olfactory organs. +Its anterior end always remains unossified, and forms the septal +cartilage of the nose. + +The brain case may then, to use the words of Sir W.H. Flower, be +described as a tube dilated in the middle and composed of three bony +rings or segments, with an aperture at each end, and a fissure or +space at the sides between each of them. + + +2. THE SENSE CAPSULES. + +Each of the three special sense organs, of hearing, of sight, and of +smell, is in the embryo provided with a cartilaginous or membranous +protecting capsule; and two of these, the auditory and olfactory +capsules, become afterwards more or less ossified, and intimately +related to the cranium proper. + + +(1) =Bones in relation to the Auditory capsules.= + +These bones lie on each side wedged into the vacuity between the +lateral parts of the occipital and parietal segments; they are three +in number, the =periotic=, the _tympanic_ and the _squamosal_. + +The =periotic= is the most important of them, as it replaces the +cartilaginous auditory capsule of the embryo, and encloses the +essential organ of hearing. It commences to ossify from three centres +corresponding to the pro-otic, epi-otic and opisthotic of lower +skulls, such as those of the Turtle and Crocodile. + +These ossifications however very early combine to form a single bone, +the =periotic=, which nevertheless consists of two portions, the +=petrous= and the =mastoid=, differing considerably from one another. + +[Illustration FIG. 74. DIAGRAM OF THE MAMMALIAN TYMPANIC CAVITY AND +ASSOCIATED PARTS (modified from LLOYD MORGAN). + + 1. external auditory meatus. + 2. tympanic membrane. + 3. malleus. + 4. incus. + 5. lenticular. + 6. stapes. + 7. fenestra ovalis. + 8. fenestra rotunda. + 9. Eustachian tube. + 10. cavity occupied by the cochlea. + 11. cavity occupied by the membranous labyrinth.] + +The =petrous portion= lies dorsally and anteriorly, and is much the +more important of the two, as it encloses the essential part of the +auditory organ. It forms an irregular mass of hard dense bone, +projecting into the cranial cavity, and does not appear on the +external surface at all. The =mastoid portion= lies ventrally and +posteriorly, is smaller, and formed of less dense bone than is the +petrous portion, from which it differs also in the fact that it +appears on the surface of the skull, just external to the exoccipital. +The petrous portion bears a ridge, which together with a ridge on the +supra-occipital, and the =tentorium= (fig. 72, 21), a transverse fold +of the dura mater[142], separates the large cerebral fossa from the +=cerebellar fossa=, which is much smaller than the cerebral fossa and +lies behind and partly beneath it. The plane of the tentorium is +called the =tentorial plane=, and the angles that it makes with the +basicranial axis and with the occipital and ethmoidal planes vary much +in different mammals. + +The periotic has its inner surface marked by important depressions, +while both inner and outer surfaces are pierced by foramina. At about +the middle of its inner surface are seen two deep pits, one lying +immediately above the other. Of these the more ventral is a foramen, +the =internal auditory meatus= (fig. 72, 20), through which the VIIth +(facial) and VIIIth (auditory) nerves leave the cranial cavity, the +facial nerve passing through the bone and afterwards leaving the skull +by the =stylomastoid foramen= (fig. 75, VII), while the auditory +passes to the inner ear. The more dorsal of the two pits is not a +foramen but the =floccular fossa= (fig. 72, 23) which lodges the +floccular lobe of the cerebellum. In some skulls another wide and +shallow but fairly prominent depression is seen dorsal to and slightly +behind the floccular fossa, this also lodges part of the cerebellum. +Behind the internal auditory meatus, between the periotic and +exoccipital is seen the internal opening of the =foramen lacerum +posterius= (fig. 72, 22). The shape of this opening varies. The +ventro-anterior border of the periotic is marked by a deep notch, the +sides of which sometimes unite, converting it into a foramen. + +On the outer side of the periotic, and clearly seen only after the +removal of the tympanic, are two holes, the =fenestra ovalis= and the +=fenestra rotunda=. + +The _tympanic_ (figs. 72, 15 and 75, 4) is a greatly expanded +boat-shaped bone, which forms the auditory bulla and lies immediately +ventral to the periotic; it is separated from the periotic by the +=tympanic cavity= into which the fenestra rotunda and the fenestra +ovalis open. + +There are several other openings into the tympanic cavity. + +(_a_) On the external surface is a large oval opening, the =external +auditory meatus= bounded by a thickened rim. + +(_b_) Into the outer and anterior part of the cavity the outer end of +the =Eustachian tube= opens; while the inner end passes through a +foramen (fig. 75, 22) just external to the foramen lacerum medium, on +its way to open into the pharynx. + +(_c_) The internal carotid artery also enters the tympanic cavity by a +canal which commences in the foramen lacerum posterius, and passes +forwards to open on the inner side of the bulla. The artery then +passes forwards, and barely appearing on the ventral surface of the +cranium, enters the brain cavity through the foramen lacerum medium +(fig. 75, 9). + +Immediately behind the tympanic, between it and the mastoid process of +the periotic and the paroccipital process of the exoccipital is the +=stylomastoid foramen= (fig. 75, VIII). + +Within the tympanic cavity are four small bones, the =auditory +ossicles= (cp. fig. 74), called respectively the =malleus=, =incus=, +=lenticular= and =stapes=; these together form a chain extending from +the fenestra ovalis to the tympanic membrane. + +The =malleus= has a somewhat rounded head (fig. 100, B, 1) which +articulates with the incus, while the other end of the bone is drawn +out into a long process, the =manubrium=, which lies in relation to +the tympanic membrane. The head is also more or less connected by a +thin plate of bone, the =lamella=, to another outgrowth, the +=processus longus=. The =incus= (fig. 100, B, 3) is somewhat +anvil-shaped, and is drawn out into a process which is connected with +the =lenticular=, a nodule of bone interposed between the incus and +the stapes, with which it early becomes united. The =stapes= (fig. +100, B, 2) is stirrup-shaped, consisting of a basal portion from which +arise two =crura=, which meet and enclose a space, the =canal=. + +The _squamosal_ (fig. 73, 7) is a large bone occupying much of the +side wall of the cranial cavity, and articulating above with the +parietal, and behind with the supra-occipital, while in front it +overlaps the frontal and alisphenoid. But though it occupies so large +a space on the outer wall, it forms very little of the internal wall +of the skull, but is really like a bony plate attached to the outer +surface of the cranial wall. The squamosal is drawn out into a strong +forwardly-directed =zygomatic process= which meets the jugal or malar. +The ventral side of the zygomatic process is hollowed out, forming the +=glenoid fossa= (fig. 75, 8), a smooth laterally elongated surface +with which the lower jaw articulates, while the hinder edge of the +glenoid fossa is drawn out into a rounded =postglenoid process= (fig. +75, 23). The articulation is such as to allow but little lateral play +of the lower jaw. + + +(2) =Bones in relation to the Optic capsules.= + +The only bone developed in relation to the optic capsule on each side +is the _lachrymal_. This is a small membrane bone lying between the +frontal and palatine behind, and the maxillae and jugal in front. It is +perforated by a prominent =lachrymal foramen= (fig. 73, 13) which +opens within the orbit. + + +(3) =Bones in relation to the Olfactory capsules.= + +In connection with the =olfactory capsules=, five pairs of bones are +developed, two pairs being membrane bones, and three pairs cartilage +bones. + +Of membrane bones, the _nasals_ (fig. 73, 4) are a pair of long narrow +bones, lying closely side by side, and forming the main part of the +roof of the olfactory chamber. Their posterior ends overlap the +frontals, and the outer margin of each is in contact with the nasal +process of the frontal, and with the maxillae and premaxillae. + +Lying immediately ventral to the nasals, and on each side of the +perpendicular mesethmoid, are the =ethmoid= or =turbinal= bones, +which have a curious character, being formed of a number of delicate +plates intimately folded on one another. The posterior pair of these +bones, the =ethmo-turbinals= (fig. 72, 10), are the larger, and form a +mass of intricately folded lamellae attached behind to the cribriform +plate, and passing laterally into two thin plates of bone, which abut +on the maxillae. The uppermost lamella of each ethmo-turbinal is larger +than the others and more distinct. It is sometimes distinguished as +the =naso-turbinal=, and forms an imperfect lower boundary to a canal, +which is bounded above by the nasals. In front of and somewhat below +the ethmo-turbinals, lie another pair of bones of similar character, +the =maxillo-turbinals= (fig. 72, 11). + +The last bone to be mentioned in connection with the olfactory +capsules is a membrane bone, the _vomer_ (fig. 72, 9). This is a +slender vertically-placed bone, whose anterior part lies between the +maxillo-turbinals, while behind it extends beyond the mesethmoid, so +as to underlie the anterior part of the presphenoid. The anterior part +of the vomer forms a kind of trough, while further back in the region +of the ethmo-turbinals it sends out a pair of strong lateral plates, +each of which, passing below the ethmo-turbinal, joins the side wall +of the nasal cavity, and forms a partition dividing the nasal cavity +into a lower =narial passage= and an upper =olfactory chamber=. + + +THE JAWS. + +In the embryo both upper and lower jaws are formed of cartilaginous +bars, but in the adult not only has the cartilage entirely +disappeared, but even cartilage bone is absent, the jaws being formed +of membrane bone. + + +3. THE UPPER JAW. + +[Illustration FIG. 75. VENTRAL VIEW OF THE CRANIUM OF A DOG (_Canis +familiaris_) × 3/5. (Camb. Mus.) + + 1. supra-occipital. + 2. foramen magnum. + 3. occipital condyle. + 4. tympanic bulla. + 5. basi-occipital. + 6. basisphenoid. + 7. external auditory meatus. + 8. glenoid fossa. + 9. foramen lacerum medium and anterior opening of carotid canal. + 10. postglenoid foramen. + 11. alisphenoid. + 12. presphenoid. + 13. vomer. + 14. jugal. + 15. pterygoid. + 16. palatal process of palatine. + 17. maxillae (palatal portion). + 18. posterior palatine foramina. + 19. anterior palatine foramen. + 20. premaxillae. + 21. alisphenoid canal. + 22. Eustachian foramen. + 23. postglenoid process of squamosal. + II. optic foramen. + III, IV, V{1}, VI. foramen lacerum anterius. + V{2}. foramen rotundum. + V{3}. foramen ovale. + VII. stylomastoid foramen. + IX, X, XI. foramen lacerum posterius. + XII. condylar foramen. + _i_ 2. second incisor. + _c._ canine. + _pm_ 1, _pm_ 4. first and fourth premolars. + _m_ 1. first molar.] + +The bones of the upper jaw are closely connected with those of the +cranium proper and olfactory capsules. The most posterior of them is +the _pterygoid_ (fig. 75, 15), a thin vertically placed plate of bone, +which articulates above with the basisphenoid, the presphenoid, and +the strong pterygoid process of the alisphenoid. The ventral end of +the pterygoid is drawn out into a small backwardly-projecting =hamular +process=. In front the pterygoid articulates with the _palatine_, a +much larger bone, consisting of (1) a vertical portion, which passes +up to meet the orbitosphenoid and frontal, and sends inwards a plate +which meets the presphenoid and vomer, forming much of the roof of the +posterior part of the narial passage; and (2) a strong horizontal +portion, the =palatal process= (fig. 75, 16), which passes inwards and +meets its fellow in the middle line, forming the posterior part of the +bridge of bone supporting the hard palate. The palatal process is +continuous in front, with a large bone, the _maxillae_, which, like +the palatine, consists of vertical and horizontal portions. The +vertical, or =facial portion= (fig. 73, 5), is the largest, and +constitutes the main part of the side of the face in front of the +orbit, forming also the chief part of the outer wall of the nasal +cavity. It is continuous in front with the premaxillae, above with the +nasal and frontal, and behind with the lachrymal, jugal, and palatine. +The horizontal, or =palatal portion= (fig. 75, 17), forms the anterior +part of the bony plate supporting the hard palate, and meets its +fellow in a long straight symphysis. The junction line between the +palatal and facial portions is called the =alveolar border=, and +along it are attached the canine, premolar, and molar teeth. + +The anterior part of the upper jaw on each side is formed by a small +bone, the _premaxillae_, which bears the incisor teeth. It, like the +maxillae, has a palatal portion (fig. 75, 20), which meets its fellow +in the middle line, and an ascending portion, which passes backwards +as the =nasal process=, tapering regularly and lying between the nasal +and the maxillae. The two premaxillae form the outer and lower borders +of the anterior nares. The last bone to be mentioned in connection +with the upper jaw and face is the _jugal_ or _malar_ (figs. 73, 8, +and 75, 14), a strong bone which forms the anterior half of the +zygomatic arch. It is firmly united in front to the maxillae, and +behind meets the zygomatic process of the squamosal, being drawn out +dorsally into a short =postorbital process= at the point of meeting. +This process lies immediately below the postorbital process of the +frontal, and if the two met, as they do in some mammals, they would +partially shut off the orbit from a larger posterior cavity, the +=temporal fossa=. In the living animal a ligament unites the two +postorbital processes. + + +(_b_) THE LOWER JAW OR MANDIBLE. + +This consists of two elongated symmetrical halves, the =rami=, which +are united to one another at the median symphysis in front, while +behind they diverge considerably, and each articulates with the +glenoid surface of the corresponding squamosal. In young animals the +rami are united at the symphysis by fibrous tissue, but in old animals +they sometimes become fused together. The upper or alveolar border +bears the teeth, and behind them is drawn out into a high laterally +compressed =coronoid process= (fig. 72, 24), which is hollowed on its +outer surface. Immediately behind the coronoid process is the +transversely elongated =condyle= (fig. 72, 25), which fits into the +glenoid cavity in such a way as to allow free up and down movement of +the jaw, with but little rolling motion. The posterior end of the jaw +below the condyle forms a short rounded process, the =angle= (fig. +72, 26). Two prominent foramina are to be seen in the lower jaw. These +are firstly the =inferior dental foramen= (fig. 72, 28), which lies on +the inner surface below the coronoid process; through it an artery and +a branch of the fifth nerve enter to supply the teeth, and secondly +the =mental foramen=, which lies on the outer side near the anterior +end, and through which a branch of the same nerve emerges. + + +(_c_) THE HYOID. + +The =Hyoid= of the dog consists of a transverse median piece, the +=basi-hyal=[1] (fig. 72, 32), from which arise two pairs of =cornua=. +The =anterior cornu= is much the longer of the two, and consists +principally of three short separate ossifications, placed end to end +and called respectively the =cerato-hyal=[143], =epi-hyal=, and +=stylo-hyal=. All of them are short rods of bone, contracted in the +middle, and expanded at the ends, where they are tipped with +cartilage. The cerato-hyal (fig. 72, 31) lies next to the basi hyal. +The stylo-hyal is terminated by a much smaller bone, the +=tympano-hyal=, which lies in a canal between the tympanic and +periotic, and is ankylosed to the periotic just to the anterior and +inner side of the stylomastoid foramen. + +The =posterior cornu= of the hyoid is much smaller than the anterior; +it consists of a short bone, the =thyro-hyal= (fig. 72, 33), which +connects the basi-hyal with the thyroid cartilage of the larynx. + +FORAMINA OF THE SKULL. + +The foramina, or apertures perforating the walls of the skull, are +very numerous, and may either be due to holes actually penetrating the +bone, or may be small vacuities between the margins of two elsewhere +contiguous bones. + +They may be divided into two groups, the first including + +I. The holes through which the =twelve cranial nerves= leave the +cranial cavity. + +_a._ The most anterior of these nerves, the olfactory, leaves the +skull by a number of small holes piercing the =cribriform plate= (fig. +72, 5). + +_b._ The second, or optic, passes out by a large hole, the =optic +foramen= (fig. 75, II) piercing the orbitosphenoid. The optic foramen +is the most anterior of the three prominent holes seen within and +immediately behind the orbit. + +_c._ The third, fourth, and sixth nerves, i.e. those supplying the eye +muscles, and with them the first or ophthalmic branch of the large +fifth or trigeminal nerve, pass out by a large hole, the =foramen +lacerum anterius= (fig. 75, III, IV, V{1}, VI), which, as has been +already mentioned, lies between the orbitosphenoid and alisphenoid. + +_d._ Immediately behind the foramen lacerum anterius, the alisphenoid +is perforated by a prominent round hole, the =foramen rotundum= (fig. +75, V{2}), through which the second branch of the trigeminal nerve +passes out. + +_e._ A quarter of an inch further back there is another prominent +hole, the =foramen ovale= (fig. 75, V{3}), through which the third +branch of the trigeminal nerve leaves the cranium. + +_f._ The seventh or facial nerve, as already mentioned, leaves the +cranial cavity and enters the auditory capsule, through an opening in +the periotic called the =internal auditory meatus=, while it finally +leaves the skull by the =stylomastoid foramen= (fig. 75, VII), which +lies between the tympanic bulla, the paroccipital process, and the +mastoid portion of the periotic. + +_g._ The eighth or auditory nerve on leaving the cranial cavity, +passes with the facial straight into the auditory capsule through the +=internal auditory meatus= (fig. 72, 20). It is then distributed to +the organ of hearing. + +_h._ The ninth, tenth and eleventh nerves leave the skull through the +=foramen lacerum posterius= (fig. 75, IX, X, XI), a large space lying +between the auditory bones and the exoccipital. + +_i._ Finally, the twelfth nerve, the hypoglossal, passes out through +the prominent =condylar foramen= (fig. 75, XII), which perforates the +exoccipital just behind the foramen lacerum posterius. + + +II. OTHER OPENINGS IN THE SKULL. + +_a._ The =anterior narial opening= lies at the anterior end of the +skull, and is bounded by the premaxillae and nasals. In the natural +condition it is divided into two by a vertical partition, formed by +the =narial septum=, the anterior unossified part of the mesethmoid. + +_b._ Penetrating the middle of the maxillae at the side of the face is +the rather large =infra-orbital foramen= (fig. 73, 11), through which +part of the second branch of the trigeminal nerve passes out from the +orbit to the side of the face. + +_c._ Several foramina are seen perforating the anterior part of the +orbit. The most dorsal of these, perforating the lachrymal bone, is +the =lachrymal foramen= (fig. 73, 13). Lying below and slightly +external to this is a large foramen, through which part of the second +branch of the trigeminal enters on its way to the infra-orbital +foramen and so to the side of the face. Lastly, lying below these, and +perforating the palatine, are two closely apposed foramina, the +=internal orbital foramina=, through which part of the first or +ophthalmic branch of the trigeminal nerve leaves the orbit, passing +into the nasal cavity. + +_d._ The anterior part of the palate between the premaxillae and the +maxillae is perforated by a pair of long closely apposed apertures, the +=anterior palatine foramina= (fig. 75, 19). They transmit part of the +trigeminal nerve. + +_e._ Towards the posterior part of the palate are two pairs of small +=posterior palatine foramina= (fig. 75, 18). These perforate the +palatine and transmit branches of the trigeminal nerve and certain +blood-vessels. + +_f._ The =posterior narial opening= is bounded chiefly by the +palatines. + +_g._ The =alisphenoid canal= (fig. 75, 21) is a short canal +penetrating the base of the alisphenoid bone, and transmitting the +external carotid artery. It lies between the foramen rotundum and the +foramen ovale. + +_h._ Between the auditory bulla and the foramen ovale are seen two +openings. The more external of these is the opening of the =Eustachian +canal= (fig. 75, 22), which communicates with the tympanic cavity. The +more internal is the =foramen lacerum medium= (fig. 75, 9), through +which the internal carotid enters the cranial cavity. + +_i._ The =external auditory aperture= (fig. 75, 7) is a large opening +with rough edges at the outer side of the tympanic bulla. + +_j._ Between it and the glenoid surface of the squamosal is the +=postglenoid= foramen (fig. 75, 10) through which a vein passes out. + +_k._ Lastly, there is the great =foramen magnum= (fig. 75, 2), between +the occipital condyles. Through it the brain and spinal cord +communicate. + + +C. THE RIBS AND STERNUM. + +[Illustration FIG. 76. STERNUM AND STERNAL RIBS OF A DOG (_Canis +familiaris_) × 1/2. + + 1. presternum. + 2. first sternebra of mesosternum. + 3. last sternebra of mesosternum. + 4. xiphisternum. The flattened cartilaginous plate terminating the + xiphisternum is not shown. + 5. first sternal rib.] + +These, together with the thoracic vertebrae, form the skeletal +framework of the thorax. Each rib is a curved rod, which at its dorsal +end is movably articulated to the vertebra, and at its ventral end is +either connected with the sternum, or ends freely. In the dog there +are thirteen pairs of ribs, nine pairs of which are directly connected +with the sternum, while the remaining four end freely and are known as +=floating ribs=. Each rib is obviously divided into two parts, a +dorsal or =vertebral part=, and a ventral or =sternal part=. The +vertebral portion, which forms about two-thirds of the whole rib, is a +flattened, regularly curved rod, completely ossified. Its dorsal end +is rounded, forming the =head= or =capitulum=, which articulates with +a concave surface furnished partly by the corresponding vertebra and +partly by the vertebra next in front. The last three or four however +articulate with one vertebra only. A short way behind the capitulum on +the dorsal side of the rib is a rounded outgrowth, the =tubercle= or +=tuberculum=, by means of which the rib articulates with the +transverse process. The portion of the rib between the head and the +tubercle is known as the =neck=. The =sternal portion= of the rib +(fig. 76) is a short bar of calcified or imperfectly ossified +cartilage, about one-third of the length of the corresponding bony +portion. The anterior sternal ribs are somewhat more cartilaginous +than the posterior ones. The vertebral portions increase in length +from the first which is very stout, and has the capitulum and +tuberculum very distinct, to about the eighth or ninth; afterwards +they gradually diminish in size. The first nine to eleven have the +capitula and tubercula separate, afterwards they gradually merge +together. + + +THE STERNUM. + +This is an elongated cylindrical structure lying in the mid-ventral +wall of the thorax, and is divided into eight segments or +=sternebrae=. The anterior segment, the =presternum= (fig. 76, 1) or +=manubrium sterni= is expanded in front; the next six segments, which, +together form the =mesosternum= are elongated, somewhat contracted in +the middle and expanded at the ends. The last segment or +=xiphisternum= (fig. 76, 4) is long and narrow, and terminates in a +flattened expanded plate of cartilage. The first pair of sternal ribs +articulate with the sides of the presternum, and the remaining pairs +between the successive sternebrae. Between the last sternebra and the +xiphisternum two pairs articulate. Development shows that the sternum +is formed by the union in the middle line of two lateral portions; +this can be well seen in the presternum and xiphisternum of the puppy, +but no traces of this median division remain in the adult dog. + + +2. THE APPENDICULAR SKELETON. + +The appendicular skeleton consists of the bones of the anterior and +posterior limbs, and of their respective supports, the pectoral and +pelvic girdles. + + +THE PECTORAL GIRDLE. + +The =pectoral girdle= lies external to the ribs, and has no bony +attachment to the axial skeleton. In almost all Mammalia it is, as +compared with that in Sauropsids, very incomplete; and in the dog it +is even more reduced than in the majority of Mammalia. The dorsal +portion or =scapula= is well developed, but the ventral portion is +almost entirely absent. + +The =scapula= is somewhat triangular in shape, the apex being +directed downwards and forwards, and being expanded to form the +shallow =glenoid cavity= with which the head of the humerus +articulates. The inner surface of the scapula is nearly flat, while +the outer is drawn out into a very prominent ridge, the =spine=, +which, arising gradually near the dorsal end, runs downwards, dividing +the surface into two nearly equal parts, the =prescapular= and +=postscapular fossae=, and ends in a short blunt process, the +=acromion=. The anterior border of the scapula is somewhat curved, and +is called the =coracoid border=; it is terminated ventrally by a +slight blunt swelling, the =coracoid process=, which ossifies from a +different centre from the rest of the scapula, and is probably the +sole representative of the =coracoid=. The dorsal or =suprascapular +border= of the scapula is rounded, while the posterior or =glenoid +border= is nearly straight. The clavicle[144] or collar bone, which in +a large proportion of mammals is well seen, in the dog is very +imperfectly developed; it is short and broad, and is suspended in the +muscles, not reaching either the scapula or sternum. + + +THE ANTERIOR LIMB. + +The anterior limb of the dog is divisible into the usual three +portions, the =brachium= or =upper arm=, the =antibrachium= or +=fore-arm=, and the =manus= or =wrist= and =hand=. + +The =brachium= or =upper arm= includes only a single bone, the +=humerus=. + +The =humerus= is a stout elongated bone, articulating by its large +proximal =head= (fig. 77, 1) with the glenoid cavity of the scapula, +and at its distal end by the =trochlea= with the bones of the +fore-arm. The head passes on its inner side into an area roughened for +the attachment of muscles and called the =lesser tuberosity= (fig. 77, +2); while in front it is divided by the shallow =bicipital groove= +from a large roughened area, the =greater tuberosity= (fig. 77, 3), +which is continued as a slight roughened ridge, extending about +one-third of the way down the outer side of the shaft. This ridge, +which in many animals is much more strongly developed than it is in +the dog, is called the =deltoid ridge=. The =trochlea= (fig. 77, 5) at +the distal end of the bone is a pulley-like surface, elevated at the +sides and grooved in the middle. It articulates with the radius and +ulna of the fore-arm. On each side of it are slight roughened +projections, the =internal= and =external condyles= (fig. 77, 7). In +the cat and many other animals there's a foramen, the =ent-epicondylar +foramen= above the internal condyle, but in the dog this is not +developed. Passing up the shaft from the external condyle is a slight +ridge, the =supinator= or =ectocondylar ridge=; this is better +developed in many mammals. Immediately above the trochlea in front and +behind are the deep =supra-trochlear fossae=, which communicate with +one another through the =supra-trochlear foramen= (fig. 77, 8). The +posterior of these, the =olecranon fossa=, is much the deeper, and +receives the olecranon process of the ulna when the arm is extended. +The head and tuberosities of the humerus ossify from one centre, the +shaft from a second, and the trochlea and condyles from a third. + +The =fore-arm= or =antibrachium= contains two bones, the =radius= and +=ulna=; they are immovably articulated with one another, but not +fused. The pre-axial bone, the =radius= (fig. 77, B), which lies more +or less in front of the ulna, is external to the ulna at its proximal +end, and at its distal end is internal to that bone. It articulates +with the external portion of the trochlea, while the ulna articulates +with the internal portion. It is a straight bone with its distal end +slightly larger than its proximal end. The proximal end articulates +with the trochlea, the distal end with the bones of the carpus. + +[Illustration FIG. 77. BONES OF THE LEFT UPPER ARM AND FORE-ARM OF A +DOG (_Canis familiaris_) × 1/2. + +A, humerus (seen from the posterior side); B, radius, C, ulna, both +seen from the anterior side. + + 1. head. + 2. lesser tuberosity. + 3. greater tuberosity. + 4. shaft of the humerus. + 5. trochlea. + 6. internal condyle. + 7. external condyle. + 8. supra-trochlear foramen. + 9. proximal end of the radius. + 10. shaft of the radius. + 11. olecranon. + 12. surface for articulation with the trochlea. + 13. surface for articulation with the radius. + 14. distal end of the ulna.] + +The postaxial bone, the =ulna= (fig. 77, C), has the proximal end much +enlarged, forming the =olecranon= (fig. 77, 11), and tapers gradually +to the distal end. Near its proximal end the ulna is marked by a deep +=sigmoid notch=, which bears on its inner side a concave surface +(fig. 77, 12) for articulation with the trochlea. The pointed proximal +end of the sigmoid notch is called the =coronoid process=. Somewhat +in front of and below the sigmoid notch is a smaller hollow (fig. 77, +13), with which the radius articulates. + +In the young animal the ends of both radius and ulna are seen to +ossify from centres different from those forming the shafts. The +epiphyses forming both ends of the radius, and the distal end of the +ulna are large, while that at the proximal end of the ulna is small, +and forms only the end of the olecranon. + +The =Manus= is divided into + +_a._ The =carpus= or =wrist=, formed of a group of small bones. + +_b._ The =hand=, which includes firstly some elongated bones, the +=metacarpals=, forming what corresponds to the palm of the hand, and +secondly the phalanges, which form the =fingers=. + +The =Carpus= or =wrist=. The carpus of the dog consists of seven small +bones, arranged in a proximal row of three, and a distal row of four. +It differs much from the simpler type met with in the newt. The +largest bone of the proximal row is the =scapho-lunar= (fig. 80, 1), +formed by the fused =scaphoid= (radiale), =lunar= (intermedium), and +=centrale=; it has a large convex proximal surface for articulation +with the radius, and articulates distally with the trapezium, +trapezoid, and magnum, and internally with the cuneiform. The +=cuneiform= (ulnare) (fig. 80, 2) has a posterior rounded surface +articulating with the ulna; it articulates in front with the unciform, +and internally with the =pisiform= (fig. 80, 7), which is a +comparatively large sesamoid bone on the ulnar side of the carpus. +Frequently also there is a small sesamoid bone on the radial side of +the carpus. The =trapezium= (carpale 1), =trapezoid= (carpale 2), and +=magnum= (carpale 3) (fig. 80, 5) are all small bones, and support +respectively the first, second, and third metacarpals. The =unciform= +(carpalia 4 and 5) (fig. 80, 6) is larger, and supports the fourth and +fifth metacarpals. + +The hand has five =digits=, each consisting of an elongated +=metacarpal=, followed by =phalanges=, the last of which, the =ungual +phalanx=, is pointed and curved, and bears the claw. Each of the +metacarpals is seen in the young animal to have its distal end formed +by a prominent epiphysis, and each of the phalanges, except those +bearing the claws, has a similar epiphysis at its proximal end. + +The =pollex= (fig. 80, A, I ) is far shorter than the other digits, +and normally does not touch the ground in walking. It has only two +phalanges, while each of the other digits has three. A pair of small +sesamoid bones are developed on the ventral or flexor side of the +metacarpo-phalangeal articulations of all the digits except the +pollex. Frequently similar sesamoid bones occur also on the dorsal +side of the phalangeal articulations. + + +THE PELVIC GIRDLE. + +The =pelvic girdle= consists of two halves, which lie nearly parallel +to the vertebral column. + +Each half is firmly united to its fellow in a ventral symphysis +behind, and is in front expanded and united to the sacrum. Each half +or =innominate bone= is seen in the young animal to consist of four +distinct parts, the =ilium= or dorsal element, the =pubis= or anterior +ventral element, the =ischium= or posterior ventral element, and a +small fourth part, the =acetabular= or =cotyloid= bone, wedged in +between the three others. These parts, though all distinct in the +young animal, are in the adult so completely fused that their +respective boundaries cannot be distinguished. At about the middle of +the outer surface of the innominate bone is a very deep cavity, the +=acetabulum= (fig. 78, A, 1) with which the head of the femur +articulates; all the bones except the pubis take part in its +formation. + +The =ilium= is a rather long bone, expanded in front and contracted +behind; it forms about half the acetabulum. On its inner or =sacral +surface= (fig. 78, 4) is a large roughened patch for articulation +with the sacrum; its outer or =gluteal surface= is concave. The +posterior part of the bone is flattened below, forming the narrow +=iliac surface= (fig. 78, A, 5). + +[Illustration FIG. 78. RIGHT INNOMINATE BONE, A, OF A FULL-GROWN +TERRIER, B, OF A COLLIE PUPPY. × 1. + +A is seen from the ventral side, B from the inner or sacral side. + + 1. acetabulum. + 2. thyroid foramen. + 3. supra-iliac border of ilium. + 4. sacral surface. + 5. iliac surface. + 6. acetabular border. + 7. pubic border. + 8. ischial border. + 9. ischium. + 10. tuberosity of ischium. + 11. ischial symphysis. + 12. pubis. + 13. pubic symphysis. + 14. cotyloid or acetabular bone.] + +The =ischium= (fig. 78, 9) is a wide flattened bone forming the +posterior part of the innominate bone. It meets the pubis ventrally, +but is separated from it for the greater part of its length by the +large =obturator= or =thyroid foramen= (fig. 78, 2). At its posterior +end externally it bears a rather prominent roughened =ischial +tuberosity= (fig. 78, A, 10). The ischium meets its fellow in a +ventral symphysis, and forms about one-third of the acetabulum. + +[Illustration FIG. 79. FRONT VIEW OF THE LEFT LEG BONES OF A DOG +(_Canis familiaris_) × 1/2. + +A femur, B tibia, C fibula, D patella. + + 1. head of femur. + 2. neck. + 3. great trochanter. + 4. shaft. + 5. external condyle. + 6. internal condyle. + 7. fabella. + 8. cnemial crest.] + +The =pubis= (fig. 78, 12) is smaller than either the ischium or ilium; +it does not take part in the formation of the acetabulum, and like the +ischium, meets its fellow in a ventral symphysis. The =acetabular +bone= (fig, 78, B, 14) is small and triangular, and is wedged in +between the other three. It forms about one-sixth of the acetabulum. + + +THE POSTERIOR LIMB. + +The =posterior limb=, like the anterior, is divisible into three +parts; these are the =thigh=, the =crus= or =shin=, and the =pes=. + +The =thigh= contains only a single bone, the =femur=. + +The =femur= is a long straight bone with a nearly smooth shaft and +expanded ends. The proximal end bears on its inner side the large +rounded =head= (fig. 79, A, 1) which articulates with the acetabulum. +External to the head and divided from it by a deep pit is a large +rough outgrowth, the =great trochanter= (fig. 79, 3). The deep pit is +the =trochanteric= or =digital fossa=. On the inner side below the +head is a smaller roughened surface, the =lesser trochanter=. The +lower or distal end of the bone bears two prominent rounded surfaces, +the =condyles=, which articulate with the tibia. They are separated +from one another by the deep =intercondylar notch=, which is continued +above and in front as a shallow groove, lodging a large sesamoid bone, +the =patella= or =knee-cap=. At the back of the knee-joint are a pair +of smaller sesamoids, the =fabellae= (fig. 79, 7). + +In the young animal there are three epiphyses to the shaft of the +femur, one forming the head, one the great trochanter, and one the +distal end. + +The =crus= or =shin= contains two bones, the =tibia= and =fibula=. The +=tibia= is a fairly thick straight bone, expanded at both ends, +especially at the head or proximal end. The proximal end is triangular +in cross section, and bears two facets for articulation with the +condyles of the femur. The anterior surface of the proximal end of the +tibia is marked by the strong =cnemial crest= (fig. 79, 8), which runs +some way down the shaft. The distal end of the tibia articulates with +the astragalus by an irregular, somewhat square surface. + +The shaft of the tibia ossifies from one centre, the distal end from a +second, and the proximal end from two more. + +[Illustration FIG. 80. A, RIGHT MANUS, B, RIGHT PES OF A DOG (_Canis +familiaris_) × 1/2 (after VON ZITTEL). + + 1. bone representing the fused scaphoid, lunar and centrale. + 2. cuneiform. + 3. trapezium. + 4. trapezoid. + 5. magnum. + 6. unciform. + 7. pisiform. + 8. first metacarpal. + 9. fifth metacarpal. + 10. astragalus. + 11. calcaneum. + 12. navicular. + 13. middle cuneiform. + 14. external cuneiform. + 15. cuboid. + 16. first metatarsal. + +The digits are numbered with Roman numerals.] + +The =fibula= (fig. 79, C) is a distinct but very slender bone, +somewhat expanded at both ends. It lies external to the tibia and +articulates by its proximal end with the head of the tibia, and by its +distal end with the calcaneum. Its shaft and proximal end ossify from +one centre, and its distal end from a second. + + +The =Pes=. + +The structure of the =pes= corresponds closely with that of the manus. +It is divided into:-- + +_a._ The =tarsus= or =ankle= formed of a group of small bones. + +_b._ The =foot=, which includes, firstly, some elongated bones, the +=metatarsals=, forming what corresponds to the sole of the foot, and +secondly the =phalanges=, which form the toes. + +The =Tarsus=. The tarsus of the dog consists of seven bones arranged +in two rows, of two and four respectively, with a =centrale= between +them. The two bones of the proximal row are the =astragalus= and +=calcaneum=. + +The =astragalus= (fig. 80, 10) corresponds to the fused =tibiale= and +=intermedium= of the typical tarsus. Its proximal end is much wider +than its distal end, and forms a large rounded =condyle= articulating +with the tibia, while its posterior end meets the navicular. It lies +to the dorsal side of the foot. + +The =calcaneum= (fibulare) (fig. 80, 11), the thickest bone in the +pes, lies somewhat behind, and to the outer side of the astragalus. It +articulates with the astragalus and fibula, and is drawn out behind +into a long rounded process, which forms the heel, and is in the young +animal terminated by an epiphysis. Between the proximal and distal +rows of tarsals is the =navicular= (centrale) (fig. 80, 12), a +somewhat flattened and square bone articulating with the astragalus. + +The distal row of tarsals consists of four bones. The =internal +cuneiform= (tarsale 1) is a smooth flattened bone lying to the inner +side of the foot; it articulates with the first metatarsal and with +the navicular. The =middle cuneiform= (tarsale 2) (fig. 80, 13) is a +still smaller bone, lying external to the internal cuneiform. It +articulates with the second metatarsal and with the navicular. The +=external cuneiform= (tarsale 3) (fig. 80, 14) is a larger, somewhat +square bone lying external to the middle cuneiform. It articulates +with the third metatarsal and with the navicular. The =cuboid= +(tarsalia 4 and 5) (fig. 80, 15) is a considerably larger bone lying +to the outer side of the foot. It articulates with the fourth and +fifth metatarsals and with the calcaneum. + +The pes has sometimes five digits, sometimes four, the hallux being +absent. Even when present the =hallux= (fig. 80, =B, I=) is commonly +much reduced, and may be quite vestigial, and represented only by a +small nodular metatarsal. + +Each of the other digits consists of a long metatarsal, which in the +young animal has a prominent epiphysis at its distal end, and of three +phalanges. The proximal and middle phalanges have epiphyses at their +proximal ends, while the distal phalanx is without epiphyses and is +claw-shaped. + + +FOOTNOTES: + +[139] W. Ellenberger and H. Baum, _Anatomie des Hundes_, Berlin, 1891. + +[140] T.H. Huxley, "Dental and cranial characters of the Canidae," +_P.Z.S._ 1880. + +[141] See p. 392. + +[142] The dura mater is a membrane which lines the cranial cavity and +is formed of tough connective tissue. + +[143] These are not strictly homologous with the basi-hyal and +cerato-hyal of the Dogfish. + +[144] See note to p. 25. + + + + +CHAPTER XXII. + +GENERAL ACCOUNT OF THE SKELETON IN MAMMALIA. + + +THE EXOSKELETON AND VERTEBRAL COLUMN. + + +EPIDERMAL EXOSKELETON. + +=Hair=, which forms the characteristic Mammalian exoskeleton, varies +much in different animals, and in different parts of the same animal. +A large proportion of mammals have the surface fairly uniformly +covered with hair of one kind only. In some forms however there are +two kinds of hair, a longer and stiffer kind alone appearing on the +surface, and a shorter and softer kind forming the under fur. In most +mammals hairs of a special character occur in certain regions, such as +above the eyes, on the margins of the eyelids, and on the lips and +cheeks, here forming the vibrissae or whiskers. + +Sometimes as in _Hippopotamus_, _Orycteropus_ and the Sirenia, the +hair, though scattered over the whole surface, is extremely scanty, +while in the Cetacea it is limited to a few bristles in the +neighbourhood of the mouth, or may even be absent altogether in the +adult. In most mammals the hairs are shed and renewed at intervals, +sometimes twice a year, before and after the winter. The vibrissae or +large hairs which occur in many animals upon the upper lip, and the +mane and tail of Equidae are probably persistent. + +In the hedgehogs, porcupines and _Echidna_ certain of the hairs are +modified and greatly enlarged, forming stiff spines. Similar spines +occur in the young of _Centetes_, and in _Acanthomys_ among the +Muridae. + +Several other forms of epidermal exoskeleton are met with in mammals, +including:-- + +(_a_) =Scales=. These overlie the bony scutes of armadillos and occur +covering the tail in several groups of mammals, such as beavers and +rats. In the Manidae the body is covered by flat scales which overlap. + +(_b_) The =horns= of Bovine Ruminants. These, which must on no account +be confused with antlers, are hollow cases of hardened epidermis +fitting on to bony outgrowths of the frontals. In almost every case +they are unbranched structures growing continuously throughout life, +and are very rarely shed entire. In the Prongbuck _Antilocapra_ +however they are bifurcated and are periodically shed. Horns are +nearly always limited to a single pair, but the four-horned antelope +_Tetraceros_ has two pairs, the anterior pair being the smaller. + +(_c_) The =horns of Rhinoceroses=. These are conical structures +composed of a solid mass of hardened epidermal cells growing from a +cluster of long dermal papillae. From each papilla there grows a fibre +which resembles a thick hair, and cementing the whole together are +cells which grow from the interspaces between the papillae. These +fibres differ from true hairs in not being developed in pits in the +dermis. Rhinoceros horns may be either one or two in number, and are +borne on the fronto-nasal region of the skull. They vary much in +length, the longest recorded having the enormous length of fifty-seven +inches. + +(_d_) _Nails_, _hoofs_ and =claws=. In almost all mammals except the +Cetacea, these are found terminating the digits of both limbs. =Nails= +are more or less flattened structures, =claws= are pointed and +somewhat curved. In most mammals the nails tend to surround the ends +of the digits much more than they do in man. Sometimes the nail of one +digit differs from that of all the others; thus the second digit of +the pes in the Hyracoidea and Lemuroidea is terminated by a long claw, +the other digits having flat nails. In the Felidae the claws are +retractile, the ungual phalanx with claw attached folding back when +the animal is at rest into a sheath, above, or by the side of the +middle phalanx. In the Sloths and Bats enormously developed claws +occur, forming hooks by which the animals suspend themselves. In +_Notoryctes_ the third and fourth digits of the manus bear claws of +great size; similar claws occur in _Chrysochloris_, being correlated +in each case with fossorial habits. The nail at its maximum +development entirely surrounds the terminal phalanx of the digit to +which it is attached, and is then called a =hoof=. Hoofs are specially +characteristic of the Ungulata. + +(_e_) =Spurs= and =beaks= are structures which are hardly represented +among mammals, while so characteristic of birds. They are however both +found in the Monotremata. In both _Echidna_ and _Ornithorhynchus_ the +male has a peculiar hollow horny spur borne on a sesamoid bone +articulated to the tibia. The jaws in _Ornithorhynchus_ are cased in +horny beaks similar to those of birds, and are provided with horny +pads which act as teeth. + +(_f_) =Horny plates= of a ridged or roughened character occur upon the +anterior portion of the palate, and of the mandibular symphysis in all +three genera of recent Sirenia; also upon the toothless anterior +portion of the palate in Ruminants. + +(_g_) The =baleen of whales= also belongs to the epidermal +exoskeleton. It consists of a number of flattened horny plates +arranged in a double series along the palate. The plates are somewhat +triangular in form and have their bases attached to the palate at +right angles to its long axis, while their apices hang downwards into +the mouth cavity. The outer edge of each plate is hard and smooth, +while the inner edge and apex fray out into long fibres which look +like hair. At the inner edge of each principal plate are subsidiary +smaller plates. The plates are formed of a number of fibres each +developed round a dermal papilla in the same way as are the fibres +forming the horns of _Rhinoceros_. Baleen and Rhinoceros horn likewise +agree in that the fibres are bound together by less hardened +epithelial cells, which readily wear away and allow the harder fibres +to fray out. The greatest development of baleen occurs in the Northern +Right whale, _Balaena mysticetus_, in which the plates number three +hundred and eighty or more on each side, and reach a length of ten or +twelve feet near the middle of the series. + + +DERMAL EXOSKELETON. + +Mammals show two principal kinds of exoskeletal structures which are +entirely or partially dermal in origin, viz. the bony scutes of +armadillos, and teeth. + +The =bony scutes of armadillos= are quadrate or polygonal in shape and +are in general aggregated together, forming several shields protecting +various regions of the body. The head is generally protected by a +_cephalic_ shield, the anterior part of the body by a _scapular_, and +the posterior by a _pelvic_ shield. The tail is also generally encased +in bony rings, and scutes are irregularly scattered over the surface +of the limbs. The mid-body region is protected by a varying number of +bands of scutes united by soft skin, so as to allow of movement. +Corresponding to each dermal scute is an epidermal plate. In +_Chlamydophorus_ the scutes are mainly confined to the posterior +region where they form a strong vertically-placed shield which +coalesces with the pelvis. The anterior part of the body is mainly +covered by horny epidermal plates with very little ossification +beneath. In the gigantic extinct Glyptodonts the body is covered with +a solid carapace formed by the union of an immense number of plates, +and there are no movable rings. The top of the head is defended by a +similar plate, the tail is generally encased in an unjointed bony +tube, and there is commonly a ventral plastron. + +In _Phocaena phocaenoides_ the occurrence of vestigial dermal ossicles +has been described, and in _Zeuglodon_ the back was probably protected +by dermal plates. + +TEETH[145]. + +Teeth are well developed in the vast majority of mammalia, and are of +the greatest morphological and systematic importance, many extinct +forms being known only by their teeth. Mammalian teeth differ from +those of lower animals in various well-marked respects. (1) They are +attached only to the maxillae, premaxillae and mandible, never to the +palatines, pterygoids or other bones. (2) They frequently have more +than one root. (3) They are always, except in some Odontoceti, placed +in distinct sockets. (4) They are hardly ever ankylosed to the bone. +(5) They are in most cases markedly heterodont. (6) They are commonly +developed in two sets, the milk dentition and permanent dentition. + +It sometimes happens that teeth after being formed are reabsorbed +without ever cutting the gum. This is the case, for instance, with the +upper incisors of Ruminants. + +The form of mammalian teeth varies much, some are simple conical +structures comparable to those of most reptiles, and these may either +have persistent pulps, as in the case of the upper canines of the +Walrus and the tusks of Elephants, or may be rooted as in most canine +teeth. Some teeth have chisel-shaped edges, and this may be their +original form, as in the human incisors, or may, as in those of +Rodents, be brought about by the more rapid wearing away of the +posterior edge, the anterior edge being hardened by a layer of enamel. +Then, again, the crown may, as in the majority of grinding teeth, be +more or less flattened. The various terms used in describing some of +the forms of the surface of grinding teeth are defined on page 345. + +[Illustration FIG. 81. SKULL OF A YOUNG INDIAN RHINOCEROS (_R. +unicornis_), SHOWING THE CHANGE OF THE DENTITION × 1/7. (Brit. Mus.) + + 1. nasal. + 2. frontal. + 3. parietal. + 4. zygomatic process of squamosal. + 5. jugal. + _mI{1}._ milk incisor. + _mc._ milk canine. + _mpm{1}._ milk premolar. + _I{1}._ first incisor. + _c._ canine. + _pm{2}_, _pm{3}_, _pm{4}_. 2nd, 3rd and 4th premolars. + _m{1}_, _m{2}_. first and second molars.] + +The teeth of the Aard Varks are compound, and differ completely from +those of all other mammals (see p. 425). + +As a rule, the higher the general organisation of an animal the +better are its milk teeth developed, and the more do they form a +reproduction on a small scale of the permanent set. This fact is well +seen in the Primates, Carnivora and Ungulata. The method of notation +by which the dentition of any mammal can be briefly expressed as a +formula has been already described. The regular mammalian arrangement +of teeth for each side is expressed by the formula + + _i_ 3/3 _c_ 1/1 _pm_ 4/4 _m_ 3/3 × 2; total, 44. + +MONOTREMATA. In _Echidna_ teeth are quite absent. In the young +_Ornithorhynchus_[146] functional molar teeth of a multi-tubercular +type resembling those of some Mesozoic mammalia are present, but in +the adult they disappear, their office being discharged by horny +plates. + +MARSUPIALIA[147] have a heterodont dentition, which has generally been +regarded as almost monophyodont, the only tooth which has an obvious +deciduous predecessor being the last premolar. The researches of +Röse[148] and Kükenthal[149] tend to show that the teeth of Marsupials +are developed in the same way as in other mammals, and are diphyodont. +In the case of the premolars, teeth which are homologous with the +permanent teeth of other mammals begin to develop as lateral +outgrowths from the milk teeth, but afterwards become absorbed, so +that the teeth which actually persist belong to the milk series. The +last premolar, however, does as a rule develop and replace its milk +predecessor; sometimes, however, as in _Didelphys_, it takes its place +among the milk molars without replacing one of them. + +The types of dentition characteristic of the different groups of +placental mammals may mostly be paralleled among the Marsupials. Thus +among the polyprotodont forms the Didelphyidae or opossums, and some +of the Dasyuridae, such as _Sarcophilus_ and _Thylacinus_, have a +typical carnivorous dentition with small incisors, large canines, and +molars with pointed compressed crowns. The dental formula of +_Thylacinus_, is _i_ 4/3 _c_ 1/1 _pm_ 3/3 _m_ 4/4, total 46. + +In _Myrmecobius_ five or six molar teeth occur on each side, and the +total number of teeth reaches fifty-two or fifty-six. The teeth bear +rows of tubercles, and resemble those of the Multituberculate mesozoic +Mammals[150], more than do those of any other living form. Calcified +teeth have recently been described[151] in _Myrmecobius_ earlier than +the functional or milk set. This would relegate the milk teeth of +mammals in general to a second series, and the permanent teeth to a +third. In _Notoryctes_ the dental formula[152] is given as _i_ 3/2 _c_ +1/1 _pm_ 2/3 _m_ 4/4, total 40. The canines are small, and the +anterior molars have strongly developed cusps, and much resemble those +of _Chrysochloris_ (Insectivora). + +Among the diprotodont types the Phascolomyidae, or Wombats, have a +dentition recalling that of the Rodents. All the teeth grow from +persistent pulps, and the incisors have enamel only on the anterior +surface as in Rodents. The dental formula is _i_ 1/1 _c_ 0/0 _pm_ 1/1 +_m_ 4/4, total 24. There are indications of a vestigial second pair of +incisors. + +The Macropodidae, or Kangaroos, have a herbivorous dentition with the +formula _i_ 3/1 _c_ (0--1)/0 _pm_ 2/2 _m_ 4/4. The incisors are sharp +and cutting, and are separated by a long diastema or gap from the +molars, which have their crowns marked by ridges or cusps. There are +indications of several vestigial incisors. + +_Coenolestes_, a remarkable form recently described from America, +belongs to the diprotodont section, and is the only living member of +the section known outside the Australian region[153]. An exceptional +dentition is seen in the case of the extinct _Thylacoleo_, in which +the functional teeth are reduced to two pairs; one pair of large +cutting incisors and one of compressed sharp-edged premolars. + +EDENTATA. Some Edentata, viz. the ant-eaters (Myrmecophagidae) are, as +far as is known, absolutely toothless at all stages of their +existence; being the only mammals except _Echidna_ in which no tooth +germs have been discovered; others, viz. the Manidae, though showing +foetal tooth germs, are quite toothless in post-foetal life; others, +viz. some of the armadillos, have the largest number of teeth met with +in land mammals. The teeth are homodont except in the Aard Varks, and +grow from persistent pulps. In the sloths (Bradypodidae) and the +Megatheriidae, there are five pairs of teeth in the upper and four in +the lower jaw. The teeth of sloths consist of a central axis of +vasodentine, surrounded firstly by a thin coating of hard dentine, and +secondly by a thick coating of cement. + +In no living Edentate have the teeth any enamel; it has, however, been +described as occurring in certain early Megatheroid forms from S. +America[154], and an enamel organ has also been discovered in an +embryo _Dasypus_[155]. In the Armadillos (Dasypodidae) the number of +teeth varies from 8/8 or 7/7 in _Tatusia_, to upwards of 25/25 in +_Priodon_, which therefore may have upwards of a hundred teeth, the +largest number met with in any land mammal. In _Tatusia_ all the teeth +except the last are preceded by two-rooted milk teeth. The Aard Varks +are diphyodont, and milk teeth are also known in a species of +_Dasypus_, but with these exceptions Edentates are, as far as is +known, monophyodont. In _Glyptodon_ the teeth are almost divided into +three lobes by two deep grooves on each side. + +The Aard Varks (Orycteropodidae) are quite exceptional as regards +their teeth, which are cylindrical in shape, and are made up of a +number of elongated denticles fused together. Each denticle contains a +pulp cavity from which a number of minute tubes radiate outwards. +These teeth are diphyodont and somewhat heterodont, eight to ten pairs +occur in the upper jaw and eight in the lower, but they are not all in +place at one time. The last three teeth in each jaw are not preceded +by milk teeth[156]. + +SIRENIA. The teeth of Sirenia show several very distinct types, the +least modified being that of the extinct Halitheriidae, which have +large incisors in the upper jaw, and five or six pairs of tuberculated +grinding teeth in each jaw, the anterior ones being preceded by milk +teeth. + +In both the living genera the dentition is monophyodont. In _Manatus_ +the dentition is _i_ 2/2 _pm_ and _m_ 11/11. The incisors are +vestigial, and disappear before maturity. The grinding teeth have +square enamelled crowns marked by transverse tuberculated ridges. They +are not all present in the jaw at the same time. In _Halicore_ the +upper jaw bears a pair of straight tusklike incisors; in the male +these have persistent pulps and project out of the mouth; in the +female they soon cease to grow and are never cut. They are separated +by a long diastema from the grinding teeth which have tuberculated +crowns and are 5/5 or 6/6 in number, but are not all in place at once. +Several other pairs of slender teeth occur in the young animal, but +are absorbed or fall out before maturity. In _Rhytina_ teeth are +altogether absent. + + +CETACEA. + +_ARCHAEOCETI._ _Zeuglodon_ has the following dentition, _i_ 3/3 _c_ +1/1 _pm_ and _m_ 5/5, total 36. The incisors and canines are simple +and conical; the cheek teeth are compressed and have serrated cutting +edges like those in some seals. + +In the _MYSTACOCETI_, or whalebone whales, calcified tooth germs +probably belonging to the milk dentition are present in the embryo, +but they are never functional, and are altogether absent in the adult. +The anterior of these germs are simple, the posterior ones are +originally complex, but subsequently split up into simple teeth like +those of the anterior part of the jaw. Hence according to Kükenthal, +who described these structures, the Cetacean dentition was originally +heterodont. + +In the living _ODONTOCETI_ the dentition is homodont and monophyodont. +In some cases traces occur of a replacing dentition which never comes +to maturity, and renders it probable that the functional teeth of the +Odontoceti are really homologous with the milk teeth of other mammals. +Some of the dolphins afford the apparently simplest type of mammalian +dentition known. The teeth are all simple, conical, slightly recurved +structures, with simple tapering roots and without enamel. The +dentition is typically _piscivorous_, being adapted for seizing active +slippery animals such as fish. The prey is then swallowed entire +without mastication. Sometimes the teeth are excessively numerous, +reaching two hundred or more (fifty to sixty on each side of each jaw) +in _Pontoporia_. This multiplication of teeth is regarded by Kükenthal +as due to the division into three parts of numbers of trilobed teeth +similar to those of some seals. + +In the Sperm whale, _Physeter_, the lower jaw bears a series of twenty +to twenty-five stout conical recurved teeth, while in the upper jaw +the teeth are vestigial and remain imbedded in the gum. An extinct +form, _Physodon_, from the Pliocene of Europe and Patagonia is allied +to the Sperm whale, but has teeth in both jaws. In the Grampus _Orca_, +the teeth number about 12/12, and are very large and strong. In some +forms the teeth are very much reduced in number; thus in _Mesoplodon_ +the dentition consists simply of a pair of conical teeth borne in the +mandible. In the Narwhal _Monodon_ the dentition is practically +reduced to a single pair of teeth, which lie horizontally in the +maxillae, and in the female normally remain permanently in the alveoli. +In the male the right tooth remains rudimentary, while the left is +developed into an enormous cylindrical tusk marked by a spiral groove. +Occasionally both teeth develop into tusks, and there is reason for +thinking that two-tusked individuals are generally or always female. +In the extinct _Squalodon_ the dentition is decidedly heterodont, and +the molars have two roots. The dental formula is + + _i_ 3/3 _c_ 1/1 _pm_ 4/4 _m_ 7/7, total 60. + +It is probable that the homodont condition of modern Odontoceti is not +primitive, but due to retrogressive evolution. + + +UNGULATA. + +Just as in the Cetacea a piscivorous dentition is most typically +developed, so the Ungulata are, as a group, the most characteristic +representatives of a _herbivorous_ dentition in its various forms. + + +UNGULATA VERA. + +_ARTIODACTYLA_. As regards the living forms, the Artiodactyla can be +readily divided into two groups, namely those with bunodont and those +with selenodont teeth. It has, however, been shown that selenodont +teeth always pass through an embryonic bunodont stage[157]. The +bunodont type is best seen in Pigs and Hippopotami and such extinct +forms as _Hyotherium_. In _Hippopotamus_ the dental formula is _i_ +(2-3)/(1-3) _c_ 1/1 _pm_ 4/4 _m_ 3/3. + +The incisors and canines of _Hippopotamus_ are very large and grow +continuously. The genus _Sus_, which affords a good instance of an +_omnivorous_ type of dentition, has the regular unmodified Mammalian +dental formula _i_ 3/3 _c_ 1/1 _pm_ 4/4 _m_ 3/3, total 44. The +canines, specially in the male, are large and have persistent pulps, +and the upper canines do not have the usual downward direction but +pass outwards and upwards. In the Wart Hog, _Phacochaerus_, they are +enormously large, but a still more extraordinary development of teeth +is found in _Babirussa_. In the male _Babirussa_ the canines, which +are without enamel, are long, curved and grow continuously. Those of +the upper jaw never enter the mouth, but pierce the skin of the face +and curve backwards over the forehead. The dental formula of +_Babirussa_ is _i_ 2/3 _c_ 1/1 _pm_ 2/2 _m_ 3/3, total 34. + +The Wart Hog has a very anomalous dentition, for as age advances all +the teeth except the canines and last molars show signs of +disappearing; both pairs of persisting teeth are however very large. + +Various extinct Ungulata such as _Anoplotherium_ have teeth which are +intermediate in character between the bunodont and selenodont types. +_Anoplotherium_ has the regular mammalian series of forty-four teeth. +The crowns of all the teeth are equal in height, and there is no +diastema--an arrangement found in no living mammal but man. + +We come now to the selenodont Artiodactyla. + +The Tylopoda--camels (Camelidae) and Llamas (Aucheniidae) when young +have the full number of incisors, but in the adult the two upper +middle ones are lost. The molars are typically selenodont and +hypsodont[158]. In the Camel the dental formula is _i_ 1/3 _c_ 1/1 +_pm_ 3/2 _m_ 3/3, total 34. The upper incisors, canines and first +premolars of the Camel are very small teeth, and the first premolar +is separated by a long diastema from the others. + +The Tragulina or Chevrotains have no upper incisors, while the canines +are largely developed, especially in the male. + +The Ruminantia or Pecora are very uniform as regards their dentition. +The upper incisors are always absent, for though their germs are +developed they are reabsorbed without ever becoming visible, and as a +rule the upper canines are absent too, while the lower canines are +incisiform. The grinding teeth are typically selenodont, and in the +lower jaw form a continuous series separated by a wide diastema from +the canines. The dental formula is usually + + _i_ 0/3 _c_ 0-1/1 _pm_ 3/3 _m_ 3/3. + +The canines are largely developed in the male Muskdeer (_Moschus_) and +in _Hydropotes_. + +_PERISSODACTYLA_. The premolars and molars have a very similar +structure and form a continuous series of large square teeth with +complex crowns. The crowns are always constructed on some modification +of the bilophodont[159] plan, as is easily seen in the case of the +forms with brachydont teeth, but in animals like the Horse, in which +the teeth are very hypsodont, this arrangement is hard to trace. All +four premolars in the upper jaw are preceded by milk teeth, while in +Artiodactyla the first has no milk predecessors. + +In the Tapiridae the grinding teeth are brachydont and the lower ones +are typically bilophodont. The last two upper molars have the +transverse ridges united by an outer longitudinal ridge. The dentition +is _i_ 3/3 _c_ 1/1 _pm_ 4/3 _m_ 3/3, total 42. + +In some of the extinct Perissodactyles such as _Lophiodon_[160], the +dentition is brachydont and bilophodont, the grinding teeth in general +resembling the posterior upper molars of the Tapir. The same type of +brachydont tooth is seen in _Palaeotherium_ but the transverse ridges +are crescentic instead of straight, and are separated from one another +by shallow valleys without cement. Some of the Palaeotheridae have the +regular series of forty-four teeth. + +A complete series of forms is known showing how from the simple +brachydont teeth of the Palaeotheridae, were derived the complicated +hypsodont teeth of the Equidae. The increase in depth of the tooth was +accompanied by increase in the depth and complexity of the enamel +infoldings, and of the cement filling them. + +Both upper and lower grinding teeth of the Equidae are much +complicated by enamel infoldings, but their derivation from the +bilophodont type can still be recognised. The diastema in front of the +premolars is longer in the living Equidae than in their extinct +allies. In the adult horse the dental formula is _i_ 3/3 _c_ 1/1 _pm_ +3/3 _m_ 3/3, total 40, with often a vestigial first upper premolar +(fig. 82, _pm_ 1). The last molar is not more complex than the others, +and in the female the canine is quite vestigial. The incisors are +large and adapted for cutting and have the enamel curiously folded in +forming a deep pit. The milk dentition is _di_ 3/3 _dc_ 0/0 _dpm_ 3/3, +total 24. The last milk premolar is not more complex than the premolar +that succeeds it. The horse affords an excellent instance of a +typically _herbivorous_ type of dentition, the cutting incisors, +reduced canines and series of large square flat-crowned grinding teeth +being most characteristic. + +In _Rhinoceros_ the grinding teeth are much like those of _Lophiodon_, +having an outer longitudinal ridge from which two crescentic +transverse ridges diverge. The upper premolars are as complex as the +molars, and there are no canines; in some species incisors also are +absent. The dental formula is + + _i_ (0--2)/(0--1) _c_ 0/(0--1) _pm_ 4/4 _m_ 3/3. + +[Illustration FIG. 82. PALATAL ASPECT OF THE CRANIUM AND MANDIBLE OF A +DONKEY (_Equus asinus_) × 1/5. (Camb. Mus.) + + 1. supra-occipital. + 2. occipital condyle. + 3. basi-occipital. + 4. vacuity representing the confluent foramen lacerum posterius + and foramen lacerum medium. + 5. auditory bulla. + 6. glenoid surface. + 7. vomer. + _i_ 1, _i_ 3. first and third incisors. + _c._ canine. + _pm_ 1, _pm_ 2. first and second premolars. + _m_ 1. first molar.] + +Among the Titanotheriidae _Palaeosyops_[161] has very brachydont teeth +whose crowns have been described as _buno-selenodont_, the inner pair +of columns being bunodont, the outer, selenodont. Similar grinding +teeth occur in _Chalicotherium_. Some of the Titanotheriidae have the +regular mammalian series of forty-four teeth. + + +SUBUNGULATA. + +_TOXODONTIA._ _Nesodon_ has the regular dental formula; its grinding +teeth are rooted and the upper ones resemble those of Rhinoceros. The +second upper and third lower incisors form ever-growing tusks. There +is a marked difference between the deciduous and permanent dentition. +_Astrapotherium_ likewise has large rooted cheek teeth of a +rhinocerotic type, and each jaw bears a pair of permanently growing +tusks, those of the lower jaw being the canines. The dental formula is + + _i_ 1/3 _c_ 0/1 _pm_ 2/1 _m_ 3/3, total 28. + +In _Toxodon_ the upper incisors and molars are large and curved and +all the teeth have persistent pulps. In _Typotherium_ there are no +tusks, but the upper incisors are chisel-like, recalling those of +Rodents. + +The _CONDYLARTHRA_ have brachydont, generally bunodont teeth, with the +premolars simpler than the molars. They generally have the regular +dental formula. + +_HYRACOIDEA._ The dental formula of _Procavia_ is usually given as _i_ +1/2 _c_ 0/0 _pm_ 4/4 _m_ 3/3, total 34; in young individuals however +there occur a second pair of upper incisors which early fall out. The +upper incisors resemble those of Rodents in being long and curved and +growing from persistent pulps. They are however triangular in +transverse section, not rectangular, having two antero-lateral faces +covered with enamel and a posterior face without enamel. Their +terminations are pointed, not chisel-shaped as in Rodents. The lower +incisors (fig. 83, _i_ 1) are pectinate or partially divided by +vertical fissures, and the grinding teeth are of the rhinocerotic +type. + +[Illustration FIG. 83. SKULL OF _Procavia (Dendrohyrax) dorsalis_ × +2/3. (Camb. Mus.) + + 1. nasal. + 2. parietal. + 3. external auditory meatus. + 4. paroccipital process of the exoccipital. + 5. jugal. + 6. lachrymal foramen. + _i_ 1. first incisor.] + +_AMBLYPODA._ Two of the best known forms belonging to this extinct +group differ much as regards dentition. For while _Coryphodon_ has the +regular dental formula, and the canines of both jaws of moderate size, +in _Uintatherium_ the dentition is very specialised, there are no +upper incisors, and the upper canines form a pair of enormous tusks. +The grinding teeth form a continuous series marked by =V=-shaped ridges +and the dental formula is _i_ 0/3 _c_ 1/1 _pm_ 3/3 _m_ 3/3 total 34. + +_PROBOSCIDEA._ The incisors are composed entirely of dentine and have +the form of conical tusks projecting greatly from the mouth. In +living forms they are confined to the upper jaw, in some species of +the extinct _Mastodon_ however they occur in the lower jaw also. In +_Dinotherium_ they are probably absent from the upper jaw, but form a +pair of downwardly and backwardly-directed tusks growing from the +elongated symphysis of the mandible. + +The grinding teeth in the various Proboscidea show a very remarkable +series of modifications. In _Dinotherium_ they are bilophodont or else +are marked by three straight transverse ridges. The dental formula is +_i_ 0?/1 _c_ 0/0 _pm_ 2/2 _m_ 3/3, and the teeth have the normal +method of succession. In _Mastodon_ as in _Dinotherium_ the grinding +teeth are marked by transverse ridges, but the ridges are subdivided +into conical or mammillary cusps, and similar cusps often occur +between the ridges. These cusps are covered with very thick enamel and +the spaces between them are not filled up with cement. There are six +of these grinding teeth for each side of each jaw but only three are +in place at once. The first three are milk teeth as they may be +succeeded vertically by others. + +In the true Elephants the number and depth of the enamel folds is much +increased, and the spaces between the folds are filled up with cement. +A very complete series of extinct forms is known with teeth +intermediate in character between those of _Mastodon_ and those of the +Mammoth and living elephants. The dental formula of _Elephas_ is + + _di_ 1/0 _i_ 1/0 _c_ 0/0 _dm_ 3--4/3--4 _m_ 3/3. + +Sir W.H. Flower describes[162] the mode of succession of teeth in +Elephants as follows: "As regards the mode of succession that of +modern Elephants is as before mentioned very peculiar. During the +complete lifetime of the animal there are but six molar teeth on each +side of each jaw with occasionally a rudimentary one in front, +completing the typical number of seven. The last three represent the +true molars of ordinary mammals, those in front appear to be milk +molars which are never replaced by permanent successors, but the whole +series gradually moves forwards in the jaw, and the teeth become worn +away and their remnants cast out in front while development of others +proceeds behind. The individual teeth are so large and the processes +of growth and destruction by wear take place so slowly, that not more +than one or portions of two teeth are ever in place and in use on each +side of each jaw at one time, and the whole series of changes +coincides with the usual duration of the animal's life. On the other +hand the _Dinotherium_, the opposite extreme of the Proboscidean +series, has the whole of the molar teeth in place and use at one time, +and the milk molars are vertically displaced by premolars in the +ordinary fashion. Among Mastodons transitional forms occur in the mode +of succession as well as in structure, many species showing a vertical +displacement of one or more of the milk molars, and the same has been +observed in one extinct species of Elephant (_E. planifrons_) as +regards the posterior of these teeth." + +In the TILLODONTIA the grinding teeth are of Ungulate type, while the +second incisors are large and grow from persistent pulps, so as to +resemble those of Rodents. + +RODENTIA have a most characteristic and very constant dentition, the +common dental formula being + + _i_ 1/1 _c_ 0/0 _pm_ (0--1)/(0--1) _m_ 3/3, total 18 or 20. + +The incisors always have chisel-like edges and persistent pulps, and +are separated by a wide diastema from the premolars. Canines are +always absent, and there are generally three grinding teeth not +preceded by milk teeth; their surface may be grooved, or may be +bunodont. Teeth are most numerous in the Duplicidentata (Hares and +Rabbits), in which the formula is _i_ 2/1 _c_ 0/0 _pm_ 3/2 _m_ 3/3, +total 28, and fewest in Hydromys and certain other forms, in which +the formula is _i_ 1/1 _c_ 0/0 _pm_ 0/0 _m_ 2/2, total 12. The hares +and rabbits are the only rodents which have well developed deciduous +incisors, though a vestigial milk incisor has been described in the +Mouse (_Mus musculus_). The last upper molar of _Hydrochaerus_ is very +complicated, its structure approaching that of the teeth of Elephants. + +[Illustration FIG. 84. CARNASSIAL OR SECTORIAL TEETH OF CARNIVORA +(from FLOWER). + +_Upper sectorial teeth_ of I. _Felis_, II. _Canis_, III. _Ursus_. 1. +anterior, 2. middle, 3. posterior cusp of blade, 4. inner lobe +supported on distinct root, 5. inner lobe posterior in position and +without distinct root, characteristic of the Ursidae. + +_Lower sectorial teeth._ 1. _Felis_, 2. _Canis_, 3. _Herpestes_. 1. +anterior, 2. posterior lobe of blade, 3. inner tubercle, 4. heel.] + +CARNIVORA have the teeth rooted and markedly diphyodont and +heterodont. The canines are greatly developed, and the incisors are +small. + +In _CARNIVORA VERA_ the incisors are almost always 3/3. The fourth +upper premolar and first lower molar are differentiated as carnassial +teeth (see p. 436), and retain fundamentally the same characters +throughout the suborder. The upper carnassial (fig. 84, I. II. III.) +consists of a more or less compressed, commonly trilobed blade borne +on two roots, with an inner tubercle borne on a third root. The lower +carnassial has only two roots; its crown consists of a bilobed blade +with generally an inner cusp, and a heel or talon (fig. 84, 4) behind +the blade. + +The most thoroughly carnivorous type of dentition is seen in the +Æluroidea, and especially in the cat tribe (Felidae). In the genus +_Felis_ the dental formula is _i_ 3/3 _c_ 1/1 _pm_ 3/2 _m_ 1/1, total +30. The incisors are very small, so as not to interfere with the +action of the large canines, the lower carnassial is reduced to simply +the bilobed blade (fig. 84, IV), and the cheek teeth are greatly +subordinated to the carnassial. The extinct _Machaerodus_ has the +upper canines comparable in size to those of the Walrus. + +The Civets and Hyaenas have a dentition allying them closely to the +cats. The hyaena-like _Proteles_ has, however, the grinding teeth +greatly reduced. + +In the Cynoidea[163] the general dentition is _i_ 3/3 _c_ 1/1 _pm_ 4/4 +_m_ 2/3, total 42. This differs from the regular mammalian dentition +only in the absence of the last upper molar. The upper carnassial +tooth (fig. 84, II.) consists of a larger middle and smaller posterior +lobe with hardly any trace of an anterior lobe. The lower carnassial +(fig. 84, V.) is typical, consisting of a bilobed blade with inner +cusp and posterior talon. + +The dentition of the Cynoidea is most closely linked with that of the +Arctoidea by means of fossil forms. + +[Illustration FIG. 85. MANDIBLE OF ISABELLINE BEAR (_Ursus +isabellinus_) × 1/2. (Camb. Mus.) + + 1. condyle. + 2. coronoid process. + _i_ 1. first incisor. + _c._ canine. + _pm_ 1, _pm_ 2. first and second premolars. + _m_ 1. first molar. The dotted + line is pointing to the posterior + half of the tooth. + This specimen has only + three premolars, there + should be four.] + +In the Arctoidea the dentition is not so typically carnivorous as in +the Æluroidea and Cynoidea. In the bears, Ursidae, the molars have +broad flat tuberculated crowns (fig. 85). The dental formula in +_Ursus_ is _i_ 3/3 _c_ 1/1 _pm_ 4/4 _m_ 2/3, total 42. The upper +carnassial (fig. 84, III.) differs from that of the Æluroidea and +Cynoidea in having no inner lobe supported on a third root. In the +large group of Mustelidae there are generally two molars in the lower +and one in the upper jaw. The grinding teeth commonly have large, +flattened, more or less tuberculated crowns, and the upper molar may +be as large or much smaller than the carnassial. + +In the _CREODONTA_ there are no specially differentiated carnassial +teeth. + +[Illustration FIG. 86. LEFT MANDIBULAR RAMUS OF THE SEA LEOPARD +(_Ogmorhinus leptonyx_) WITH THE ROOTS OF THE TEETH EXPOSED × 1/3. +(Camb. Mus.) + + 1. condyle. + 2. coronoid process. + _i_ 3. third incisor. + _c._ canine. + _pm_ 1, _pm_ 4. first and fourth premolars. + _m._ molar.] + +In the _PINNIPEDIA_ the dentition differs considerably from that of +the Carnivora vera. The milk dentition is always vestigial, and the +teeth are frequently absorbed before birth. There are four premolars +and one molar, forming an uniform series of cheek teeth, all of which +except in the Walrus have compressed and pointed, never flattened, +crowns. There is no special carnassial tooth, and the incisors are +always fewer than 3/3. In _Otaria_ the dentition is + + _i_ 3/2 _c_ 1/1 _pm_ 4/4 _m_ 1 or 2/1, total 34 or 36. + +In the Walrus the upper canines form immense tusks. The other teeth +are all small and one-rooted, and the molars have flat crowns. In the +true seals the dentition is strikingly piscivorous, the cheek teeth +often having accessory cusps (fig. 86). + +The INSECTIVORA are diphyodont and heterodont, having well-developed +rooted teeth. The canines are usually weak, the incisors pointed, and +those of the two jaws often meet like a pair of forceps. The crowns of +the molars are characteristically studded with short cusps. Some +genera, such as _Gymnura_ and the mole, _Talpa_, have the regular +mammalian dentition. In the hedgehog, _Erinaceus_, the dentition is + + _i_ 3/2 _c_ 1/1 _pm_ 3/2 _m_ 3/3, total 36. + +In the genus _Sorex_ (Shrews) the teeth differ in the following two +marked respects from those of most other Monodelphia, (1) they are +monophyodont, (2) the lower incisors sometimes become fused to the +jaws. Most Insectivora have square molar teeth, but in _Potamogale_, +_Chrysochloris_, _Solenodon_ and the Centetidae the molar teeth are +triangular in section. Four molars occur in _Centetes_. + +In the aberrant genus _Galeopithecus_ the dentition is _i_ 2/3 _c_ 1/1 +_pm_ 2/2 _m_ 3/3, total 34. The upper incisors are placed at some +distance from the anterior end of the jaw, and the outer upper +incisors and canines of both jaws have two roots,--a very unusual +character. The lower incisors are deeply grooved or pectinated in the +same way as are the lower incisors of _Procavia_. The upper incisors +and canines of both jaws bear many cusps, and are very similar in +appearance to the cheek teeth of some Seals. + +The dentition of the CHIROPTERA is diphyodont and heterodont, and the +dental formula never exceeds + + _i_ 2/3 _c_ 1/1 pm 3/3 _m_ 3/3, total 38. + +The milk teeth are very slender and have sharp recurved cusps; they +are quite unlike the permanent teeth. The permanent teeth are of two +types. In the Insectivorous forms the molar teeth are cusped, and +resemble those of Insectivora. In the blood-sucking Vampire bat +_Desmodus_, the teeth are peculiarly modified; the canines and the +single pair of upper incisors are much enlarged and exceedingly sharp, +while all the other teeth are much reduced in size. + +In the Frugivorous bats the molar teeth have nearly always smooth +crowns. The dental formula in the chief genus _Pteropus_ is _i_ 2/2 +_c_ 1/1 _pm_ 3/3 _m_ 2/3, total 34. + +The PRIMATES have a diphyodont and heterodont dentition, generally of +an omnivorous type, with cheek teeth adapted for grinding. The +incisors are generally 2/2, and the molars, except in the Hapalidae, +are 3/3. In the Lemurs the upper canines are large, and the lower +incisors slender and directed almost horizontally forwards. The Aye +Aye, _Chiromys_, has the following singular dentition: _i_ 1/1 _c_ 0/0 +_pm_ 1/0 _m_ 3/3, total 18. The incisors much resemble those of +rodents having persistent pulps, and enamel only on the anterior face. + +In Man and in the Anthropoid and Old World Apes the dental formula is +always _i_ 2/2 _c_ 1/1 _pm_ 2/2 _m_ 3/3, total 32. + +In the Cebidae there is an extra premolar in each jaw bringing the +number up to 36. In the Hapalidae, as in the Cebidae, there is a third +premolar, but the molars are reduced to 2/2. Man is the only Primate +that has the teeth arranged in a continuous series. In all the others +there is a gap or diastema of larger or smaller size between the +incisors and canines. In all except man also the canines are enlarged, +especially in the males. + +The Exoskeletal structures of mammals may be summarised in the +following table: + + I. _Epidermal exoskeletal structures._ + + 1. Hairs (_a_) ordinary hair, + (_b_) vibrissae and bristles, + (_c_) spines of hedgehog, porcupine, _Echidna_, + _Centetes_, _Acanthomys_. + 2. Scales { of Manidae, + { on tails of rats, beavers, &c. + 3. Horns of Rhinoceros. + 4. Horns of Bovine Ruminants. + 5. Nails, claws, hoofs. + 6. Spurs of male _Ornithorhynchus_ and _Echidna_. + 7. Horny beak and teeth of _Ornithorhynchus_. + 8. Horny pads on jaws of Sirenians and Ruminants. + 9. Baleen of whales. + 10. Enamel of teeth. + + II. _Dermal exoskeletal structures._ + + 1. Dentine and cement of teeth. + 2. Bony scutes of Armadillos. + + +ENDOSKELETON. + +VERTEBRAL COLUMN. + +CERVICAL VERTEBRAE. + +The cervical vertebrae of all mammals have certain characters in +common. However long the neck may be, the number of cervical +vertebrae, with very few exceptions, is seven. Movable ribs are +generally absent, and if present are small and do not reach the +sternum. The transverse processes are generally wide but not long, and +are perforated near the base by the vertebrarterial canals, through +which the vertebral arteries pass; they generally bear downwardly +directed inferior lamellae which are sometimes as in the seventh human +cervical seen to ossify from centres distinct from those forming the +rest of the transverse process, and are really of the nature of ribs. +The atlas and axis always differ much from the other vertebrae. + +We may pass now to the special characters of the cervical vertebrae in +the different groups. In MONOTREMES and MARSUPIALS the number of +cervical vertebrae is always seven. With the exception of the atlas of +_Echidna_ the cervical vertebrae of Monotremes are without +zygapophyses. In Monotremes the transverse processes ossify from +centres distinct from that forming the body, and remain suturally +connected with the rest of the vertebra until the adult condition is +reached. The method of the ossification of the atlas in Marsupials +varies considerably, thus in some forms such as the Wombats +(_Phascolomys_) there is an unossified gap in the middle of the +inferior arch of the atlas, which may remain permanently open; in +_Thylacinus_ this gap is filled up by a distinct heart-shaped piece of +bone, while in _Didelphys_ and _Perameles_ the atlas is ossified below +in the same way as in other mammals. In _Notoryctes_ the second to +sixth cervical vertebrae are ankylosed together. + +The cervical vertebrae of the EDENTATA have some remarkable +peculiarities. In the three-fingered Sloth, _Bradypus_, there are nine +cervical vertebrae, all except the last of which have their transverse +processes perforated by the vertebrarterial canals. In a two-fingered +sloth, _Choloepus hoffmanni_, there are only six cervical vertebrae. +In the Megatheriidae, Anteaters (Myrmecophagidae), Pangolins +(Manidae), and Aard Varks (Orycteropodidae), the cervical vertebrae +are normal, but in the Armadillos (Dasypodidae), and still more in the +Glyptodonts, several of them are commonly fused together. The fusion +affects not only the centra, but also the neural arches, so that the +neural canals form a continuous tube. + +In the Glyptodonts there is a complex joint at the base of the neck to +allow the partial retraction of the head within the carapace. This +arrangement recalls that in Tortoises. + +As a rule the SIRENIA possess seven short cervical vertebrae, not +fused together and not presenting any marked peculiarities. In +_Manatus_ however there are only six cervical vertebrae and they are +very variable. + +[Illustration FIG. 87. CERVICAL VERTEBRAE OF A YOUNG FIN WHALE +(_Balaenoptera musculus_) × 1/10. (Camb. Mus.) + + 1. surface on the atlas for articulation + with the occipital condyle of the skull. + 2. foramen for exit of the first spinal nerve. + 3. upper transverse process. + 4. lower transverse process. + +In the fresh specimen these two transverse processes are united by +cartilage, in adult individuals the whole transverse process is +ossified. + + 5. epiphyses of centrum. + 6. neural spine.] + +In the CETACEA there are invariably seven cervical vertebrae, but they +are always very short and are frequently even before birth fused +together by their centra into one continuous mass (see fig. 67). +Sometimes the last one or two are free. In the Rorquals +(_Balaenoptera_) however, the cervical vertebrae are quite separate +and distinct (fig. 87), and in the fluviatile Odontoceti, +_Platanista_, _Inia_, and _Pontoporia_, and also in _Beluga_ and +_Monodon_, though very short they are free. In _Physeter_ the first +vertebra is free while the others are fused. An odontoid process is +not commonly present even in Cetaceans with free cervical vertebrae, +but a very short one occurs in the Rorquals. The cervical vertebrae of +Rorquals give off on each side two transverse processes (fig. 87, 3 +and 4) which enclose between them a wide space. These processes are +not completely ossified till the animal is adult. + +[Illustration FIG. 88. ATLAS (B) AND AXIS (A) VERTEBRAE OF AN OX (_Bos +taurus_) × 1/4. (Camb. Mus.) + + 1. neural canal. + 2. transverse process. + 3. surfaces for articulation with the occipital condyles of the skull. + 4. spout-like odontoid process. + 5. hypapophysis. + 6. anterior opening of the vertebrarterial canal. + 7. foramen for the exit of the second spinal nerve. + 8. neural spine. + 9. postzygapophysis.] + +In all UNGULATA the number of cervical vertebrae is seven. Among the +Artiodactyla two forms of the odontoid process of the axis occurs; in +the Suina and Tragulina it is conical, in the Ruminantia and Tylopoda +it is spout-like (fig. 88, 4). The atlas in the Suina and to a less +extent in the Ruminantia has long flattened transverse processes, and +the remaining cervical vertebrae are opisthocoelous. Those of the +Giraffe and Llama (fig. 103) are noticeable for their great length. In +the Tylopoda the posterior half of the vertebrarterial canal is +confluent with the neural canal. + +The Perissodactyla have remarkably opisthocoelous cervical vertebrae. +Those of _Macrauchenia_ have the posterior half of the vertebrarterial +canal confluent with the neural canal as in Tylopoda. In the +Proboscidea they are short flattened discs slightly opisthocoelous; +the axis and seventh vertebra and to a less extent the sixth have high +neural spines. + +In the RODENTIA the atlas generally has broad wing-like transverse +processes, and the axis a large and long neural spine, while the +odontoid process is much developed. In the Jerboas (_Dipus_) all the +cervical vertebrae except the atlas are fused together, a condition +recalling that in armadillos. + +In the CARNIVORA the wings of the atlas are well developed (fig. 69, +A, 1), and it is deeply cupped for articulation with the condyles of +the skull. The axis has a long odontoid process and a high compressed +neural spine (fig. 69, B, 4). The third to sixth cervical vertebrae +have large transverse processes with prominent perforated inferior +lamellae, whose ventral margins in the third and fourth vertebrae +diverge as they pass backwards, while in the fifth they are parallel +and in the sixth convergent. The transverse processes of the seventh +vertebra have no inferior lamellae and are not perforated. +Metapophyses are often developed. + +In the INSECTIVORA the cervical vertebrae vary considerably. The +neural spines except in the case of the axis are generally very small +and in the Shrews and Moles the neural arches are exceedingly slender. + +In the CHIROPTERA all the cervical vertebrae are broad and short with +slender neural arches. + +PRIMATES. In Man the cervical vertebrae have short blunt transverse +processes and small often bifid neural spines. The neural and +vertebrarterial canals are large. The atlas forms a ring surrounding a +large cavity, and has a very slender inferior arch and small +transverse processes. Traces of a pro-atlas have been described in +_Macacus_ and _Cynocephalus_. The axis has a prominent spine and +odontoid process and short transverse processes. In most Primates the +cervical vertebrae are very similar to those of man, but the inferior +lamellae of the transverse processes are better developed. In the +Anthropoid Apes the neural spines are as a rule much elongated. + + +THORACO-LUMBAR, OR TRUNK VERTEBRAE. + +In the MONOTREMATA there are nineteen thoraco-lumbar vertebrae, +sixteen (_Echidna_) or seventeen (_Ornithorhynchus_) of which bear +ribs. The transverse processes are very short and do not articulate +with the ribs, which are united to the centra only. + +In the MARSUPIALIA there are always nineteen thoraco-lumbar vertebrae, +thirteen of which generally bear ribs. The lumbar vertebrae frequently +have large metapophyses and anapophyses, these being specially well +seen in the Kangaroos and Koala (_Phascolarctus_). + +The EDENTATA are very variable as regards their trunk vertebrae. The +two genera of Sloths differ much as regards the number, for while +_Bradypus_ has only nineteen, fifteen or sixteen of which bear ribs, +_Choloepus_ has twenty-seven, twenty-four of which are thoracic, and +bear ribs. In _Bradypus_ a small outgrowth from the transverse process +articulates with the neural arch of the succeeding vertebra. In both +genera the neural spines are all directed backwards. + +In the Megatheriidae as in the sloths the neural spines are all +directed backwards, and in the lumbar region additional articulating +surfaces occur, better developed than are those in _Bradypus_. + +In the ant-eaters (Myrmecophagidae) there are seventeen or eighteen +thoraco-lumbar vertebrae, all of which except two or three bear ribs. +The posterior thoracic and anterior lumbar vertebrae articulate in a +very complex fashion, second, third, and fourth pairs of zygapophyses +being progressively developed in addition to the ordinary ones, as the +vertebrae are followed back. + +In the Armadillos the lumbar vertebrae have long metapophyses which +project upwards and forwards and help to support the carapace. In +_Glyptodon_ almost all the thoraco-lumbar vertebrae are completely +ankylosed together. + +In the Manidae there are no additional zygapophyses but the normal +ones of the lumbar and posterior thoracic regions are very much +developed, the postzygapophyses being semi-cylindrical and fitting +into the deep prezygapophyses of the succeeding vertebra. + +In the SIRENIA the number of lumbar vertebrae is very small; in the +dugong there are nineteen thoracic and four lumbar, and in the manatee +seventeen thoracic and two lumbar. + +In the CETACEA the number of thoracic vertebrae varies from nine in +_Hyperoödon_ to fifteen or sixteen in _Balaenoptera_, and the number +of lumbar vertebrae from three in _Inia_ to twenty-four or more in +_Delphinus_. The lumbar vertebrae are often very loosely articulated +together and the zygapophyses sometimes as in the Dolphins are placed +high up on the neural spines. The centra are large, short in the +anterior region but becoming longer behind. The epiphyses are +prominent, and so are the neural spines and to a less extent the +metapophyses. The transverse processes are well developed, anteriorly +they arise high up on the neural arch, but when the vertebral column +is followed back they come gradually to be placed lower down, till in +the lumbar region they project from the middle of the centra. This can +be well traced in the Porpoise (_Phocaena_). In the Physeteridae the +transverse processes of the anterior thoracic vertebrae are similar to +those of most Cetacea, but when followed back, instead of shifting +their position on the vertebrae, they gradually disappear, and other +processes gradually arise from the point where the capitulum of the +rib articulates. + +UNGULATA. In the Ungulata vera the thoraco-lumbar vertebrae are +slightly opisthocoelous. The anterior thoracic vertebrae commonly have +exceedingly high backwardly-projecting neural spines (fig. 89, 1); but +those of the lumbar and posterior thoracic vertebrae often point +somewhat forwards so that the spines all converge somewhat to a point +called the _centre of motion_ (cp. fig. 101). In the Artiodactyla +there are always nineteen thoraco-lumbar vertebrae, and in the +Perissodactyla twenty-three. + +_Procavia_ sometimes has thirty thoraco-lumbar vertebrae, a greater +number than occurs in any other terrestrial mammal; twenty-two of +these are thoracic and eight lumbar. In _Phenacodus_ the convergence +of the neural spines to a centre of motion is well seen. + +[Illustration FIG. 89. FIRST AND SECOND THORACIC VERTEBRAE OF AN OX +(_Bos taurus_) × 1/3. (Camb. Mus.) + + 1. neural spine. + 2. neural canal. + 3. prezygapophysis. + 4. facet for articulation with the tuberculum of the rib. + 5. facet for articulation with the capitulum of the rib. + 6. postzygapophysis. + 7. foramen for exit of spinal nerve.] + +In the Proboscidea there are twenty-three thoraco-lumbar vertebrae, of +which nineteen or twenty bear ribs. + +In the RODENTIA there are generally nineteen thoraco-lumbar vertebrae +but occasionally the number rises as high as twenty-five. In the Hares +(Leporidae) the number is nineteen, twelve or thirteen of which are +thoracic. The anterior thoracic vertebrae have short centra and high +backwardly-directed neural spines, the lumbar vertebrae have large +forwardly-and downwardly-directed transverse processes with expanded +ends. Metapophyses, anapophyses and hypapophyses are all present. In +the Agouti (_Dasyprocta_) the convergence of the neural spines to a +centre of motion is very strongly marked. + +In the CARNIVORA the trunk vertebrae are nearly always twenty or +twenty-one in number; in the genera _Felis_ and _Canis_ thirteen of +these are thoracic and seven lumbar. The anterior thoracic vertebrae +have long backwardly-projecting neural spines, while the posterior +thoracic and lumbar vertebrae have shorter and thicker neural spines +which project slightly forwards. In the Pinnipedia there is no change +in the direction of the neural spines, and anapophyses are but little +developed. + +In the INSECTIVORA the number of trunk vertebrae varies much from +nineteen--thirteen thoracic and six lumbar--in _Tupaia_, to +twenty-four--nineteen thoracic and five lumbar--in _Centetes_. The +development of the various processes varies in accordance with the +habits of the animals, being great in the active forms, slight in the +slowly moving or burrowing forms. In _Talpa_ and _Galeopithecus_ the +intervertebral discs of the thoraco-lumbar region instead of being +cartilaginous have ossified forming inter centra, a condition met with +in very few mammals. + +In the CHIROPTERA there are seventeen or eighteen thoraco-lumbar +vertebrae, eleven to fourteen of which may bear ribs. The development +of processes is slight. + +Among PRIMATES the number of trunk vertebrae is generally nineteen, of +which twelve to fourteen bear ribs; in man and the Gorilla and +Chimpanzee the number is, however, seventeen, and in the Orang +(_Simia_) sixteen. In some of the Lemuroidea there are as many as +twenty-three or twenty-four. In most cases the neural spines converge +more or less to a centre of motion, and this is especially marked in +some of the Lemurs; it does not occur in man and the anthropoid apes. + + +SACRAL AND CAUDAL VERTEBRAE. + +At the posterior end of the trunk in all mammals a certain number of +vertebrae are found fused together forming the sacrum. But of these +only two or three answer to the definition of true sacral vertebrae in +being united to the ilia by small ribs. The others which belong to the +caudal series may be called pseudosacral vertebrae. In different +individuals of the same species it sometimes happens that different +vertebrae are attached to the pelvis and form the sacrum. Sometimes +even different vertebrae are attached to the pelvis at successive +periods in the life history of the individual. This is owing to a +shifting of the pelvis and has been especially well seen in man. In +young human embryos the pelvis is at a certain stage attached to +vertebra 30, but as development goes on it becomes progressively +attached to the twenty-ninth, twenty-eighth, twenty-seventh, +twenty-sixth and twenty-fifth vertebrae. As the attachment to these +anterior vertebrae is gained, the attachment to the posterior ones +becomes lost, so that in the adult the pelvis is generally attached to +vertebrae 25 and 26. But there are no absolutely pre-determined sacral +vertebrae, as sometimes the pelvis does not reach vertebra 25, +remaining attached to vertebrae 26 and 27; sometimes it becomes +attached even to vertebra 24. This shifting of the pelvis is seen in +_Choloepus_ in a more marked degree even than in man. + +Of the MONOTREMATA, _Ornithorhynchus_ has two sacral vertebrae +ankylosed together, while _Echidna_ has three or four[164]. + +In MARSUPIALIA as a rule only one vertebra is directly united to the +ilia, but one or two more are commonly fused to the first. In the +Wombats there may be as many as four or five vertebrae fused together +in the sacral region. In _Notoryctes_ there is extensive fusion in the +sacral region, six vertebrae, owing mainly to the great development of +their metapophyses, being united with one another, and with the ilia, +and the greater part of the ischia. + +In most EDENTATA there is an extensive fusion of vertebrae in the +sacral region. This is especially marked in the Armadillos and +Megatheriidae, and to a less extent in the Sloths and Aard Varks. + +In the SIRENIA the vestigial pelvis is attached by ligament to the +transverse processes of a single vertebra, which hence may be regarded +as sacral. + +In CETACEA there is no sacrum, the vestigial pelvis not being +connected with the vertebral column. + +In most UNGULATA the sacrum consists of one large vertebra united to +the ilia, and having a varying number of smaller vertebrae fused with +it behind. + +The same arrangement obtains in most RODENTIA, but in the Beavers +(Castoridae) all the fused vertebrae are of much the same size, the +posterior ones having long transverse processes which nearly meet the +ilia. + +In CARNIVORA there may be two sacral vertebrae as in the Hyaena, three +as in the Dog, four or five as in Bears and Seals. + +In INSECTIVORA from three to five are united, while in many CHIROPTERA +all the sacral and caudal vertebrae have coalesced. Among PRIMATES, in +Man and Anthropoid Apes there are usually five fused vertebrae forming +the sacrum, but of these only two or three are connected to the ilia +by ribs. In most of the other Anthropoidea there are two or three +fused vertebrae, and in the Lemuroidea two to five. + +FREE CAUDAL VERTEBRAE. The free caudal vertebrae vary greatly in +number and character. When the tail is well developed, the anterior +vertebrae are comparatively short and broad, with well-developed +neural arches and zygapophyses; but as the tail is followed back, the +centra gradually lengthen and become cylindrical, and at the same time +the neural arches and all the processes gradually become reduced and +disappear, so that the last few vertebrae consist of simple rod-like +centra. Chevron bones are frequently well-developed. + +Of the MONOTREMES _Echidna_ has twelve caudal vertebrae, two of which +bear irregular chevron bones. In _Ornithorhynchus_ there are twenty or +twenty-one caudal vertebrae with well-developed hypapophyses, but no +chevron bones. + +In MARSUPIALS there is great diversity as regards the tail. In the +Wombat and Koala the tail is small and without chevron bones. In most +other Marsupials it is very long, having sometimes as many as +thirty-five vertebrae in the prehensile-tailed opossums. In the +Kangaroos the tail is very large and stout. Chevron bones are almost +always present, and in _Notoryctes_ are large and expanded. + +Most EDENTATES have large tails with well-developed chevron bones. The +length of the tail varies greatly from the rudimentary condition in +Sloths to that in the Pangolins, one of which has forty-six to +forty-nine caudal vertebrae--the largest number in any known mammal. +Chevron bones are much developed, sometimes they are Y-shaped, +sometimes as in _Priodon_, they have strong diverging processes. The +caudal vertebrae of Glyptodonts, though enclosed in a continuous bony +sheath, have not become ankylosed together. + +The SIRENIA have numerous caudal vertebrae with wide transverse +processes. In the CETACEA also the tail is much developed, and the +anterior vertebrae have large chevron bones and prominent straight +transverse processes; the posterior caudal vertebrae, which in life +are enclosed in the horizontally expanded tail fin, are without +transverse processes. + +In UNGULATA the tail is simple, formed of short cylindrical vertebrae, +which in living forms are never provided with chevron bones. The +number of caudal vertebrae varies from four, sometimes met with in +_Procavia_, to thirty-one in the Elephant. The tail is exceedingly +long in _Anoplotherium_ and in _Phenacodus_, in which there are thirty +caudal vertebrae. + +In RODENTIA the tail is variable. In the Hares, Guinea pig (_Cavia_) +and _Capybara_ it is very small, in _Pedetes_ and the Beaver it is +very long and has well-developed chevron bones. + +Most of the CARNIVORA except the Bears and Seals have very long tails, +the greatest number of vertebrae, thirty-six, being met with in +_Paradoxurus_. Bears have only eight to ten caudal vertebrae. Chevron +bones are not often much developed. + +In INSECTIVORA the tail is very variable as regards length, the number +of vertebrae varying from eight in _Centetes_ to forty-three in +_Microgale_. + +In CHIROPTERA the tail is sometimes quite rudimentary, and as in +_Pteropus_, composed of a few coalesced vertebrae, sometimes it is +formed of a large number of slender vertebrae. + +In PRIMATES also the tail is very variable. In Man all the four caudal +vertebrae are rudimentary and are fused together, forming the +_coccyx_. In the Anthropoid apes, too, there are only four or five +caudal vertebrae. In many monkeys of both the eastern and western +hemispheres the tail is very long, having thirty-three vertebrae in +_Ateles_, in which genus it is also prehensile. Chevron bones are +present in all Primates with well-developed tails. In the Lemuroidea +the number of caudal vertebrae varies from seven to twenty-nine. + + +FOOTNOTES: + +[145] See W.H. Flower, "Remarks on the homologies and notation of the +teeth in Mammalia," _J. Anat. and Physiol. norm. path._, Vol. III., p. +262; R. Owen, _Odontography_, London, 1840--45; C.S. Tomes, _Manual of +Dental Anatomy_, London, 1876. See also H.F. Osborn, "Recent +researches on succession of teeth in Mammals," _Amer. Natural._, +XXVII., p. 493, and "Rise of Mammalia in N. America," _Stud. Biol. +Lab. Columb. Coll._, Zool. I., no. 2. + +[146] See E.B. Poulton, _P.R.S._, Feb. 1888, and _Quart. J. Micr. +Sci._, Vol. XXIX. 1889; also Oldfield Thomas, _P.R.S._, XLVI. (1889). + +[147] W.H. Flower, _Phil. Trans._, vol. 156, pp. 631--641, 1867; also +Oldfield Thomas, _Phil. Trans._, pp. 443--462, 1887. + +[148] C. Röse, _Anat. Anz._ VII., p. 639. + +[149] W. Kükenthal, _Anat. Anz._ VI., p. 364. + +[150] See p. 348. + +[151] W. Leche, _Morph. Jahrb._ XX., pp. 113--142 (1893). + +[152] E.C. Stirling, _P.Z.S._ 1891, p. 327. + +[153] O. Thomas, _P.Z.S._, 1895, p. 870. + +[154] F. Ameghino, _Bull. Ac. Argen._ XII. p. 437. According to H. +Burmeister, _Annal. Mus. Buenos Aires_, III. 401 (1891), enamel does +not occur, osteodentine having been mistaken for it. + +[155] E. Ballowitz, _Arch. Mikr. Anat._ XL. p. 133. + +[156] See Oldfield Thomas, _P.R.S._, vol. XLVII., p. 246 (1890). + +[157] J. Taeker, "_Fur Kenntniss der Odontogenese bei Ungulaten_." +Dorpat, 1892. + +[158] See p. 345. + +[159] See p. 345. + +[160] According to H.F. Osborn, _Amer. Natural._, XXVI. p. 763, a +number of not very closely allied forms have been included under +_Lophiodon_. + +[161] C. Earle, _J. Ac. Philad._, vol. IX., 1892, p. 267. + +[162] _Encyclopaedia Britannica_, article _Mammalia_, p. 424. + +[163] See T.H. Huxley, "The dental and cranial characters of the +Canidae," _P.Z.S._, 1880, p. 238. + +[164] G.B. Howes, _Journ. of Anat. and Phys._ XXVII., p. 544. + + + + +CHAPTER XXIII. + +GENERAL ACCOUNT OF THE SKELETON IN MAMMALIA (CONTINUED). + + +THE SKULL AND APPENDICULAR SKELETON. + + +THE SKULL. + +MONOTREMATA. In both genera the cranium is thin-walled, has a fairly +large cavity, and is very smooth and rounded externally. The sutures +between many of the bones early become obliterated in a manner +comparable to that in birds, and the facial portion of the skull is +much prolonged. + +In _Echidna_ the face is drawn out into a gradually tapering rostrum, +formed mainly by the premaxillae, maxillae and nasals. The zygomatic +arch is very weak, and the palate extends very far back. The tympanic +forms a slender ring. The mandible is extremely slight, with no +ascending portion, and but slight traces of the coronoid process and +angle. The hyoid has a wide basi-hyal and stout thyro-hyals, while the +anterior cornua are slender, and include ossified epi-hyals and +cerato-hyals. + +In _Ornithorhynchus_ the zygomatic arch is much stouter than in +_Echidna_. The face is produced into a wide beak, mainly supported by +the premaxillae, between whose diverging anterior ends there is a +dumb-bell-shaped bone. The maxillae are flattened below, and each +bears a large horny tooth, which meets a corresponding structure borne +on a surface near the middle of the mandible. The mandible is +considerably stouter than in _Echidna_, but the angle and coronoid +process are but little developed. The infra-orbital foramen and the +inferior dental and mental foramina of the mandible are all very +large. + +[Illustration FIG. 90. HALF FRONT VIEW[165] OF THE SKULLS OF A +TASMANIAN WOLF (_Thylacinus cynocephalus_) (to the left) × 3/8; AND OF +A HAIRY-NOSED WOMBAT (_Phascolomys latifrons_) (to the right) × 3/8. +(Camb. Mus.) + + 1. premaxillae. + 2. nasal. + 3. frontal. + 4. infra-orbital foramen. + 5. lachrymal. + 6. jugal. + 7. coronoid process of the mandible. + 8. lachrymal foramen. + i. 1. first upper incisor. + C. canine.] + +MARSUPIALIA. The skulls of the various types of the Marsupials +frequently bear a strong superficial resemblance to those of some of +the different groups of placental mammals. Thus the skull of the +Dasyuridae resembles that of the Carnivora, the resemblance being most +marked between the skulls of _Thylacinus_ and the dog. The skull of +_Notoryctes_ is strongly suggestive of that of an Insectivore, and +that of other Marsupials such as the wombat, recalls equally the +characteristic features of a Rodent's skull. But, however much they +may differ from one another, the skulls of all Marsupials agree in the +following respects. (1) The brain cavity, and especially the cerebral +fossa, has a very small comparative size. (2) The nasals are always +large, and the mesethmoid is extensively ossified, and terminated by a +prominent vertical edge. (3) Processes from the jugal and frontal in +living forms never meet and enclose the orbit, but the zygomatic arch +is always complete. (4) The jugal always extends back to form part of +the glenoid fossa. (5) The lachrymal canal opens either external to or +upon the margin of the orbit, and the nasal processes of the +premaxillae never quite reach the frontals. (6) The posterior part of +the palate is commonly pierced by large oval vacuities. (7) The +tympanic is small and never fused to the bones of the cranium. (8) The +carotid canal perforates the basisphenoid and not the tympanic bulla. +(9) The optic foramen and sphenoidal fissure are confluent. (10) In +every case except _Tarsipes_ the angle of the mandible is more or less +inflected. + +The skull of the extinct _Thylacoleo_ differs from that of all other +Marsupials in the fact that the postorbital bar is complete. The hyoid +is constructed on much the same plan in all Marsupials. It consists of +a small basi-hyal, a pair of broad cerato-hyals, and a pair of strong +thyro-hyals. The epi-hyals and stylo-hyals are generally unossified. + +EDENTATA. In Sloths (Bradypodidae) the sutures become early +obliterated, the cranial portion of the skull is rather high, and the +facial portion very short. The lachrymal is very small, and its canal +opens outside the orbit. The zygomatic arch is incomplete, and the +jugal (fig. 91, 5) is curiously forked, but in a manner differing in +the two genera. The premaxillae are very small,--in _Bradypus_ quite +vestigial. The mandible is well developed, the angle being specially +marked in _Bradypus_. In _Choloepus_ the symphysial part is drawn out +in a somewhat spout-like manner (fig. 91, 6). In both genera the +thyro-hyals are ankylosed with the basi-hyal. + +[Illustration FIG. 91. SKULL OF A TWO-FINGERED SLOTH (_Choloepus +didactylus_) × 1/2. (Camb. Mus.) + + 1. anterior nares. + 2. postorbital process of the frontal. + 3. coronoid process. + 4. angle of the mandible. + 5. jugal. + 6. spout-like prolongation of the mandible.] + +In _Megatherium_ the general appearance of the skull is distinctly +sloth-like, but the facial portion is more elongated, partly owing to +the development of a prenasal bone, and the zygomatic arch is +complete. The mandible is very deep in the middle, and is drawn out +into a long spout-like process in front. + +Anteaters (Myrmecophagidae) have a much modified skull, and this is +especially the case in the Great Anteater, _Myrmecophaga_. The skull +is smooth and evenly-rounded, in these respects recalling that of +_Echidna_, but it is longer and tapers much more gradually than in +_Echidna_. The occipital condyles are remarkably large. The premaxillae +are small, and the long rostrum is chiefly composed of the maxillae and +nasals with the mesethmoid and vomer. The zygomatic arch is +incomplete, and there is no trace of a separation between the orbit +and the temporal fossa. The palate is much elongated, the pterygoids +meeting in the middle line just like the palatines. The mandible is +very long and slender, there being no definite coronoid process, and a +short and slight symphysis. The hyoid arch is noticeable for the +length of the anterior cornu. + +In the Armadillos (Dasypodidae) the skull varies a good deal in shape, +but the facial portion is always tapering and depressed. The zygomatic +arch is complete. In _Dasypus_ and _Chlamydophorus_ the tympanic bulla +is well ossified. + +In the Glyptodontidae the skull is very short and deep; the zygomatic +arch is complete, and has a long downwardly projecting maxillary +process. The mandible is massive, and has a very high ascending +portion. + +In the Manidae the skull is smooth and rounded, the zygomatic arch is +incomplete, and the orbit is inconspicuous. The palate is long and +narrow, but the pterygoids do not take part in its formation. The +mandible is slightly developed and has no angle or coronoid process. + +In _Orycteropus_ the zygomatic arch is complete, and there is a small +postorbital process to the frontal. The mandible is well-developed, +having a coronoid process and definite ascending portion, and the +hyoid is well ossified. + +SIRENIA. The skull, and especially the brain case of all Sirenia, is +remarkable for the general density of the component bones, which, +though often very thick, are without air sinuses. It is noticeable +also for the roughness of the bones, and the irregular manner in which +they are united together. + +[Illustration FIG. 92. LATERAL VIEW OF THE SKULL OF _Rhytina stelleri_ +× 1/8. (Brit. Mus.) + + 1. frontal. + 2. parietal. + 3. zygomatic process of the squamosal. + 4. squamosal. + 5. exoccipital. + 6. occipital condyle. + 7. pterygoid process of the alisphenoid. + 8. jugal. + 9. premaxillae. + 10. angle of the mandible. + 11. maxillae.] + +The cranial cavity is decidedly small, the reduction being specially +noticeable in the cerebral fossa, which is not much larger than the +cerebellar fossa. The foramen magnum is large, and the dorsal surface +of the cranium narrow. The zygomatic arch is very strongly developed, +the squamosal (fig. 92, 4) being especially prominent, and being drawn +out not only into the zygomatic process, but also into a large +post-tympanic process which articulates with the exoccipital. At the +side of the skull between the squamosal, supra-occipital and +exoccipital, there is a wide vacuity in the cranial wall, partially +filled up by the very large periotic, which is ankylosed to the +tympanic, but is not united to any other bones of the skull. The +foramen lacerum medium is confluent with the foramen lacerum anterius, +and the two together form an enormous vacuity on the floor of the +skull, bounded chiefly by the exoccipital, basi-occipital, alisphenoid +and squamosal. The jugal (fig. 92, 8) is large and in _Manatus_ sends +up a strong process, which nearly or quite meets the postorbital +process of the frontal, completing the orbit. In the other Sirenia the +orbit is completely confluent with the very large temporal fossa. The +lachrymal in _Manatus_ is very small, but is larger in _Halicore_. The +premaxillae (fig. 92, 9) are large, but smaller in _Manatus_ than in +the other genera, in all of which they are curiously bent down in +front. Their upper margin forms the anterior border of a very large +aperture lying high on the roof of the skull and extending back for a +considerable distance. This aperture is formed by the union of the two +anterior nares. The nasals are quite vestigial or absent, and the +narial aperture is bounded above by the frontals; in its floor are +seen the slender vomer and large mesethmoid. The palate is long and +narrow, and formed mainly by the maxillae; behind it there is a large +irregular process formed by the union of the palatine, pterygoid, and +pterygoid plate of the alisphenoid. The mandible is very massive and +has a very high ascending portion, a rounded angle (fig. 92, 10), and +a prominent coronoid process; the two rami are firmly ankylosed +together. The hyoid consists principally of the broad flat basi-hyal; +the anterior cornua are but slightly ossified, while the thyro-hyals +are not ossified at all. + +CETACEA. The skull in all Cetacea, especially in the Odontoceti, is a +good deal modified from the ordinary mammalian type. + +In the _ARCHAEOCETI_ this modification is less marked than in either +of the other suborders. The nasals and premaxillae are a good deal +larger than they are in living forms, and the anterior nares are +placed further forward. The maxillae do not extend back over the +frontals, and there is a well-marked sagittal crest. + +In the _MYSTACOCETI_ the skull is always quite bilaterally +symmetrical, and is not so much modified from the ordinary mammalian +type as in the Odontoceti. The parietals are not, as in the +Odontoceti, separated by a wide interparietal, but meet; they are, +however, hidden under the very large supra-occipital. The nasals are +developed to a certain extent, and the nares, though placed very far +back and near the top of the head, terminate forwardly-directed narial +passages. Turbinal bones are also developed to some extent; this fact, +and the occurrence of a definite though small olfactory fossa +constituting important distinctions from the Odontoceti. The maxillae +are large, but do not extend back to cover the frontals as in the +Odontoceti. The zygomatic process of the squamosal is very large. The +mandibular rami are not compressed, but are rounded and arched +outwards, and never meet in a long symphysis. + +_ODONTOCETI._ The skull departs widely from the ordinary mammalian +type. The following description will apply to any of the following +genera of the Delphinidae, _Phocaena_, _Globicephalus_, +_Lagenorhynchus_, _Delphinus_, _Tursiops_, _Prodelphinus_, _Sotalia_. + +The upper surface of the skull is more or less asymmetrical. The +cerebral cavity is high, short and broad; and formed mainly by the +cerebral fossa, the olfactory fossa being entirely absent. The +supra-occipital (fig. 93, 3) is very large, and forms much of the +posterior part of the roof of the skull. It has the interparietal +(fig. 93, 7) fused with it, and completely separates the two +parietals. The frontal (fig. 93, 10) is large and laterally expanded, +forming the roof of the orbit, but is almost completely covered by an +extension of the maxillae. The zygomatic arch is very slender, and is +mainly formed by a rod-like process from the jugal (fig. 93, 15), the +zygomatic process of the squamosal being short and stout. + +The nasal passages are peculiarly modified, instead of passing +horizontally forwards above the roof of the mouth, they pass upwards +and even somewhat backwards towards the top of the skull (fig. 93, +23). They are bounded laterally by two processes from the premaxillae, +the left of which is shorter than the right. The nasal cavities are +narrow and without turbinals and the nasals (fig. 93, 19) are almost +as much reduced as in Sirenia. + +[Illustration FIG. 93. A, LATERAL VIEW, AND B, LONGITUDINAL SECTION OF +THE SKULL OF A YOUNG CA'ING WHALE (_Globicephalus melas_) × 1/6. +(Brit. Mus.) + + 1. basi-occipital. + 2. exoccipital. + 3. supra-occipital. + 4. basisphenoid. + 5. alisphenoid. + 6. parietal. + 7. interparietal. + 8. presphenoid. + 9. orbitosphenoid. + 10. frontal. + 11. mesethmoid. + 12. tympanic. + 13. periotic. + 14. squamosal. + 15. jugal. + 16. vomer. + 17. palatine. + 18. pterygoid. + 19. nasal. + 20. maxillae. + 21. premaxillae. + 22. mandible. + 23. anterior nares.] + +In front of the nasal openings the face is prolonged as a narrow beak +or rostrum of varying length, formed by the maxillae and premaxillae +surrounding the vomer and large mesethmoid (fig. 93, 11), which sends +forwards a long partially cartilaginous process, and is fused behind +with the presphenoid (fig. 93, 8). The basi-occipital (fig. 93, 1) too +is fused with the basisphenoid. The foramen rotundum is confluent with +the sphenoidal fissure, and the foramen ovale with the foramen lacerum +medium and the foramen lacerum posterius. The palate is mainly formed +by the maxillae; the premaxillae and palatines (fig. 93, 17), though +both meet in symphyses, forming very little of it. The pterygoids vary +in size in the different genera, sometimes as in _Lagenorhynchus_ and +_Delphinus_ meeting in the middle line, sometimes as in _Phocaena_ and +_Globicephalus_ (fig. 93, 18) being widely separated. The tympanic and +periotic are not fused together, and the periotic has generally no +bony union with the rest of the skull. The mandible is rather slightly +developed, with the rami straight, compressed and tapering to the +anterior end. The condyle is not raised at all above the edge of the +ramus; the angle is rounded and the coronoid process is very small. +_Platanista_ has a curiously modified skull; the rostrum and mandible +are exceedingly long and narrow, and arising from the maxillae are two +great plates of bone which nearly meet above. + +In the Physeteridae the skull is raised into a very prominent crest at +the vertex behind the nares. In front of this in _Hyperoödon_ a pair +of ridges occur, formed by outgrowths from the maxillae. In the old +male these ridges reach an enormous size and almost meet in the middle +line. In _Physeter_, the Sperm whale, these ridges are not developed; +the maxillae and premaxillae unite with the other bones of the crest +enclosing an enormous half basin-shaped cavity, at the base of which +are the very asymmetrical anterior narial apertures. + +In all living Cetacea the hyoid has the same general shape, consisting +firstly of a crescentic bone formed by the fusion of the thyro-hyals +with the basi-hyal, and secondly of the anterior cornu formed +principally by the strong stylo-hyal. + +UNGULATA. None of the distinctive characters separating the Ungulata +from the other groups of mammals are drawn from the skull. But in the +Ungulata vera as opposed to the Subungulata a distinguishing feature +is found in the fact that the lachrymal and jugal form a considerable +part of the side of the face, and that the jugal always forms the +anterior part of the zygomatic arch, the maxillae taking no part in it. + + +UNGULATA VERA. + +_ARTIODACTYLA._ The skull in Artiodactyla differs from that in +Perissodactyla in the fact that the posterior end of the nasal is not +expanded and there is no alisphenoid canal. + +The skulls in the different groups of Artiodactyla differ considerably +from one another. + +[Illustration FIG. 94. A, CRANIUM AND B, MANDIBLE OF A PIG (_Sus +scrofa_) × 1/5. (Camb. Mus.) + + 1. jugal. + 2. postorbital process of the frontal. + 3. zygomatic process of the squamosal. + 4. supra-occipital. + 5. glenoid cavity. + 6. occipital condyle. + 7. foramen magnum. + 8. paroccipital process of the exoccipital. + 9. tympanic bulla. + 10. pterygoid. + 11. anterior palatine foramen. + 12. palatal plate of maxillae. + 13. coronoid process. + 14. mandibular condyle. + _i_ 1, _i_ 2, _i_ 3. first, second, and third incisors. + _c._ canine. + _pm_ 1, _pm_ 2, _pm_ 3, _pm_ 4. first, second, third, and fourth + premolars. + _m_ 1, _m_ 2, _m_ 3. first, second, and third molars.] + +The skull of the Pig[166] will be described as illustrative of the +skull in the Suina. In the Pig as in most Artiodactyla the face is +bent sharply down on the basicranial axis, the commencement of the +vomer being situated below the mesethmoid instead of in front of it as +in most skulls. The occipital region of the skull is small, and the +line of junction of the supra-occipital and parietals is raised into a +prominent occipital crest. The parietal completely fuses at an early +stage with its fellow, and the exoccipital is drawn out into a long +paroccipital process (fig. 94, A, 8). The frontal is large and broad +and drawn out into a small postorbital process. The lachrymal too is +large and takes a considerable part in forming the side of the face in +front of the orbit, as does also the jugal, though to a less extent. +The face is long and tapers much anteriorly. The nasals are long and +narrow, as are the nasal processes of the premaxillae, which do not +however reach the frontals. A prenasal ossicle is developed in front +of the mesethmoid. The palate is long and narrow, the pterygoid (fig. +94, A, 10) is small, but the pterygoid process of the alisphenoid is +prominent. The squamosal is small and has the tympanic fused with it; +the tympanic is dilated below, forming a bulla (fig. 94, A, 9) filled +with cancellous bone, and above forms the floor of a long +upwardly-directed auditory meatus. The mandible has a high ascending +portion and a small coronoid process (fig. 94, B, 13). The hyoid +differs from that of most Ungulates, the stylo-hyal being very +imperfectly ossified. + +[Illustration FIG. 95. MANDIBLE OF A HIPPOPOTAMUS (_H. amphibius_) × +1/7. (Camb. Mus.) + +The second incisor of the left side is missing and the crowns of the +grinding teeth are much worn. + + 1. condyle. + 2. coronoid process. + 3. mental foramina. + _i_ 1, _i_ 2. first and second incisors. + _c._ canine. + _pm_ 3. third premolar. + _m_ 1, _m_ 3. first and third molar.] + +In _Hippopotamus_ the skull though essentially like that of the pig is +much modified in detail. The brain cavity is very small, while the +jaws are immensely developed. The face contracts in front of the +orbits and then expands again greatly, to lodge the enormous incisor +and canine teeth. The postorbital bar is complete or nearly so, and +the orbits project curiously outwards and slightly upwards; the +lachrymal is thin and much dilated. The squamosal is drawn out into a +postglenoid process, and the hamular process of the pterygoid is +prominent. The tympanic bulla is filled with cancellous bone. The +mandible is enormously large, the symphysis is long, the angle much +expanded and drawn out into a process which projects outwards and +forwards. + +Among extinct forms related to the Suina, _Cyclopidius_ is noticeable +for having large vacuities in the lachrymo-nasal region, while +_Cotylops_ has the postorbital bar complete; both these forms are from +the North American Miocene. + +In the Tylopoda and Tragulina the skull resembles in most respects +that of the Ruminants, shortly to be described; but it is allied to +that of the Suina in having the tympanic bulla filled with cancellous +bone. The tympanic bulla is better developed in the Tragulina than in +most Ungulates. + +Among Ruminants, the Bovidae, that large group including the Oxen, +Sheep, and Antelopes, as a rule have the face bent on the basicranial +axis much as in the Suina. The parietals are generally small and early +coalesce, the frontals are large and are usually drawn out into horn +cores, which are however absent in the skulls of some domestic +varieties of sheep and oxen, and also in some of the earlier extinct +forms of Bovidae. These horn cores are formed internally of cancellous +bone, and on them the true epidermal horns are borne. In young animals +there is a distinct interparietal, but this early fuses with the +supra-occipital, and in the oxen also with the parietals. The +occipital crest is generally well marked, but in the genus _Bos_ +becomes merged in a very prominent straight ridge running between the +two horn cores; this ridge, which contains air cells communicating +with those in the horn cores, is not nearly so well marked in _Bison_. +There is often, as in _Gazella_, a vacuity on the side of the face +between the nasal, frontal, lachrymal, and maxillae, but this is not +found in oxen or sheep. The premaxillae are small, the nasals are long +and pointed, and the turbinals are much developed. The Saiga antelope +has a curiously specialised skull; the nasals are absent or have +coalesced with the frontals and the anterior nares are enormously +large. In all Ruminants the lachrymal is large and forms a +considerable part of the side of the face; it often bears a +considerable depression, the _suborbital_ or _lachrymal fossa_, well +seen in most of the smaller antelopes. The postorbital bar is +complete, and the orbit is prominent and nearly circular. The +palatines and pterygoids are moderately large, and the pterygoids have +a backwardly-projecting hamular process. The squamosal is small, but +has a postglenoid process. The tympanic is not fused to the periotic +and has a small bulla not filled with cancellous bone. There is a +large paroccipital process to the exoccipital and the mandible has a +long slender coronoid process. + +In the Cervidae and Giraffidae the face is not bent down on the +basicranial axis as it is in the Bovidae. The frontals are drawn out, +not into permanent horn cores as in the Bovidae, but into short +outgrowths, the pedicels, upon which in the Cervidae long antlers are +annually developed. These _antlers_ are outgrowths of bone, and are +covered during development by vascular integument, which dries up and +peels off when growth is complete. Every year they are detached, by a +process of absorption at the base, and shed. They may occur in both +sexes, as in the Reindeer, but as a rule they are found only in the +male. They are generally more or less branched, and are sometimes of +enormous size and weight, as in the extinct _Cervus megaceros_. In +young animals they are always simple, but become annually more and +more complicated as the animal grows older. + +In the Giraffe the frontals bear a small pair of bony cores, which are +at first distinct, but subsequently become fused to the skull. In the +allied _Sivatherium_, a very large form from the Indian Pliocene, the +skull bears two pairs of bony outgrowths, a pair of short conical +outgrowths above the orbits, and a pair of large expanded outgrowths +on the occiput. + +The opening of the lachrymal canal is commonly double and the +lachrymal fossa is large in the Cervidae and the Giraffidae except +_Sivatherium_. The vacuity between the frontal, lachrymal, maxillae, +and nasal is specially large. + +The hyoid of Ruminants is noticeable for the development of the +anterior cornua, which include stout and short cerato-hyals and +epi-hyals, long and strong stylo-hyals and large tympano-hyals which +are more or less imbedded in the tympanics. + +_PERISSODACTYLA._ In the skull of Perissodactyles an alisphenoid canal +is found and the nasals are expanded behind. Among the living animals +belonging to this group the skull least modified from the ordinary +type is that in _Rhinoceros_. In this form the skull is considerably +elongated, the facial portion being very large. The occipital region +is elevated, but the cranial cavity is small, the boundary line +between the occipital and parietal regions being drawn out into a +prominent crest, which is occupied by air cells. There is no +postorbital process to the frontal, and the orbit is completely +confluent with the temporal fossa. The nasals are fused together and +are very strongly developed, extending far forwards, sometimes +considerably beyond the premaxillae. In some extinct species, such as +_Elasmotherium_ and the Tichorhine Rhinoceros, _R. antiquitatis_, the +mesethmoid is ossified as far forwards as the end of the nasals. The +nasals are arched and bear one or two roughened surfaces to which the +great nasal horns are attached. The premaxillae are very small and the +pterygoids are slender. The palate is long, narrow, and deeply +excavated behind. The postglenoid process of the squamosal is well +developed, and generally longer than the paroccipital process of the +exoccipital. The tympanic and periotic are both small and are fused +together. The condyle of the mandible is very wide, the angle rounded, +and the coronoid process moderately developed. + +In the Titanotheriidae, a family of extinct Perissodactyla from the +Miocene of North America, the occipital region is much elevated, as is +also the fronto-nasal region, the nasals (perhaps only in the male) +bearing a pair of blunt bony outgrowths. Between these two elevated +regions the skull is much depressed. The cranial cavity is very small, +the orbit confluent with the temporal fossa, and the zygomatic arch +massive. + +In _Tapirus_ the orbit and temporal fossa are confluent. The nasals +are small, wide behind and pointed in front, and are supported by the +mesethmoid; the anterior nares are exceedingly large and their lateral +boundaries are entirely formed by the maxillae. The postglenoid and +post-tympanic processes of the squamosal are large. The periotic is +not fused to the squamosal or to the small tympanic. The mandible is +large and has the angle much developed and somewhat inflected. + +_Palaeotherium_, which lived in early Tertiary times, has a skull much +like that of the Tapir, especially as regards the nasal bones. + +In the Horse and its allies (Equidae) the facial portion of the skull +is very large as compared with the cranial portion, the nasals and +nasal cavities being specially large. In the living species of the +genus _Equus_ there is no fossa between the maxillae and lachrymal, but +it occurs in some extinct species. The lachrymal and jugal form a +considerable part of the side of the face; and the orbit though small +is complete and prominent. The postorbital bar is formed by a strong +outgrowth from the frontal, which unites with a forward extension of +the squamosal. The squamosal may extend forwards and form part of the +wall of the orbit, a very unusual feature, as in most mammals the +squamosal stops before the postorbital bar. The palate is narrow and +excavated behind as in _Rhinoceros_; the palatines take very little +part in its formation. The glenoid surface for the articulation of the +mandible is very wide. The squamosal gives rise to small postglenoid +and post-tympanic processes, and the exoccipital to a large +paroccipital process. The tympanic and periotic are ankylosed +together, but not to any other bones. + +In the SUBUNGULATA, the lachrymal and jugal do not form any +considerable part of the side of the face, and the maxillae commonly +takes part in the formation of the zygomatic arch. + +_TOXODONTIA._ The skull in the Toxodontia shows several Artiodactyloid +features, while the manus and pes are of a more Perissodactyloid type. +The Artiodactyloid features are (1) the absence of an alisphenoid +canal, (2) the fact that the palate is not excavated behind, and that +the palatines form a considerable part of it, and (3) the fusion of +the tympanic to the squamosal and exoccipital, forming the floor of an +upwardly directed auditory meatus. The frontal has a fairly well +developed postorbital process, but the orbit is confluent with the +temporal fossa. The premaxillae is well developed, as is the +paroccipital process of the exoccipital, especially in _Typotherium_. +The mandible has a rounded angle and a coronoid process of moderate +size. In _Typotherium_ the ascending portion is very massive. + +_CONDYLARTHRA._ As far as is known the skull of these generalised +Ungulates is depressed, and is frequently marked by a strong sagittal +crest. The cranial cavity is small, the cerebral fossa in _Phenacodus_ +being exceptionally small. The orbit is completely confluent with the +temporal fossa. + +_HYRACOIDEA._ The skull of _Procavia_ resembles that of +Perissodactyles more than that of any other Ungulates, but differs +strongly in the comparatively small size of its facial portion. The +posterior portion of the cranium is rather high, the occipital plane +being nearly vertical. There is a small interparietal. The nasals are +wide behind, and the zygomatic arch is strongly developed, its most +anterior part being formed by the maxillae. The jugal and parietal give +rise to postorbital processes which sometimes meet, but as a rule the +orbit is confluent with the temporal fossa; it is very uncommon for +the parietal to give rise to a postorbital process, and even in +_Procavia_ the frontal often forms part of the process. The +alisphenoid canal, and postglenoid and paroccipital processes are well +developed. The tympanic bulla is large and the periotic and tympanic +are fused together, but not as a rule to the squamosal. The ascending +portion of the mandible is very high and broad, the angle rounded and +the coronoid process moderate in size. The hyoid is singular, there is +a large flat basi-hyal prolonged laterally into two broad flattened +thyro-hyals. Articulating with its anterior end are two large +triangular cerato-hyals, which are drawn out into two processes +meeting in the middle line. + +_AMBLYPODA._ In the Uintatheriidae (Dinocerata) the skull has a very +remarkable character, being long and narrow and drawn out into three +pairs of rounded protuberances, a small pair on the nasals, a larger +pair on the maxillae in front of the orbits, and the largest pair on +the parietals. The cranial cavity, and especially the cerebral fossa, +is extraordinarily small. The orbit is not divided behind from the +temporal fossa. The mandible has a prominent angle, and a long curved +coronoid process; its symphysial portion bears a curious flattened +outgrowth to protect the great upper canines. + +In _Coryphodon_ the skull is of a more normal character, being without +the conspicuous protuberances. The cranial cavity though very small is +not so small as in _Uintatherium_. + +[Illustration FIG. 96. SKULL OF A YOUNG INDIAN ELEPHANT (_Elephas +indicus_), SEEN FROM THE RIGHT SIDE, THE ROOTS OF THE TEETH HAVE BEEN +EXPOSED. × 1/8. (Camb. Mus.) + + 1. exoccipital. + 2. parietal. + 3. frontal. + 4. squamosal. + 5. jugal. + 6. premaxillae. + 7. maxillae. + 9. supra-occipital. + 13. basi-occipital. + 14. postorbital process of the frontal. + 15. lachrymal. + 16. pterygoid process of the alisphenoid. + _i_ 1. incisor. + _mm_ 3., _mm_ 4. third and fourth milk molars. + _m_ 1. first molar.] + +_PROBOSCIDEA._ The character of the skull in the young elephant +differs much from that in the old animal. In very young individuals +the skull is of a normal character, and the cranial cavity is +distinctly large in proportion to the bulk of the skull. But as the +animal gets older, while its brain does not grow much, the size of its +trunk and especially of its tusks increases greatly; and consequently +the skull wall is required to be of very great superficial extent in +order to afford space for the attachment of the muscles necessary for +the support of these heavy weights. This increase in superficial +extent is brought about without much increase in weight of bone by the +development of an enormous number of air cells in nearly all the bones +of the skull; sometimes, as in the case of the frontal, separating the +inner wall of the bone from the outer, by as much as a foot. This +development of air cells is accompanied by the obliteration of the +sutures between the various bones. The most noticeable point with +regard to the cranial cavity is the comparatively large size of the +olfactory fossa. The supra-occipital (figs. 96 and 97, 9) is +large--exceedingly large in the adult skull; the parietals (figs. 96 +and 97, 2) are also very large. The frontals send out small +postorbital processes, but these do not meet processes from the small +jugal, which forms only the middle part of the slender zygomatic +arch, the anterior part being formed by the maxillae. The lachrymal +(fig. 96, 15) is small and lies almost entirely inside the orbit. The +anterior narial aperture (fig. 97, 8) is wide and directed upwards, +opening high on the anterior surface of the skull. It is bounded above +by the short thick nasals and below by the premaxillae. The narial +passage is freely open, maxillo-turbinals not being developed. The +palatine is well developed, the pterygoid is small and early fuses +with the pterygoid process of the alisphenoid. The tympanic is united +with the periotic but not with the squamosal, and forms a large +auditory bulla. There are no paroccipital or postglenoid processes. +The exoccipital is not perforated by the condylar foramen,--a very +exceptional condition. + +[Illustration FIG. 97. LONGITUDINAL SECTION TAKEN RATHER TO THE RIGHT +OF THE MIDDLE LINE OF THE SKULL OF A YOUNG INDIAN ELEPHANT (_E. +Indicus_) × 1/8. (Camb. Mus.) + + 8. anterior nares. + 10. periotic. + 11. palatine. + 12. pterygoid. + 17. nasal. + +Other numbers as in Fig. 96.] + +The mandible has a high ascending portion, is rounded off below and +has no angle. The symphysial portion is long, narrow, and spout-like, +and the coronoid process is small. The thyro-hyals are ankylosed with +the basi-hyal, which is connected with the large forked stylo-hyals by +ligament only. + +RODENTIA. The cranial cavity is depressed, elongated, and rather +small, and the cerebral fossa lies entirely in front of the cerebellar +fossa. The occipital plane is vertical or directed somewhat backwards, +and the supra-occipital does not form much of the roof of the cranium. +The paroccipital processes of the exoccipitals are generally of +moderate size; in the Capybara (_Hydrochaerus_), however, they are +very long, and are laterally compressed and directed forwards. The +parietals are small, and often become completely fused together; there +is sometimes a small interparietal. The frontals in most genera have +no trace of a postorbital process; in Squirrels, Marmots and Hares, +however, one occurs, but in no case does it meet a corresponding +process from the zygomatic arch, so the orbit and temporal fossa are +completely confluent. In Hares the postorbital process of the frontal +is much flattened, and has an irregular margin. The temporal fossa is +always small, and in _Lophiomys_ is arched over by plates arising +respectively from the parietal and jugal; a secondary roof is thus +partially developed in a manner unique among mammals, but carried to a +great extent in many Chelonia. The nasal bones and cavities are large, +attaining their maximum development in the Porcupines (fig. 98, 1). +The premaxillae is always very large, and sends back a long process +which meets the frontal. The vomer is occasionally found persisting in +two separate halves, a feature recalling the arrangement in +Sauropsids. In many Rodents there is an enormous vacuity at the base +of the maxillary portion of the zygomatic arch. It is sometimes as +large as the orbit, and attains its maximum development in the +Capybara and other Hystricomorpha; in the Marmots, Beavers, and +Squirrels (Sciuromorpha), and in the Hares it is undeveloped. In +_Lagostomus_ the maxillae bears an upwardly directed plate of bone, +shutting off from this vacuity a space which is the true infra-orbital +foramen. + +[Illustration FIG. 98. HALF FRONT VIEW OF THE SKULL OF A PORCUPINE +(_Hystrix cristata_) × 1/2. (Camb. Mus.) + + 1. nasal. + 2. maxillo-turbinals. + 3. infra-orbital vacuity. + 4. maxillae. + 5. premaxillae. + 6. jugal. + _i_ 1. upper incisor.] + +The zygomatic arch is always complete, and in many cases the jugal +extends back to form part at least of the glenoid surface for +articulation with the mandible. In _Coelogenys_ the jugal and +maxillary portion of the zygomatic arch is greatly expanded and +roughened, and the maxillary portion encloses a large cavity. The +palate in Rodents is narrow, and the space between the incisor and +molar teeth passes imperceptibly into the sides of the face. The +anterior palatine foramina form long, rather narrow slits in this +region. The bony palate between the grinding teeth is sometimes as in +the Hares very short, sometimes as in the Capybara very long. The +maxillae extends back beneath the orbit to unite with the squamosal. +The pterygoid is always small, but sometimes has a well-marked hamular +process which in _Hystrix_, _Lagostomus_, and some other genera unites +with the tympanic bulla. The periotic is large, and fused with the +tympanic, which forms a prominent bulla, and is generally drawn out +into a tubular meatus. The bulla attains its maximum development in +_Chinchilla_ and _Dipus_. + +The mandible is narrow and rounded in front, the two halves meeting in +a long symphysis. The angle is generally drawn out into a long +backwardly-projecting process, which is often pointed and directed +upwards. In the Hares the angle is rounded. The coronoid process is +never large. + +There are a number of points in which the skull of the Duplicidentata +(Hares and Rabbits) differs from that of other Rodents. (_a_) The +sutures between the basi-occipital and basisphenoid, and between the +basisphenoid and presphenoid remain open throughout life. (_b_) Much +of the maxillae forming the side of the face in front of the orbit is +fenestrated. (_c_) The optic foramina are united to form a single +hole, much as in birds. (_d_) The coronoid process is slightly +differentiated from the ascending portion of the mandible. The first +two of these points have been thought to indicate degradation of the +hares and rabbits as compared with higher mammals. + +CARNIVORA[167]. It is characteristic of the skull in Carnivora that +the glenoid fossa is deep, and the postglenoid process (fig. 75, 23) +well developed. The condyle of the mandible is much elongated +transversely. The orbit and temporal fossa in the great majority of +forms communicate freely, the postorbital bar being incomplete. + +_CARNIVORA VERA._ The axis of the facial portion of the skull is a +direct continuation of that of the cranial portion. The cranial +cavity though rather depressed is large, and generally long, though in +Cats it is comparatively short and wide. The occipital plane is nearly +vertical, and the exoccipitals are developed into fairly prominent +paroccipital processes. The interparietal is commonly distinct, and +the parietals unite in a long sagittal suture, which is often +developed into a crest. The nasals (fig. 73, 4) are well developed, +especially in Cats, and the nasal processes of the premaxillae do not +nearly reach the frontals. A considerable part of the palate is formed +by the palatine, and the maxillary portion is pierced by rather long +anterior palatine foramina. The pterygoid has a hamular process. The +zygomatic arch is strong, especially in Cats. Postorbital processes +are developed on the frontal (fig. 73, 10) and jugal, but never form a +complete postorbital bar. A carotid canal is well seen in the Ursidae, +and to a less extent in the Felidae; in the Canidae there is an +alisphenoid canal (fig. 75, 21). + +The auditory bulla differs a good deal in the different groups. In the +Bears (Ursidae) it is not much inflated, and is most prominent along +its inner border; it is not closely connected with the paroccipital +process. In the Cats it is very prominent, and its cavity is almost +divided by a septum into two parts, the inner of which contains the +auditory ossicles. The paroccipital process is closely applied to the +bulla. In the Dogs the bulla is intermediate in character between that +of the Cats and that of the Bears; it is partially divided by a +septum, and is moderately expanded. + +The mandible is well developed with a prominent angle (fig. 72, 26), +and a large coronoid process. The hyoid consists of a broad basi-hyal, +a long many-jointed anterior cornu and short thyro-hyals (fig. 72, +33). + +The skull in the _CREODONTA_ is in most respects allied to that of the +Canidae, but presents some ursine affinities. The tympanic bulla is +fairly prominent, but has no well-developed septum. The cranial cavity +is very small and narrow, the zygomatic arch standing away from it. +The temporal fossa is of great size. + +In the _PINNIPEDIA_ the cranial cavity is large and rounded. The skull +is much compressed in the interorbital region, and in correlation with +this compression the ethmo-turbinals are little developed, while the +maxillo-turbinals are large. The orbit is large, and the temporal +fossa smaller than in the Carnivora vera. In the Walrus (_Trichechus_) +the anterior part of the face is distorted by the development of the +huge canines. The Otariidae have an alisphenoid canal. The tympanic +bulla is small in _Otaria_, large in the Phocidae, and flattened in +the Walrus. The hyoid is similar to that in Carnivora vera. + +INSECTIVORA. The skull varies much in the different members of the +order Insectivora, but the following points of agreement are found. +The cranial cavity is of small size, and is never much elevated. The +facial part of the skull is generally considerably elongated, and the +nasals and premaxillae are well developed. The zygomatic arch is +usually slender or incomplete, and the coronoid process and angle of +the mandible are commonly prominent. + +In some Insectivora, such as _Galeopithecus_, _Tupaia_, and +_Macroscelides_, the skull shows a higher type of structure than is +met with in most members of the order. In these genera the cranial +cavity is comparatively large, and the occipital plane is nearly +vertical. The zygomatic arch is fairly strong, and the frontal and +jugal give rise to postorbital processes which nearly or quite +(_Tupaia_) meet. The tympanic bulla is well developed, and produced +into a tubular auditory meatus, this being specially well marked in +_Macroscelides_. + +In the other Insectivora the cranial cavity is of smaller comparative +size, and the orbit and temporal fossa are completely confluent, often +without any trace of a postorbital bar. The occipital plane commonly +slopes forwards. In the Hedgehogs (Erinaceidae) and Centetidae the +tympanic is very slightly developed, forming a small ring. The +zygomatic arch of Hedgehogs and _Gymnura_ is very slender, the jugal +being but little developed and the squamosal and maxillae meeting one +another; in the Centetidae the jugal is absent and the arch is +incomplete. + +The Moles (Talpidae) have an elongated, depressed and rounded skull +with a very slender zygomatic arch formed by the squamosal and +maxillae. The nasals are fused together, and the mesethmoid is ossified +very far forwards. In the Shrews (Soricidae) there is no zygomatic +arch; the tympanic is ring-like, and the angle of the mandible is very +prominent. The hyoid has a transversely extended basi-hyal, a long +anterior cornu with three ossifications, and thyro-hyals which are +sometimes fused to the basi-hyal. + +CHIROPTERA. In the frugivorous Flying Foxes (Pteropidae) the skull is +elongated, and the cranial cavity is large and arched, though +considerably contracted in front. There are commonly strong sagittal +and supra-orbital crests. The parietals take a great part in the +formation of the walls of the cranial cavity, the supra-occipital and +frontals being small. The frontal is drawn out into a long postorbital +process, but the zygomatic arch, which is slender, and formed mainly +by the squamosal and maxillae, gives rise to only a small postorbital +process, so that the orbit and temporal fossa are confluent. There is +no alisphenoid canal, and the tympanics are very slightly connected +with the rest of the skull. The mandible has a large coronoid process, +a rounded angle, and a transversely expanded condyle. + +In Insectivorous Bats the skull is generally shorter and broader than +in the Pteropidae. The cranial cavity is large and rounded, and has +thin smooth walls. The zygomatic arch is slender, and postorbital +processes are not generally well developed. The premaxillae is +generally small, sometimes absent. The tympanics are ring-like and are +not connected with the surrounding bones. The angle of the mandible is +distinct. The hyoid in most respects resembles that of the +Insectivora. + +PRIMATES. The characters of the skull differ greatly in the two +suborders of Primates, the Anthropoidea and the Lemuroidea. + +In the _LEMUROIDEA_ the general relative proportions of the cranium +and face are much as in most lower mammals, and the occipital plane +forms nearly a right angle with the basicranial axis. The postorbital +processes of the frontals are commonly continued as a pair of ridges +crossing the roof of the cranium and meeting the occipital crest. +Though the postorbital bar is complete, the orbit and temporal fossa +communicate freely below it. The lachrymal canal opens outside the +orbit, and the lachrymal forms a considerable part of the side of the +face. The tympanic is developed into a large bulla. The hyoid +apparatus much resembles that of the Dog. + +In the _ANTHROPOIDEA_ the skull differs greatly from that in the +Lemuroidea. The cranial portion of the skull is very large as compared +with the facial portion, though the comparative development varies, +some monkeys, such as the baboons (Cynocephali) having the facial +portion relatively large. The comparative size of the jaws does not +vary inversely with the general development of the animal, some of the +Cercopithecidae having comparatively larger jaws than some of the +Cebidae. The great size of the cranial part of the skull is mainly due +to the immense development of the cerebral fossa, which commonly +completely overlaps the olfactory fossa in front, and the cerebellar +fossa behind. This development also has the effect of making the +ethmoidal and occipital planes lie, not at right angles to the +basicranial axis, but almost in the same straight line with it. This +is, however, not always the case, as the Howling Monkey (_Mycetes_) +and also some of the very highest monkeys, the Gibbons (_Hylobates_), +have the occipital plane nearly vertical to the basicranial axis. In +adult Man the basi-occipital, exoccipitals and supra-occipital +coalesce, forming the so-called occipital bone; while the +basisphenoid, presphenoid, alisphenoids, orbitosphenoids and +pterygoids form the sphenoid bone. The roof of the skull is partly +formed by the large supra-occipital and frontals, but mainly by the +parietals (fig. 99, 1), which in Man are of enormous extent. + +[Illustration FIG. 99. HALF FRONT VIEW OF THE SKULLS, _A_ OF AN OLD, +_B_ OF A YOUNG GORILLA (_Gorilla savagei_) × 1/4. (Camb. Mus.) + + 1. parietal. + 2. sagittal crest. + 3. frontal. + 4. supra-orbital ridge. + 5. squamosal. + 6. maxillae. + 7. external auditory meatus.] + +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 the frontals 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. In _Mycetes_ the mandible is very +large, its ascending portions being specially developed. The hyoid of +Primates is remarkable 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. In _Mycetes_ the +basi-hyal is enormously large, forming a somewhat globular thin-walled +capsule. + + +AUDITORY OSSICLES. + +[Illustration FIG. 100. MALLEUS, STAPES AND INCUS OF _A._ MAN. _B._ +DOG. _C._ RABBIT. (After DORAN) x 1. + + 1. head of malleus. + 2. canal of stapes. + 3. incus. + 4. processus longus (or gracilis). + 5. manubrium of malleus. + 6. processus brevis. + 7. lamella.] + +There are in mammals four auditory ossicles forming a chain extending +from the fenestra ovalis to the tympanic membrane. Three of these, the +=malleus=, =incus= and =stapes=, are always distinct, while the +fourth, the =lenticular=, 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 +homologous as 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. The =malleus= when typically developed consists of a +rounded _head_ (fig. 100, 1) which bears a surface articulating with +the incus, and a short _neck_ continued into a process, the +_manubrium_ (fig. 100, 5), which comes into relation with the tympanic +membrane. From the junction of the neck and manubrium two processes +are given off, a _processus longus_ or _gracilis_ (fig. 100, 4), which +in the embryo is continuous with Meckel's cartilage, and a _processus +brevis_ (fig. 100, 6). The =incus= generally 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 small =lenticular=. +The =stapes= is generally stirrup shaped, consisting of a basal +portion from which arise two _crura_ separated 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. + +In MONOTREMES the 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. + +In MARSUPIALS the 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 developed +ossicles, the malleus much resembling that of many Insectivores, and +the stapes having two definite crura separated by a canal. + +In EDENTATES the 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 in +_Manis_, 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. _Priodon_ has a lower type +of stapes than _Dasypus_ and _Tatusia_. + +The ossicles of the SIRENIA differ widely from those of all other +mammals in their great density and clumsy form. + +In CETACEA the 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 the UNGULATA do 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 of _Procavia_, 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 the RODENTIA (fig. 100, C) the malleus is generally characterised +by a very broad manubrium. In many genera such as _Bathyergus_, and +most of the Hystricomorpha such as _Hystrix_, _Chinchilla_ and +_Dasyprocta_, 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. + +In INSECTIVORA the 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. +_Chrysochloris_ has 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. + +In CHIROPTERA the 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]. + +In MONOTREMES and most MARSUPIALS the sternum does not present any +characters of special importance. The presternum is strongly keeled in +_Notoryctes_. + +The sternum in EDENTATES is very variable: in the Sloths it is very +long, the mesosternum of _Choloepus_ having twelve segments. In the +ant-eaters and armadillos the presternum is broad and sometimes as in +_Priodon_ strongly keeled. In _Manis macrura_ the xiphisternum is +drawn out into a pair of cartilaginous processes about nine inches +long. + +In the SIRENIA the 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 in +_Manatus_. + +Two distinct types of sternum are met with in the CETACEA. 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 in _Hyperoödon_. 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 in UNGULATA is 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 the RODENTIA the sternum is long and narrow and generally has a +large presternum, and a xiphisternum terminated by a broad +cartilaginous plate. + +In the CARNIVORA, 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. + +In INSECTIVORA the 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 the CHIROPTERA the presternum is strongly keeled and so is +sometimes the mesosternum. + +Among PRIMATES, 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. + +In MONOTREMES (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. + +In MARSUPIALS there are almost always thirteen pairs of ribs, whose +sternal portions are very imperfectly ossified. _Notoryctes_ has +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 the EDENTATES the Sloths have very numerous ribs; twenty-four pairs +occur in _Choloepus_, 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 the SIRENIA there 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 the UNGULATA the ribs are generally broad and flattened, and this +is especially the case in the genera _Bos_ and _Bubalus_ (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, and _Procavia_ twenty +to twenty-two. The Elephant has nineteen to twenty-one pairs, seven of +which may be floating ribs. + +[Illustration 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. + 2. nasal. + 3. orbit. + 4. neural spine of first thoracic vertebra. + 5. scapula. + 6. rib. + 7. femur. + 8. patella. + 9. tibia. + 10. metatarsals. + 11. radius. + 12. metacarpals.] + +In the RODENTIA there are generally thirteen pairs of ribs, which do +not present any marked peculiarities. + +The CARNIVORA have 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 either +INSECTIVORA or CHIROPTERA. + +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 the MONOTREMATA. 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 large =T=-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. + +[Illustration FIG. 102. _A_, SIDE VIEW, _B_, DORSAL VIEW OF THE +SHOULDER GIRDLE AND PART OR THE STERNUM OF THE SPINY ANTEATER +(_Echidna aculeata_) × 1. (After PARKER.) + + 1. scapula. + 2. suprascapula. + 3. clavicle. + 4. interclavicle. + 5. coracoid. + 6. epicoracoid. + 7. glenoid cavity. + 8. presternum. + 9. second sternal rib. + 10. second vertebral rib.] + +In MARSUPIALS, 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 in _Perameles_. Unossified remains of the precoracoids +are found at either end of the clavicle. The scapula of _Notoryctes_ +has a very high overhanging spine, and there is a second strong ridge +running along the proximal part of the glenoid border. + +The shoulder girdle of the EDENTATA shows some very curious +variations. In _Orycteropus_ the 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_, and +_Myrmecophaga_, 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. In _Bradypus_ the clavicle is +very small, and is attached to the coracoid, which sometimes forms a +distinct bone[170]. + +In the SIRENIA the 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 +as _Balaenoptera_, have a broad, fan-shaped scapula, with a long +acromion and coracoid process, extending parallel to one another. +Others, such as _Balaena_, have a higher and narrower scapula, with a +smaller coracoid process. + +In UNGULATA the scapula is always high and rather narrow, and neither +acromion nor coracoid process is ever much developed. In no adult +Ungulate except _Typotherium_ is there any trace of a clavicle, but a +vestigial clavicle has been described in early embryos of sheep[171]. + +[Illustration FIG. 103. SKELETON OF A LLAMA (_Auchenia glama_) × 1/18. +(Brit. Mus.) + + 1. hyoid. + 2. atlas vertebra. + 3. seventh cervical vertebra. + 4. scapula. + 5. imperfectly ossified suprascapula. + 6. olecranon process of ulna. + 7. metacarpals. + 8. ilium. + 9. patella. + 10. calcaneum.] + +UNGULATA VERA. In the Ruminantia the suprascapular region (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. In +_Hippopotamus_ the 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 and +_Hyracotherium_ have 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 in _Rhinoceros_. + +_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. + +_Procavia_ has 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. + +In RODENTIA the 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 the CARNIVORA VERA the 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. + +[Illustration FIG. 104. DORSAL VIEW OF THE STERNUM AND RIGHT HALF OF +THE SHOULDER-GIRDLE OF _Mus sylvaticus_ × 4. (After PARKER.) + + 1. postscapular fossa. + 2. prescapular fossa. + 3. spine. + 4. suprascapular border unossified. + 5. coracoid process. + 6. acromion. + 7. cartilaginous vestige of precoracoid at scapular end of clavicle. + 8. clavicle. + 9. cartilaginous vestige of precoracoid at sternal end of clavicle. + 10. omosternum. + 11. epicoracoid. + 12. presternum. + 13. first segment of mesosternum. + 14. xiphisternum. + 15. cartilaginous termination of xiphisternum. + 16. 2nd sternal rib. + 17. 1st vertebral rib.] + +In the INSECTIVORA the 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. In +_Potamogale_, 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 the CHIROPTERA the 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. + +In PRIMATES the 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 the MONOTREMATA the 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 the MARSUPIALIA the humerus is generally a strong bone, broad at +the distal end and having well marked deltoid and supinator ridges, +which are specially large in _Notoryctes_. An ent-epicondylar or +supracondylar foramen (fig. 105, 5) is almost always present except in +_Notoryctes_. The radius and ulna are always distinct and well +developed, and a certain amount of rotation can take place between +them. The ulna of _Notoryctes_ has an enormous hooked olecranon which +causes the bone to be nearly twice as long as the radius. + +[Illustration FIG. 105. ANTERIOR SURFACE OF THE RIGHT HUMERUS OF A +WOMBAT (_Phascolomys latifrons_). (After OWEN.) + + 1. head. + 2. greater tuberosity. + 3. lesser tuberosity. + 4. deltoid ridge. + 5. ent-epicondylar (supracondylar) foramen. + 6. supinator ridge. + 7. external condyle. + 8. internal condyle. + 9. articular surface for radius. + 10. articular surface for ulna.] + +EDENTATA. The Sloths have long slender arm bones; the humerus is +nearly smooth and has a very large ent-epicondylar foramen in +_Choloepus_, but not in _Bradypus_. The radius and 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 the SIRENIA the 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 and _Rhytina_ there 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 the CETACEA the 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 the UNGULATA VERA the humerus is stout and rather short. The great +tuberosity is always large and often overhangs the bicipital groove, +it is especially large in _Titanotherium_ (_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 in +_Tapirus_, _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 process which 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. + +In _Procavia_ the 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. + +In RODENTIA the 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 the INSECTIVORA the 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 the CHIROPTERA both 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. + +All PRIMATES have 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. the union of carpalia, 4 and 5 to form the +_unciform_ bone. Two sesamoid bones are commonly developed, one on +each side of the carpus, the _pisiform_ or 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. In _Echidna_ the 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. In _Ornithorhynchus_ the manus is more slender, +but the general arrangement is the same as in _Echidna_. + +MARSUPIALIA. The carpus has no centrale and the lunar is generally +small or absent. Five digits are almost always present. In _Choeropus_ +however 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 in _Notoryctes_ is 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 the EDENTATA there is a great diversity in the structure of the +manus, the centrale is however always wanting, and except in _Manis_ +the 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. In _Myrmecophaga_ all five +digits are fairly well though irregularly developed, in _Cycloturus_ +the 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. In _Dasypus_ they are regular, but are +remarkably irregular in Priodon. The pollex is absent in Glyptodonts +and in _Megatherium._ In _Megatherium_ the 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. In _Orycteropus_ the pollex is absent, +while the other digits are terminated by pointed ungual phalanges. + +In SIRENIA the general structure of the manus is quite of the ordinary +mammalian type. In _Manatus_ most of the bones of the carpus are +distinct, but in _Halicore_ many, especially those of the distal row, +have coalesced. The digits are always five in number and have the +normal number of flattened phalanges. + +In the CETACEA, 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. In _Physeter_ the carpal bones also have epiphyses. The +carpus generally consists of six bones arranged in two rows of three +each. Five digits are generally present, but sometimes as in +_Balaenoptera 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 in +_Globicephalus_, 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 the UNGULATA VERA the manus is never plantigrade, and there are not +more than four digits, the pollex being almost always completely +suppressed: in _Cotylops_ among 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 as _Anoplotherium_, have their ungual phalanges flattened +on their contiguous surfaces. + +In all _ARTIODACTYLA_ the 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 in +_Dorcatherium_ the 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. + +[Illustration FIG. 106. MANUS OF PERISSODACTYLES. + +_A._ LEFT MANUS OF _Tapirus_. (After VON ZITTEL.) + +_B._ RIGHT MANUS OF _Titanotherium_. (After MARSH.) + +_C._ LEFT MANUS OF _Chalicotherium gigantium_. (After GERVAIS.) + + 1. scaphoid. + 2. lunar. + 3. cuneiform. + 4. trapezoid. + 5. magnum. + 6. unciform. + 7. trapezium. + II, III, IV, V. second, third, fourth and fifth digits.] + +While the manus of the Artiodactyla is symmetrical about a line drawn +between the third and fourth digits, that of the _PERISSODACTYLA_ is +symmetrical about a line drawn through the middle of the third digit, +which is larger than the others and has its ungual phalanx evenly +rounded and symmetrical in itself. The most reduced manus in the whole +of the mammalia is found in the Horse and its allies, in which the +third digit, terminated by a very wide ungual phalanx, is the only one +functional. Small splint bones representing the second and fourth +metacarpals are attached to the upper part of the third metacarpal. In +_Hipparion_[174] and other early horse-like animals the second and +fourth digits, though very small and functionless, are complete and +are terminated by small hoofs. In _Rhinoceros_ the second and fourth +digits are equally developed and nearly as large as the third, and +reach the ground in walking, a vestige of the fifth is also present. +In the Tapir (fig. 106, A) and _Hyracotherium_ the fifth digit is +fully developed but is scarcely functional. In _Titanotherium_ +(_Brontops_) (fig. 106, B) it is nearly as well developed as any of +the others, and there is little or no difference between the relative +development of the third and fourth digits. + +The Chalicotheriidae[175], though distinctly Perissodactyles in +various respects such as their cervical vertebrae and teeth, differ +not only from all other Perissodactyles, but from almost all other +Ungulates, in the very abnormal character of their manus. For while +the carpus and metacarpus are like those of ordinary Perissodactyles, +the phalanges resemble those of Edentates, each second phalanx having +a strongly developed trochlea, and each distal one being curved, +pointed and deeply cleft at its termination (fig. 106, C). + +The Macraucheniidae, while agreeing with Perissodactyles in having +only three digits, with the limb symmetrical about a line drawn +through the middle of the third, have a carpus which approaches +closely to the subungulate condition, the magnum articulating +regularly with the lunar, and only to a slight extent with the +scaphoid. + +In the SUBUNGULATA the manus sometimes has five functional digits, and +a considerable part of it rests on the ground in walking. The bones of +the carpus retain their primitive relation to one another, the magnum +articulating with the lunar, but not with the scaphoid. This character +does not however hold in the Toxodontia, for in most of the animals +belonging to this group the magnum does articulate with the scaphoid. +The corner of the scaphoid just reaches the magnum also in Amblypoda. + +As far as is known the _TOXODONTIA_ generally have three, sometimes +five digits to the manus, and the third is symmetrical in itself--a +Perissodactyloid feature. + +In _Phenacodus_ (fig. 107, B) (_CONDYLARTHRA_) all five digits are +well developed, the pollex being the smallest. The carpal bones retain +their primitive arrangement, the magnum articulating with the lunar +and not with the scaphoid. There is no separate centrale. + +[Illustration FIG. 107. LEFT MANUS OF + +_A. Coryphodon hamatus._ (After MARSH.) × 1/5. + +_B. Phenacodus primaevus._ (After COPE.) × 1/3. + +_C. Procavia (Dendrohyrax) arboreus._ (After VON ZITTEL.) × 6/7. + + 1. scaphoid. + 2. lunar. + 3. cuneiform. + 4. trapezium. + 5. trapezoid. + 6. magnum. + 7. unciform. + 8. centrale. + 9. pisiform. + I, II, III, IV, V. first, second, third, fourth and fifth digits + respectively.] + +In the _HYRACOIDEA_ (fig. 107, C) the manus is very similar to that in +_Phenacodus_, but a centrale is present and the pollex is much +reduced. + +The manus of the _AMBLYPODA_, such as _Coryphodon_ (fig. 107, A) and +_Uintatherium_, is short and broad, with five well developed digits +and large carpal bones. The carpals however interlock to a slight +extent, and the corner of the magnum reaches the scaphoid. + +In the _PROBOSCIDEA_ the manus is very short and broad, with large +somewhat cubical carpals which articulate by very flat surfaces and do +not interlock at all. All five digits are present, and none of them +are much reduced in size. The manus in Proboscidea and in _Coryphodon_ +is subplantigrade. + +In the Tillodontia the manus is plantigrade and has pointed ungual +phalanges, in this respect approaching the Carnivora. It differs +however from that of all living Carnivora in having the scaphoid and +lunar distinct. + +In RODENTIA the manus nearly always has five digits with the normal +number of phalanges: the pollex may however be very small as in the +Rabbit, or absent as sometimes in the Capybara. The scaphoid and lunar +are generally united, and a centrale may be present or absent. In +_Pedetes caffer_ the radial sesamoid is double and the distal bone +bears a nail-like horny covering. In _Bathyergus_ the pisiform is +double. It is upon these facts that the contention for the former +existence of prehallux and post-minimus digits has partly been based. + +In living CARNIVORA the scaphoid, lunar and centrale are always +united, forming a single bone. All five digits are present, but as a +rule in Carnivora vera the pollex is small, and in _Hyaena_ is +represented only by a small metacarpal. Sometimes, as in Cats and +Dogs, the manus is digitigrade, sometimes, as in Bears, plantigrade. +The ungual phalanges are large and pointed, and in forms like the +Cats, whose claws are retractile, they can be folded back into a deep +hollow on the ulnar side of the middle phalanx; a small radial +sesamoid is often present. + +In Pinnipedia the manus is large and flat and the digits are +terminated by ungual phalanges which are blunt (sea lions and walrus), +or slightly curved and pointed (seals). The pollex is nearly or quite +as long as the second digit, and as a rule the digits then +successively diminish in size. + +The Creodonta differ from living Carnivora in the fact that the +scaphoid and lunar are usually separate. + +In INSECTIVORA the scaphoid and lunar are sometimes united, sometimes +separate, and a separate centrale is usually present. There are +generally five digits, but sometimes the pollex is absent. In the Mole +the manus is greatly developed and considerably modified. It is very +wide, its breadth being increased by the great development of the +radial sesamoid which is very large and sickle-shaped. The ungual +phalanges are also large and are cleft at their extremities. + +In the CHIROPTERA the manus is greatly modified for the purpose of +flight. The pollex is short and is armed with a rather large curved +claw, the other digits are enormously elongated, the elongation in the +case of the Insectivorous bats being mainly due to the metacarpals, +and in the Frugivorous bats to the phalanges. In the Frugivorous bats +the second digit is clawed as well as the pollex, in other bats this +claw is always absent, and so is often the ungual phalanx, the middle +phalanx then tapering gradually to its termination. + +In PRIMATES as a rule the manus is moderately short and wide. The +carpus has the scaphoid and lunar distinct, and generally also the +centrale; sometimes however, as in Man, the Gorilla, Chimpanzee, and +some Lemurs, the centrale has apparently fused with the scaphoid. +There are almost always five well-developed digits, but in the genera +_Colobus_ and _Ateles_ the pollex is vestigial. + +The magnum in man is the largest bone of the carpus. The pisiform also +is well developed, but there is no radial sesamoid. In Man, the +Gorilla, Chimpanzee, and Orang, the carpus articulates only with the +radius, in most Primates it articulates also with the ulna. The third +digit of the Aye-Aye (_Chiromys_) is remarkable for its extreme +slenderness. + + +THE PELVIC GIRDLE. + +The pelvic girdle in all mammals except the Sirenia and Cetacea +consists of two halves, usually united with one another at the +symphysis in the mid-ventral line, and connected near their upper +ends, with the sacral vertebrae. Each half forms one of the +_innominate_ bones, and includes at least three separate elements, a +dorsal bone, the ilium, and two ventral bones, the ischium and pubis. +Very often a fourth pelvic element, the acetabular or cotyloid bone, +occurs. + +In the MONOTREMATA the pelvis is short and broad, and the pubes and +ischia meet in a long symphysis. The acetabulum is perforated in +_Echidna_ as in birds, but not in _Ornithorhynchus_. A pair of +elongated slender bones project forwards from the edge of the pubes +near the symphysis; these are sesamoid bones formed by ossifications +in the tendons of the external oblique abdominal muscles, and are +generally called _marsupial bones_. + +In the MARSUPIALIA the ilia are generally very simple, straight, and +narrow, while the pubes and ischia are well developed and meet in a +long symphysis. Marsupial bones are nearly always prominent, but are +not developed in _Thylacinus_ or _Notoryctes_. The ischium often has a +well-marked tuberosity and in Kangaroos the pubis bears a prominent +pectineal process on its anterior border close to the acetabulum. The +pelvis in _Notoryctes_ differs much from that in all other Marsupials, +the ilium and ischium being ankylosed with six vertebrae in a manner +comparable to that of many Edentates. + +In the EDENTATA the pelvis is generally well developed, but the +symphysis is very short. In the Sloths the pelvis is rather weak and +slender, the obturator foramina are very large and the ischia do not +meet in a symphysis. In the Megatheriidae the pelvis is exceedingly +wide and massive, and is firmly ankylosed with a number of vertebrae. +In the Armadillos, Glyptodonts, Anteaters, and Pangolins it is much +developed and firmly united to the vertebral column by both the ilia +and the ischia. In _Orycteropus_ however the ischium does not become +united to the vertebral column, and the pubis generally has a strongly +developed pectineal process. + +In the SIRENIA the pelvis is quite vestigial. In the Dugong it +consists on each side of two slender bones, one of which represents +the ilium and the other the ischium and pubis; the two bones are +placed end to end and are commonly fused together. The ilium is +attached by ligament to the transverse process of one of the +vertebrae. In the Manatee each half of the pelvis is represented by a +triangular bone connected by ligaments with its fellow and with the +vertebral column. In neither Manatee nor Dugong is there any trace of +an acetabulum but one can be made out in _Halitherium_. + +In the CETACEA the pelvis is even more vestigial than in the Sirenia, +consisting simply of a pair of small straight bones which probably +represent the ischia, and lie parallel to and below the vertebral +column at the point where the development of chevron bones commences. + +In UNGULATA VERA the pelvis is generally rather long and narrow. The +ilium is flattened and expanded in front (fig. 103, 8), but becomes +much narrower and more cylindrical before reaching the acetabulum. +Both pubis and ischium contribute to the symphysis which is often very +long. The ischia are large and have prominent tuberosities, especially +in Artiodactyles. In most Ruminantia there is a deep depression, the +supra-acetabular fossa above the acetabulum, but this is not found in +the Suina or Tylopoda. + +SUBUNGULATA. In _Procavia_ the pelvis is long and narrow, and bears +resemblance to that in Artiodactyles. + +The Proboscidea have a very large pelvis set nearly at right angles to +the vertebral column; the ilium is very wide, having expanded iliac[1] +and gluteal[1] surfaces, and a narrow sacral[176] surface. The pubes +and ischia are rather small, but both meet their fellows in the +symphysis. _Uintatherium_ (suborder Amblypoda) also has a large and +vertically placed pelvis (fig. 108) with a much expanded ilium. The +pelvis however differs from that of the Proboscidea in the fact that +the ischia do not meet in a ventral symphysis. + +In many RODENTIA the ilia have their gluteal, iliac, and sacral +surfaces of nearly equal extent; in the Hares, however, the gluteal +and iliac surfaces are confluent. The pubes and ischia are always well +developed and sometimes, as in the Hares, the acetabular bone also. In +these animals the pubis does not take part in the formation of the +acetabulum, and the ischium bears on its outer side a well-marked +ischial tuberosity. + +In the CARNIVORA the pelvis is long and narrow. The iliac surfaces +(fig. 78, A, 5) are very small and the sacral large; the crest or +supra-iliac border is formed by the union of the sacral and gluteal +surfaces. The symphysis is long and includes part of both pubis and +ischium. The ischial tuberosity (fig. 78, A, 10) is often well marked, +and sometimes as in _Viverra_ the acetabular bone is distinct. In the +Pinnipedia the pelvic symphysis is little developed, or sometimes not +developed at all, and the obturator foramina are remarkably large. + +In some INSECTIVORA such as _Galeopithecus_, there is a long pelvic +symphysis, in others such as _Erinaceus_ and _Centetes_, it is very +short, in others again such as _Talpa_ and _Sorex_, there is no pelvic +symphysis. The acetabular bone is exceptionally large in _Talpa_ and +_Sorex_. + +In CHIROPTERA the pelvis is small and narrow, and in the great +majority of cases the two halves do not meet in a ventral symphysis. +The pubis has a strongly developed pectineal process, which +occasionally unites with a process from the ilium enclosing a large +pre-acetabular foramen. + +PRIMATES. In Man and the Anthropoid Apes the pelvis is very large and +wide, and the ilium has much expanded iliac and gluteal surfaces. The +symphysis is rather short and formed by the pubis alone. The +acetabulum is deep and the obturator foramen large, and there is +frequently a well-marked ischial tuberosity. In the lower Anthropoidea +the ilium is long and narrow and has a small iliac surface. The +ischial tuberosities are large in the old world monkeys. + +[Illustration FIG. 108. LEFT ANTERIOR AND POSTERIOR LIMB AND LIMB +GIRDLE OF _Uintatherium mirabile_. The anterior limb is to the left, +the posterior to the right × 1/10. (From casts, Brit. Mus.) + + 1. ilium. + 2. head of femur. + 3. great trochanter. + 4. patella. + 5. fibula. + 6. tibia. + 7. second digit of pes. + 8. ungual phalanx of fifth digit of pes. + 9. calcaneum. + 10. postscapular fossa. + 11. prescapular fossa. + 12. coracoid process. + 13. humerus. + 14. radius. + 15. ulna. + 17. unciform. + 18. cuneiform. + 20. lunar. + 21. first metacarpal. + 22. fifth metacarpal.] + + +THE THIGH AND SHIN. + +In the MONOTREMATA the femur is short, rather narrow in the middle, +and expanded at each end. The great and lesser trochanters are large +and about equally developed, but there is no third trochanter. The +fibula is very large and is expanded at its proximal end, forming a +flattened plate much resembling an olecranon. The patella is well +developed. + +In the MARSUPIALIA there is no third trochanter to the femur, the +fibula is well developed but not the patella as a general rule. +_Notoryctes_ has a femur with a prominent ridge extending some little +way down the shaft from the great trochanter; the tibia has a +remarkably developed crest, and the fibula has its proximal end much +expanded and perforated; there is an irregularly shaped patella +closely connected with the proximal end of the tibia. + +EDENTATA. In the Sloths the leg bones are all long and slender. The +femur has no third trochanter, and the fibula is complete and nearly +equal in size to the tibia. In the Megatheriidae the leg bones are +extraordinarily massive, the circumference of the shaft of the femur +in _Megatherium_ equalling or exceeding the length of the bone. There +is no third trochanter in _Megatherium_. In most of the remaining +Edentata the leg bones are strongly developed. The femur in the +Armadillos and Aard Varks has a strong third trochanter, and the +tibia and fibula are both large and are commonly ankylosed together at +either end. The limb bones are very massive also in the Glyptodonts. + +SIRENIA. In no living Sirenian is there any trace of a hind limb, but +in _Halitherium_ a vestigial femur is found, which articulates with +the pelvis by a definite acetabulum. + +[Illustration FIG. 109. LEFT FEMUR OF AN OX (_Bos taurus_) (to the +left) AND OF A SUMATRAN RHINOCEROS (_R. sumatrensis_) (to the right). +× 1/6. (Camb. Mus.) + + 1. head. + 2. great trochanter. + 3. lesser trochanter. + 4. third trochanter. + 5. shaft. + 6. condyles.] + +In the Mystacoceti among the CETACEA small nodules of bone or +cartilage occur connected with the vestigial pelvis, and may represent +the femur and tibia. No trace of the skeleton of the hind limb is +known in the Odontoceti. + +In the UNGULATA VERA the femur is noticeable for the size of the great +trochanter (fig. 109, 2); there is no definitely constricted neck +separating the head from the rest of the bone, and the lesser +trochanter (fig. 109, 3) is not very prominent. All Perissodactyles +except the Chalicotheriidae show a strongly marked third trochanter, +but this is absent in all known Artiodactyles. The development of the +fibula in general corresponds to that of the ulna. In _Rhinoceros_, +_Macrauchenia_, _Tapirus_ and the Suina it is distinct and fairly well +developed; in the Tragulina on the other hand it is vestigial, being +reduced to the proximal end only. In the Ruminantia and Tylopoda also, +it is much reduced forming merely a small bone attached to the distal +end of the tibia, sometimes, as in the Red deer a slender vestige of +the proximal end also is preserved quite detached from the distal +portion; in the Horse this proximal portion is all that there is found +of the fibula. The progressive diminution of the fibula can be well +seen in the series of forms that are regarded as the ancestors of the +Horse. The patella of the Ungulata vera is well ossified, but +fabellae[177] are not usually found. + +SUBUNGULATA. Of the Toxodontia, _Toxodon_ has no third trochanter +while _Typotherium_ and _Astrapotherium_ have one. In the Condylarthra +the femur has well-marked lesser and third trochanters, and the fibula +and patella are well developed. In the Hyracoidea there is a slight +ridge on the femur in the place of the third trochanter, the fibula is +complete, but is generally fused to the tibia at its proximal end. + +Of the Amblypoda, _Coryphodon_ has a third trochanter, but +_Uintatherium_ has none; in this respect, in the vertical position and +general appearance (fig. 108) of the limb, and in the articulation of +the fibula with the calcaneum, the leg of _Uintatherium_ closely +approaches that of the Proboscidea. + +In the Proboscidea the femur is very long and straight, the +development of trochanters is slight, and the fibula though slender is +complete and articulates with the calcaneum. + +A third trochanter is found in the Tillodontia. + +In RODENTIA the femur is variable, the great trochanter is generally +large and so sometimes is the third as in the Hares. In most Rodents +as in the Beaver the fibula is distinct, sometimes as in the Hares it +is united distally with the tibia. The patella is well developed, and +so too are the fabellae as a general rule. + +CARNIVORA. In the Carnivora vera the femur (fig. 79, A) is generally +rather straight and slender, and has a very distinct head. The fibula +(fig. 79, C) is always distinct and there is generally a considerable +interval between it and the tibia. Fabellae (fig. 79, 7) are commonly +present. + +In the Pinnipedia the femur is short, broad and flattened, having a +prominent great trochanter. The fibula is nearly as large as the +tibia, and the two bones are generally ankylosed together at their +proximal ends. + +The Creodonta differ from all living Carnivores in having a femur with +a third trochanter. + +In the INSECTIVORA a third trochanter is sometimes developed. The +fibula is sometimes distinct, sometimes fused distally with the tibia, +thus differing from that of a Carnivore. + +In CHIROPTERA the femur is straight, slender and rather short, with a +small but well-developed head. The fibula may be well developed or +quite vestigial or absent. Owing to the connection of the hind limb +with the wing membrane the knee joint is directed backwards. + +In PRIMATES the femur is rather long and slender, having a nearly +spherical head and large great trochanter. The tibia and fibula are +always distinct and well developed. Fabellae are not found in the +highest forms but are generally present in the others. + + +THE PES. + +The skeleton of the pes is in most respects a counterpart of that of +the manus. Just as in the manus if one digit is absent it is the +pollex, so in the pes it is the hallux. But while in the manus the +third digit is always well developed, however much the limb may be +modified, in the pes any of the digits may be lost. In all mammals the +tibiale and intermedium fuse to form the _astragalus_, and the fourth +and fifth tarsalia to form the _cuboid_. Sesamoid bones are +considerably developed. In almost every case the phalanges and first +metatarsal have epiphyses only on their proximal ends, while the +remaining four metatarsals have epiphyses only on their distal ends. + +In the MONOTREMATA all the usual tarsal bones are distinct, and the +five digits have the normal number of phalanges. Several sesamoid +bones are developed, the most important one, found only in the male, +being articulated to the tibia and bearing the curious horny spur. The +ungual phalanges of the pes like those of the manus, are deeply cleft +at their extremities. In the Echidnidae the pes is turned outwards and +backwards in walking. + +In the MARSUPIALIA the pes is subject to great modifications, but in +every case the seven usual tarsal bones are distinct. In the +Didelphyidae the foot is broad, all five digits are well developed, +and the hallux is opposable to the others. In the Dasyuridae the foot +is narrow, and the hallux may be very small, or as in _Thylacinus_ +completely absent. In _Notoryctes_ the pes is much less abnormal than +the manus, and all five digits have the usual number of phalanges. The +fifth metatarsal has a curious projecting process, and there is a +large sesamoid above the hallux. In the Wombats (Phascolomyidae) the +foot is short and broad, the digits are all distinct, and the hallux +is divaricated from the others. + +In the remaining marsupials the second and third metacarpals and +digits are very slender, and are enclosed within a common integument. +This condition is known as _syndactylism_, and its effect is to +produce the appearance of one toe with two claws. In the Kangaroos +(Macropodidae) the pes is very long and narrow, owing to the +elongation of the metacarpals. The fourth digit is greatly developed, +the fifth moderately so, while the hallux is absent, and the second +and third digits are very small. The Peramelidae have the foot +constructed on the same plan as in the Kangaroos, and in one genus +_Choeropus_ the same type of foot is carried to a greater extreme than +even in the Kangaroos. Thus the fourth digit is enormously developed, +the second and third are small, and the fifth smaller still, while the +hallux is absent. In the Phalangers and Koalas though the second and +third toes are very slender, the hallux is well developed and +opposable. + +EDENTATA. In the Sloths the pes much resembles the manus, being long +and narrow, but in both genera the second, third and fourth digits are +well developed. Most of the other Edentates have a but little modified +pes with the normal number of tarsal bones and the complete series of +digits. In _Cycloturus_ however the hallux is vestigial and it is +absent in Glyptodonts. _Megatherium_ has a greatly modified pes, the +hallux is absent, and the second digit vestigial, while the third is +very large, having an enormous ungual phalanx. The calcaneum too is +abnormally large. + +No trace of the pes occurs in either SIRENIA or CETACEA. + +In the UNGULATA the pes like the manus is subject to much variation +and is of great morphological importance. + +In the UNGULATA VERA the pes is never plantigrade and never has more +than four digits, the hallux being absent. The cuboid always +articulates with the astragalus, and the tarsal bones strongly +interlock. As was the case also with the manus, the pes is formed on +two well-marked types characteristic respectively of the Artiodactyla +and Perissodactyla. + +_ARTIODACTYLA._ Just as in the manus, the third and fourth digits are +well and subequally developed; their ungual phalanges have the +contiguous sides flat, and the axis of the limb passes between them, +and between the cuboid and navicular. The astragalus has both the +proximal and distal surfaces pulley-like, and articulates with the +navicular and cuboid by two facets of nearly equal size. The calcaneum +articulates with the lower end of the fibula if that bone is fully +developed. + +In the Suina four toes are developed, and though in the Peccaries the +third and fourth metatarsals are united, they are all distinct in most +members of the group, as are all the tarsal bones. In the Hippopotami +the four digits are of approximately equal size, and the middle ones +do not have the contiguous faces of their ungual phalanges flattened. + +In the Tragulina the cuboid, navicular, and two outer cuneiforms are +united forming a single bone; all four metatarsals are complete and +the two middle ones are united. In the Tylopoda and _Anoplotherium +commune_ only the third and fourth digits are developed, their +metatarsals are free distally, but are elsewhere united. In the +Ruminantia the cuboid and navicular are always united and so are the +second and third cuneiforms, while in _Cervulus_ all four bones are +united together. The third and fourth metatarsals in Ruminants are +always united in the same way as are the third and fourth metacarpals, +while the second and fifth are always wanting. In Deer the second and +fifth digits are usually each represented by three small phalanges, +but in the Giraffe and most Bovidae the bones of these digits are +wanting. + +[Illustration + + FIG. 110. _A._ LEFT PES OF A TAPIR (_Tapirus americanus_). × 1/6. + _B._ RIGHT PES OF A RHINOCEROS (_R. sumatrensis_). × 1/8. + _C._ (CAST OF) RIGHT PES OF _Hipparion gracile_. × 1/7. + _D._ RIGHT PES OF A HORSE (_Equus caballus_). × 1/10. (All Camb. Mus.) + + 1. calcaneum. + 2. astragalus. + 3. navicular. + 4. cuboid. + 5. external cuneiform. + 6. middle cuneiform. + 7. internal cuneiform.] + +In the _PERISSODACTYLA_ the pes like the manus is symmetrical about a +line drawn through the third digit; this line when continued passes +through the external cuneiform, navicular and astragalus. The +astragalus has its distal portion abruptly truncated, and the facet by +which it articulates with the cuboid is much smaller than that by +which it articulates with the navicular. The calcaneum does not +articulate with the fibula. The tarsus in _Macrauchenia_ like the +carpus differs from that of other Perissodactyles and resembles that +of Subungulates in having the bones arranged in lines with little or +no interlocking. The calcaneum resembles that of Artiodactyles in +having a small facet for articulation with the fibula. _Tapirus_ (fig. +110, A), _Rhinoceros_ (fig. 110, B) and _Titanotherium_ have a short +and broad foot with the usual tarsal bones and three well-developed +digits,--a number never exceeded by any Perissodactyle. From this +tridactylate limb a series of stages is exhibited by various extinct +forms leading gradually to the condition met with in the Horse (fig. +110, D) in which the third toe is greatly developed, while the second +and fourth are reduced to slender metatarsals attached to the proximal +half of the third metatarsal. + +In _Chalicotherium_ and _Agriochoerus_ the pes has the same abnormal +characters as the manus, the digits being clawed and the ungual +phalanges in _Chalicotherium_ deeply cleft. + +In the SUBUNGULATA the pes is sometimes plantigrade and +pentedactylate, the cuboid sometimes does not articulate with the +astragalus, and the tarsal bones sometimes do not interlock. + +In _Typotherium_ (_TOXODONTIA_) the hallux is absent and the other +four digits are well developed; in _Toxodon_ and _Nesodon_ the pes is +tridactylate. The tarsal bones have the regular Subungulate +arrangement, the cuboid not articulating with the astragalus. The +calcaneum articulates with the fibula as in Artiodactyles. The +astragalus in most forms, but not in _Astrapotherium_, resembles that +of the Ungulata vera in having a grooved proximal surface. + +In _Phenacodus_ (_CONDYLARTHRA_) the tarsus is very little modified, +five digits are present, the first and fifth being small and not +reaching the ground. + +In _Procavia_ only the three middle digits are present with a vestige +of the fifth metacarpal. + +In the _AMBLYPODA_ the pes (fig. 108) is very short and broad, all +five digits are functional, and at any rate in _Coryphodon_ +plantigrade, the hallux being the smallest. The astragalus is very +flat, and the tarsals interlock to a slight extent, the cuboid +articulating with both calcaneum and astragalus. + +The pes in the _PROBOSCIDEA_ much resembles that in the Amblypoda, but +differs in that the astragalus does not articulate with the cuboid, +the tarsals not interlocking at all. + +In the RODENTIA the structure of the foot is very variable. In Beavers +the foot is very large, all five digits being well developed; the +fifth metatarsal articulates with the outer side of the fourth +metatarsal, and not with the cuboid, and there is a large sesamoid +bone on the tibial side of the tarsus. In the Rats, Porcupines and +Squirrels, there are five digits, in the Hares only four, and in the +Capybara and some of its allies only three. In the Jerboa (_Dipus_) a +curious condition of the pes is met with, as it consists of three very +long metatarsals fused together and bearing three short toes, each +formed of three phalanges. _Lophiomys_ differs from all other Rodents +in having the hallux opposable. + +CARNIVORA. In the Carnivora vera the pes is regular and shows little +deviation from the normal condition. All the usual tarsal bones are +present, but sometimes as in the Dogs, Cats, and Hyaenas, the hallux +is vestigial. Sometimes as in the Bears the pes is plantigrade, +sometimes as in the Cats and Dogs it is digitigrade. In this respect +and in the character of the ungual phalanges, the pes closely +corresponds with the manus. In the Sea Otter (_Latax_) the foot is +large and flattened and approaches in character that of the +Pinnipedia. + +In the Pinnipedia the pes differs much from that in the Carnivora +vera. In the Seals in which the foot cannot be used for walking, and +is habitually directed backwards, the first and fifth digits are much +longer and stouter than any of the others. In the Sea Lions which can +use the pes for walking, the digits are all of nearly the same length, +and in the Walrus the fifth is somewhat the longest. + +In the INSECTIVORA the pes is almost always normal, and provided with +five digits. + +In the CHIROPTERA the pes is pentedactylate, and the digits are +terminated by long curved ungual phalanges. In some genera the toes +have only two phalanges. The calcaneum is sometimes produced into a +long slender process which helps to support the membrane between the +leg and the tail. + +Among the PRIMATES Man has the simplest form of pes. In Man all five +digits are well developed, the hallux being considerably the largest. +Sesamoid bones occur only under the metatarso-phalangeal joint of the +hallux. + +In the other Primates the internal cuneiform has a saddle-shaped +articulating surface for the hallux, which is obliquely directed to +the side of the foot and opposable to the other digits. Two sesamoid +bones are usually developed below each metatarso-phalangeal joint, and +one below the cuboid. The second digit in Lemurs, and all except the +hallux in _Chiromys_ have pointed ungual phalanges; in all other cases +the ungual phalanges are flat. In some of the Lemuroidea, especially +_Tarsius_, the tarsus is curiously modified by the elongation of the +calcaneum and navicular. + + +FOOTNOTES: + +[165] The figure was drawn from a photograph and the size of the jaws +relatively to the cranium is exaggerated. + +[166] See W.K. Parker, "On the Structure and Development of the Skull +in the Pig." _Phil. Trans._ pp. 289-336, 1874. + +[167] See W.H. Flower, "On the value of the characters of the base of +the cranium in the classification of the order Carnivora." _P.Z.S._ +1869, p. 5. + +[168] See W.K. Parker, _Monograph of the shoulder-girdle and sternum +of the Vertebrata_, _Ray Soc._ 1868. + +[169] See p. 405. + +[170] See E. Lydekker, _P.Z.S._ 1895, p. 172. + +[171] See H. Wincza, _Morph. Jahr._ XVI. p. 647, 1890. + +[172] See K. Bardeleben, _P.Z.S._, 1889, p. 259. + +[173] See E. Cope, "The origin of the foot structures of Ungulata," +_Journ. of Philad. Acad._ 1874. H.F. Osborn, "The evolution of the +Ungulate foot," _T. Amer. Phil. Soc._ 1889. + +[174] See O.C. Marsh, various papers including "Fossil horses in +America," _Amer. Natural._ 1874; "Polydactyl horses," _Amer. J. Sci._ +1879 and 1892. M. Pavlow, "Le développement des Equidés," _Bul. Soc. +Moscou_, 1887, and subsequent papers in the same. Osborn and Wortman, +"On the Perissodactyls of the White River beds," _Bull. Amer. Mus._ +Dec. 23rd, 1895. + +[175] See H.F. Osborn, _Chalicotherium and Macrotherium, Amer. +Natural._ 1889--91--92. + +[176] See p. 409. + +[177] See p. 412. + + + + +LIST OF AUTHORS REFERRED TO. + + + Abbott, E.C., 112 + + Ameghino, F., 351, 424 + + Andrews, C.W., 299 + + + Balfour, F.M., 16 + + Ballowitz, E., 424 + + Bardeleben, K., 504 + + Bateson, W., 50, 344 + + Baum, H., 374 + + Baur, G., 27, 189, 190, 344, 346 + + Beneden, P.J. van, 353 + + Benham, W.B., 51 + + Bettany, G.T., 16, 87, 154 + + Boulenger, G.A., 169 + + Brandt, J.F., 352 + + Bridge, T.W., 123 + + Brühl, C.B., 210 + + Burmeister, H., 351, 424 + + + Cope, E.D., 135, 199, 204, 351, 359, 361, 363, 368 + + Credner, H., 135 + + + Dean, B., 63, 104 + + Dobson, G.E., 369, 370 + + + Earle, C., 432 + + Ecker, A., 151 + + Ellenberger, W., 374 + + + Flower, W.H., 28, 42, 351, 420, 422, 434 + + Fritsch, A., 135 + + Fürbringer, M., 295 + + + Gadow, H., 40, 112, 190, 295, 343, 350 + + Gegenbaur, C., 127 + + Gervais, P., 353 + + Günther, A.C.L.G., 70, 104 + + + Haslam, G., 151 + + Hasse, C., 112, 113 + + Haswell, W.A., 127 + + Hertwig, O., 169 + + Hoffmann, C.K., 190, 202, 210 + + Howes, G.B., 164, 451 + + Hubrecht, A.A.W., 104 + + Hulke, J.W., 192, 204 + + Hurst, C.H., 71, 297 + + Hutton, F.W., 299 + + Huxley, T.H., 11, 13, 133, 135, 191, 210, 295, 297, 334, 343, 351, + 374, 437 + + + Kirkaldy, J.W., 51 + + Klein, E., 11 + + Kölliker, A., 9 + + Kükenthal, W., 349, 422 + + + Lankester, E. Ray, 51 + + Leche, W., 344, 423 + + Lindsay, B., 336 + + Lydekker, R., 36, 42, 190, 195, 495 + + + Macbride, E.W., 50 + + Marsh, O.C., 204, 209, 299, 348, 361, 364, 365, 508 + + Marshall, A.M., 71, 151 + + Masterman, A.T., 51 + + Meyer, H. v., 135 + + Miall, L.C., 135, 243 + + Mivart, St G., 369 + + Morgan, C. Lloyd, 11 + + + Newton, E.T., 283 + + + Osborn, H.F., 348, 420, 429, 508 + + Owen, R., 191, 204, 210, 297, 348, 351, 420 + + + Parker, T.J., 83, 96, 299 + + Parker, W.K., 16, 24, 53, 87, 154, 173, 200, 243, 465, 489 + + Pavlow, M., 358, 508 + + Pollard, H.B., 119 + + Poulton, E.B., 422 + + Pycraft, W.P., 297 + + + Ridewood, W.G., 106, 164 + + Röse, C., 422 + + + Sagemehl, M., 104 + + Schäfer, E., 11 + + Scott, W.B., 368 + + Seeley, H.G., 191, 212 + + Selenka, E., 40, 295 + + Shufeldt, R., 123 + + Smith, E. Noble, 11 + + Stirling, E.C., 423 + + Swirski, G., 103 + + + Taeker, J., 427 + + Thomas, O., 349, 362, 370, 422, 424, 425 + + Tomes, C.S., 420 + + Traquair, R.H., 55, 58 + + + Vogt, C., 297 + + + Wiedersheim, R., 25, 134, 136 + + Wincza, H., 358, 495 + + Woodward, A. Smith, 34, 54, 58, 62, 127, 210 + + Wortman, J.L., 508 + + Wray, R.S., 303 + + + Zittel, K.A. v., 36, 205, 212 + + + + +INDEX. + + + Every reference is to the page: words in italics are names of + genera or species; figures in italics indicate that the + reference relates to systematic position; figures in thick type + refer to an illustration; _f._ = and in following page or + pages; _n._ = note. + + + Aard Vark, _44_, _352_; + femur, 517; + sacrum, 452; + teeth, 425; + see _Orycteropus_ + + Aard wolf, _48_; + see _Proteles_ + + Abdominal ribs, crocodile, 260; + reptiles, 286 + + Abdominal shield, turtle, 215 + + _Acanthias_, _32_; + calcification of vertebrae, 114; + pectoral fins, 130 + + _Acanthodes_, _32_, _64_ + + Acanthodii, _32_; + general characters, 64; + spines, 106 + + _Acanthomys_, _47_; + spines, 417 + + Acanthopterygii, _34_ + + Accipitres, _41_ + + Acetabular bone, 25, 513; + dog, 409 f.; + frog, 165 + + Acetabulum, 25; + crocodile, 266; + dog, 409; + duck, 324; + frog, 165; + newt, 149; + turtle, 235 + + _Acipenser_, _32_, 117; + exoskeleton, 67; + distribution, 66; + pectoral fins, 131; + plates, 104; + skull, 121, =122=; + spinal column, 112 + + Acipenseridae, _32_ + + Acrodont, defined, 199; + teeth of reptiles, 273 + + _Acrodus_, _32_; + teeth, 109 + + Acromion, dog, 405 + + _Actinotrocha_, _30_; + organ regarded as double notochord, 51 + + Ad-digital quill, duck, 303 + + Adjutant, _41_; + clavicles, 338 + + Ægithognathous, 335 + + Æluroidea, _48_, _369_; + teeth, 437 + + Æpyornis, _40_; + tibio-tarsus, 341 + + Æpyornithes, _40_, _299_ + + Aftershaft, 328 + + _Agama_, _38_; + teeth, 273 + + Agamidae, _38_; + premaxillae, 284 + + Aglossa, _36_ + + Agouti, _48_; + see _Dasyprocta_ + + Agriochoeridae, _45_ + + _Agriochoerus_, _45_; + pes, 525 + + Ala spuria, duck, 304 + + Alcidae, _42_; + see Auks + + Alisphenoid, 19; + crocodile, 247; + duck, 317; + dog, 386 + + _Alligator_, _39_, _210_, _212_; + hyoid, =285=; + limbs, =264=; + pectoral girdle, =262=; + pelvis and sacrum, =267=; + scutes, 271; + skull, =245=, =248=, =253= + + Alligatoridae, 39 + + _Alytes_, _36_; + fronto-parietal fontanelle, 179; + vertebrae, 172 + + Amblypoda, _47_; + general characters, 363; + manus, 510; + pes, 525; + skull, 473; + teeth, 433; + thigh and shin, 519 + + _Amblystoma_, _35_; + skull, 175 + + American monkeys, _373_; + see _Cebidae_ + + American vultures, _41_; + vomers, 335 + + _Amia_, _33_; + distribution, 66; + exoskeleton, 67; + pectoral fin, 131; + scales, 105; + skull, 123; + tail, 115, 117; + vertebrae, 114 + + Amiidae, _33_ + + _Ammocoetes_, _31_, _55_ + + Amphibia, _35_; + anterior limb, 185; + exoskeleton, 168; + general characters, 133; + hyoid apparatus, 180; + pectoral girdle, 184; + pelvic girdle, 187; + posterior limb, 188; + ribs, 182; + skull, 173; + sternum, 182; + teeth, 169; + vertebral column, 170 + + Amphicoelous, defined, 14 + + _Amphioxus_, _30_; + skeleton, 51 f.; + spinal column, 112 + + _Amphisbaena_, _38_, 272; + loss of limbs, 289 + + Amphisbaenidae, _38_, _200_; + pectoral girdle, 288; + skull, 277; + vertebral column, 275 + + Amphitheriidae, _43_ + + _Amphiuma_, _35_, _135_; + manus, 187; + pes, 188; + skull, 174 + + Amphiumidae, _35_ + + Anacanthini, _33_ + + Anal shield, turtle, 215 + + _Anas_, _41_; + _A. boschas_, see Duck + + Ankylosis, defined, 12 + + Angel fish, _32_ + + Angler, attachment of teeth, 107 + + Anguidae, _38_ + + _Anguilla_, _33_; + see Eel + + _Anguis_, _38_; + loss of limbs, 289; + scutes, 271 + + Angular, 22; + cod, 100; + crocodile, 258; + duck, 319; + salmon, 94; + turtle, 231 + + Angulo-splenial, frog, 161 + + Ankle joint, duck, 327; + reptiles, 294 + + Anomodontia, _36_ + + Anoplotheriidae, _45_ + + _Anoplotherium_, _45_; + manus, 506; + pes, 523; + tail, 454; + teeth, 428 + + _Anser_, _41_ + + Anseres, _41_; + aftershaft, 329; + claws, 330 + + Anseriformes, _41_ + + Anteaters, _352_; + absence of teeth, 424; + manus, 505; + pectoral girdle, 495; + pelvis, 513; + skull, 458; + thoraco-lumbar vertebrae, 447; + Spiny --, _43_; + Great and Two-toed --, _44_ + + Antelope, _359_; + manus, 507; + Four-horned A., _46_ + + Anterior limb, 26; + Amphibia, 185; + birds, 338; + crocodile, 263; + dog, 405; + duck, 322; + frog, 164; + newt, 147; + reptiles, 290; + turtle, 232 + + Anthropoid apes, _373_; + arm-bones, 503; + pelvis, 515 + + Anthropoidea, 49; + general characters, 372; + sacrum, 452; + skull, 482; + teeth, 441 + + _Anthropopithecus_, _49_; + ribs, 493 + + Antiarcha, _31_; + general characters, 55 + + Antibrachium, see fore-arm + + _Antilocapra_, _46_; + horns, 417 + + Antilocapridae, _46_ + + Antitrochanter, duck, 325 + + Antlers, 8, 358; + Cervidae, 469 + + Antorbital process, 18 + + Anura, _36_; + general characters, 136; + hyoid apparatus, 180; + pelvis, 187; + posterior limb, 188; + skull, 179; + sternum, 182; + vertebrae, 172 + + _Apatornis_, _40_; + vertebrae, 332 + + Apteria, 328 + + Apteryges, _40_ + + _Apteryx_, _40_, _299_; + aftershaft, 329; + anterior nares, 333; + claws, 330; + foot, 342; + manus, 338; + pectineal process, 341; + pectoral girdle, 338; + pneumaticity of skeleton, 331; + _A. oweni_, pelvic girdle and sacrum, =340= + + Aqueductus vestibuli, dogfish, 74 + + Arcade: + infratemporal--, crocodile, 255; + _Sphenodon_, 283; + inner --, duck, 318; + outer --, duck, 318; + supratemporal --, crocodile, 257; + reptiles, 281 + + Archaeoceti, _44_; + general characters, 356; + skull, 461; + teeth, 426 + + _Archaeopteryx_, _40_, _297_; + claws, 330; + fibula, 341; + mandible, 335; + metatarsals, 342; + pelvis, 341; + ribs, 336; + sacrum, 333; + skull, 333; + tail, 333; + teeth, 330; + wing, 338 + + Archaeornithes, _40_; + characters, 297 + + _Archegosaurus_, _35_, _136_; + palatines, 177 + + Archipterygium, _Ceratodus_, 127; + Ichthyotomi, 62 + + Arcifera, _36_, 185 + + Arctoidea, _48_, 369; + teeth, 438 + + _Ardea_, _41_; + see Heron + + Ardeae, _41_ + + Arm, see fore-arm and upper arm + + Armadillo, _44_, _352_; + cervical vertebrae, 443; + femur, 517; + humerus, 501; + lumbar vertebrae, 447; + manus, 505; + pectoral girdle, 495; + pelvis, 513; + ribs, 491; + sacrum, 452; + scales, 417; + scutes, 419; + skull, 459; + teeth, 424 + + Armour plates, 8 + + Arthrodira, _34_; + characters, 70 + + Articular, 22; + cod, 100; + crocodile, 258; + duck, 319; + newt, 144; + salmon, 94; + turtle, 231 + + Artiodactyla, _45_; + characters, 358; + manus, 506; + odontoid process, 445; + pes, 522; + ribs, 491; + skull, 465; + teeth, 427; + thoraco-lumbar vertebrae, 448 + + Asses, _360_ + + _Asterolepis_, _31_, _55_ + + Asterospondyli, 114 + + Astragalus, 27; + crocodile, 268; + dog, 414; + mammals, 521 + + Astrapotheriidae, _46_ + + _Astrapotherium_, _46_, 361; + dental formula, 432; + femur, 519; + pes, 525 + + _Ateles_, _49_; + pollex, 512; + tail, 454 + + Atlantosauridae, _38_ + + Atlas, 15; + crocodile, 240; + dog, =379=, 380; + duck, 309; + ox, =445=; + turtle, 219 + + Attachment of teeth, 4; + in fish, 107 + + _Auchenia_, _45_; + see Llama + + Auditory aperture or meatus: + external --, crocodile, 250; + dog, 402; + turtle, 228; + internal --, crocodile, 246, 251; + dog, 392; + turtle, 228 + + Auditory capsule, 20; + cod, 96; + crocodile, 250; + dog, 390; + dogfish, 74; + frog, 156; + newt, 143; + turtle, 227 + + Auditory ossicles, crocodile, 251; + dog, 393; + duck, 320; + mammals, 485 f.; + turtle, 228 + + Auks, _42_; + thoracic vertebrae, 332 + + Autostylic, 61, 119 + + Aves, _40_; + characters, 295; + see Birds + + Axial skeletal rods, 50 + + Axial skeleton, crocodile, 239; + cod, 83; + dog, 377; + dogfish, 72; + duck, 307; + frog, 152; + newt, 138; + turtle, 218 + + Axis vertebra, crocodile, 241; + dog, 380; + duck, 309; + turtle, 220 + + Axolotl, _35_; + see _Siredon_ + + Aye Aye, _49_; + see _Chiromys_ + + + _Babirussa_, _45_; + dental formula, 428 + + Baboon, _49_; + see _Cynocephalus_ + + _Balanoglossus_, _30_, 50 + + _Balaena_, _44_, _357_; + scapula, 495; + _B. mysticetus_ baleen, 419 + + Balaenidae, _44_ + + Balaenoidea, _44_; + general characters, 356 + + _Balaenoptera_, _44_, 357; + manus, 506; + thoracic vertebrae, 448; + scapula, 495; + _B. musculus_, cervical vertebrae, =444= + + Baleen, 3, 418 + + _Balistes_, _33_; + teeth, 111 + + Balistidae, _33_ + + Ball and socket joints, 13 + + Bandicoot, _43_ + + Barb, 302 + + Barbule, 303 + + Barramunda, _34_; + see _Ceratodus_ + + Basalia, dogfish, 79 + + Basibranchial, dogfish, 78; + cod, 101; + duck, 320; + newt, 145; + salmon, 95 + + Basi-branchiostegal, cod, 101; + salmon, 95 + + Basicranial axis, 19; + dog, 384 + + Basidorsalia, dogfish, 72 + + Basi-hyal, dogfish, 78; + dog, 399; + duck, 320 + + Basilar plate, 17 + + Basilingual plate, Anura, 180; + crocodile, 259; + frog, 161; + turtle, 231 + + Basi-occipital, 19; + crocodile, 246; + cod, 97; + dog, 386; + duck, 315; + salmon, 89; + turtle, 224 + + Basipterygium, cod, 103; + dogfish, 82 + + Basisphenoid, 19; + crocodile, 247; + dog, 386; + salmon, 91; + turtle, 225 + + Bastard wing, duck, 304 + + Bathyerginae, palate, 366 + + _Bathyergus_, _47_; + auditory ossicles, 488; + manus, 511 + + Batoidei, _32_, 64 + + _Batrachoseps_, _35_; + teeth, 169 + + Bats, claws, 418; + Horseshoe bats, _49_; + see Chiroptera + + _Bdellostoma_, _31_, _55_; + teeth, 57 + + Beak, 3; + birds, 329; + duck, 302; + _Siren_, 168; + tadpoles of Anura, 168; + turtle, 215 + + Bears, _48_, _369_; + manus, 511; + pes, 526; + sacral vertebrae, 452; + skull, 479; + Isabelline -- mandible, =438= + + Beaver, _47_; + fibula, 520; + humerus, 502; + pes, 526; + sacrum, 452; + tail, 454 + + _Belodon_, _39_, 211; + frontals, 277; + palate, 281; + vertebrae, 275 + + Bichir, _33_; + see _Polypterus_ + + Bicipital groove, dog, 405 + + Bilophodont, defined, 345; + teeth of Tapiridae, 429 + + _Bipes_, _38_; + limbs, 289 + + Birds, anterior limb, 338; + endoskeleton, 331 f.; + exoskeleton, 328 f.; + general characters, 295; + hyoid, 336; + pectoral girdle, 336; + pelvic girdle, 339; + posterior limb, 341; + ribs, 336; + skull, 333; + sternum, 336; + teeth, 330; + vertebral, 332 + + _Bison_, _46_; + occipital crest, 468 + + Blind snake, _38_; + see _Typhlops_ + + Blind worm, _38_; + see _Anguis_ + + Boidae, _38_ + + _Bombinator_, _36_; + vertebrae, 172 + + Bone, development of, 10 f. + + Bone cells, 10 + + Bony Ganoids, fins, 105; + pelvic fin, 132; + ribs, 126; + skull, 123; + vertebral column, 114; + see _Holostei_ + + Border: + alveolar --, of dog's jaw, 398; + coracoid, glenoid, and suprascapular -- of dog's scapula, 405 + + _Bos_, _46_; + occipital crest, 468; + ribs, 491; + see Ox + + Bottlenose, _44_; + see _Hyperoödon_ + + Bovidae, _46_; + pes, 523; + skull, 468 + + Bow-fin, _33_; + see _Amia_ + + Brachial ossicles, cod, 103 + + Brachium; see upper arm + + Brachydont, defined, 345; + teeth of Ungulates, 429 f. + + _Brachycephalus_, _36_; + bony plates of, 168 + + Brain case, crocodile, 245; + dog, 384; + duck, 314; + frog, 154; + newt, 140; + turtle, 224 + + Bradypodidae, _43_; + see Sloths + + _Bradypus_, _43_; + cervical vertebrae, 443; + pectoral girdle, 495; + skull, 457; + thoraco-lumbar vertebrae, 447 + + Branchial arches, Amphibia, 180 f.; + cod, 101; + dogfish, 78; + fish, 120 f.; + newt, 145; + salmon, 95; + -- basket, Marsipobranchii, 38; + -- skeleton, _Amphioxus_, 52; + Balanoglossus, 50 + + _Branchiosaurus_, _35_; + branchial arches, 180 + + Branchiostegal rays, cod, 100 + + _Brontops_, _46_; + see _Titanotherium_ + + _Brontosaurus_, _38_, _207_; + sternum, 288 + + _Bubalus_, _46_; + ribs, 491; + see Buffalo + + Buccal skeleton, _Amphioxus_, 52 + + _Buceros_, _42_; + see Hornbill + + Buckler, of Labyrinthodonts, 168, 184 + + Buffalo, _46_; + Cape --, skeleton of, =492= + + _Bufo_, _36_; + hyoid, 182; + jaws, 169; + _B. viridis_, carpus, 186 + + Bufonidae, _36_ + + Bunodont, defined, 345; + teeth of Ungulata, 427 f. + + Buno-selenodont, defined, 432 + + + Caeciliidae, _35_ + + _Caiman_, _39_; + _C. latirostris_ hyoid, =285=, + limbs, =264=, + lateral view of skull, =248=, + palatal view of cranium and mandible, =245=, + longitudinal section of skull, =253=, + pectoral girdle, =262=, + pelvic girdle and sacrum, =267=; + _C. sclerops_, scutes, 271 + + Ca'ing whale, _45_; + see _Globicephalus_ + + _Calamoichthys_, _33_; + distribution, 66 + + Calamus, 302 + + Calcaneum, 27; + crocodile, 268; + dog, 414 + + Calcar, of frog, 167 + + _Callorhynchus_, _32_, _66_; + teeth, 110 + + Camel, _45_, _359_; + manus, 507; + teeth, 428 + + Camelidae, _45_ + + _Camelus_, _45_; + see Camel + + Camptosauridae, _39_ + + Canal: + alisphenoid --, dog, 402; + carotid --, duck, 315; + Eustachian --, crocodile, 247; + dog, 402; + duck, 316; + interorbital --, dogfish, 76 + + Canaliculi, 10 + + Canidae, _48_; + humerus, 502; + skull, 479; + see Dog + + Canine, 344; + dog, 376 f. + + _Canis_, _48_; + thoraco-lumbar vertebrae, 450; + see Dog + + _Capitosaurus_, _35_; + skull, =176= + + Capybara, _48_; + manus, 511; + pes, 526; + skull, 476; + tail, 454 + + Carapace, Chelonia, 271; + _Dermochelys_, 272; + Glyptodonts, 419; + Green turtle, 215; + Loggerhead turtle, =216= + + Carcharidae, _32_ + + Carina sterni, duck, 321 + + Carinatae, _40_; + general characters, 300; + quadrate, 334 + + Carnassial teeth, 368; + carnivora, =436=; + dog, 376 f. + + Carnivora, _48_; + arm bones, 502; + auditory ossicles, 488; + cervical vertebrae, 446; + general characters, 367; + manus, 511; + pelvis, 515; + pes, 526; + ribs, 493; + sacral vertebrae, 452; + skull, 478; + sternum, 490; + tail, 454; + teeth, 437; + thigh and shin, 520; + thoraco-lumbar vertebrae, 450 + + Carnivora vera, _48_; + general characters, 368; + scapula, 497 + + Carp, _33_; + pharyngeal teeth, 111 + + Carpo-metacarpus, duck, 324 + + Carpus, 26; + crocodile, 265; + dog, 408; + duck, 323; + frog, 164; + newt, 147; + turtle, 233 + + Cartilage, structure of, 10 + + Cartilaginous ganoids, cranium, 121; + pelvic fin, 132; + spinal column, 112; + see Chondrostei + + Cassowary, _40_, _299_; + aftershaft, 328; + bony crest, 334; + claws, 330; + pelvic girdle and sacrum, =340=; + secondaries, 329 + + _Castor_, _47_; + see Beaver + + Castoridae, _47_ + + _Casuarius_, _40_; + see Cassowary + + Cataphracti, _34_ + + Cat, _48_, _369_; + hallux, 526; + manus, 511; + skull, 479 + + Cat-fish, _33_ + + Cathartae, _41_ + + _Cathartes_, _41_; + see American vulture + + Caudal fin, Cetacea, 453; + fish, 116; + -- vertebrae, crocodile, 243; + cod, 85; + dog, 383; + duck, 312; + general characters, 16; + newt, 140; + turtle, 222 + + _Cavia_, _48_; + tail, 454 + + Caviidae, _48_ + + Cebidae, _49_, _373_; + ribs, 493; + skull, 484; + teeth, 441 + + _Coenolestes_, _43_, 424 + + Cement, 5 + + _Centetes_, _49_; + caudal vertebrae, 454; + pelvic symphysis, 515; + spines, 417; + teeth, 440; + thoraco-lumbar vertebrae, 450 + + Centetidae, _49_; + auditory ossicles, 488; + skull, 480 + + Centrale, 27; + see Carpus and Tarsus + + Centre of motion, 448 + + Centrum, 14 + + _Cephalaspis_, _31_, _55_ + + Cephalic shield, armadillos, 419 + + _Cephalochordata_, _30_, 51 + + _Cephalodiscus_, _30_, 50 + + _Ceratodus_, _34_, _70_; + branchial arches, 124; + cranium, =125=; + skeleton, =128=; + skull, 117, 124; + spinal column, 113; + teeth, 111 + + Cerato-branchial, cod, 101; + dogfish, 78; + duck, 320; + salmon, 95 + + Cerato-hyal, 23; + cod, 100; + dog, 399; + dogfish, 78; + salmon, 95 + + _Ceratophrys_, _36_; + bony plates of, 168; + teeth, 170 + + _Ceratops_, _39_; + see _Polyonax_ + + Ceratopsia, _39_; + characters, 209; + premaxillae, 284 + + Ceratopsidae, _39_ + + _Ceratosaurus_, _38_, 208; + supratemporal fossae, 283; + _C. nasicornis_, skeleton, =206= + + Cercopithecidae, _49_, _373_ + + Cervical ribs, crocodile, 260; + reptiles, 285 + + Cervical vertebrae, crocodile, 239; + dog, 380; + duck, 307; + general characters, 15; + mammals, 442; + turtle, 219 + + Cervidae, _46_; + skull, 469 + + _Cervulus_, _46_; + pes, 523 + + _Cervus_, 46; + _C. megaceros_, antlers, 469 + + _Cestracion 32_; + calcification of vertebrae 114; + external branchial arches 121; + pectoral fin 130; + skull =118=; + suspensorium 119; + teeth 109; + vertebral column 114 + + Cestraciontidae _32_ + + Cetacea _44_, 522; + arm bones 501; + auditory ossicles 487; + caudal vertebrae 453; + cervical vertebrae 444; + characters 353; + exoskeleton 416 f.; + hind limb 518; + manus 505; + pectoral girdle 495; + pelvis 514; + position of limbs 28; + ribs 491; + skull 461 f.; + sternum 489; + teeth 426; + thoraco-lumbar vertebrae 448 + + Cetiosauridae _38_ + + _Chalcides_ _38_; + limbs 289 + + Chalicotheriidae _46_; + femur 519; + manus 509 + + _Chalicotherium_ _46_; + femur 360; + pes =508=, 525; + teeth 432 + + _Chamaeleon_ _38_, 199 f.; + epipubis 293; + ilia 291; + manus 291; + skull 278 + + Chamaeleonidae _38_ + + Charadriidae _42_ + + Charadriiformes _42_ + + _Chauna_ _41_; + interorbital septum 333; + ribs 336; + _C. derbiana_, spurs 330 + + _Chelone_ _37_, _194_; + plastron 271, =218=; + see Turtle + + Chelonia _37_; + beaks 271; + carapace 271; + general characters 193; + humerus 290; + limbs 290; + palate 281; + pectoral girdle 288; + pelvic girdle 291; + skull 277 f.; + tarsus 293; + vertebrae 275 f. + + Chelonidae _37_ + + Chelydae _37_ + + _Chelydra_ _37_; + carpus 26, 291 + + Chelydridae _37_ + + _Chelys_ _37_, _195_ + + Chersidae _37_ + + Chevron bones 16; + crocodile 243; + mammals 453 f.; + reptiles 276 + + Chevrotain _45_, _359_; + teeth 429 + + _Chimaera_ _32,_ _66_; + attachment of fins 130; + pelvic girdle 127; + skull =65=; + teeth 110 + + Chimaeridae _32_ + + Chimaeroidei, general characters 65 + + Chimpanzee _49_; + carpus 512; + ribs 493; + thoraco-lumbar vertebrae 450 + + _Chinchilla_ _47_; + auditory ossicles 488 + + Chinchillidae _47_ + + Chiromyidae _49_ + + _Chiromys_ _49,_ _372_; + manus 512; + pes 527; + teeth 441 + + Chiroptera _49_; + auditory ossicles 488; + arm bones 503; + cervical vertebrae 446; + general characters 370; + manus 512; + pelvis 515; + pes 527; + sacrum 452; + shoulder girdle 499; + skull 481; + sternum 490; + tail 454; + teeth 440; + thigh and shin 520; + thoraco-lumbar vertebrae 450 + + _Chirotes_ _38_; + limbs 289 + + _Chlamydophorus_ _44,_ _272_; + scutes 419; + skull 459 + + _Chlamydoselache_ _31_; + branchial arches 121 + + _Choeropus_ _43_; + manus 504; + pes 522 + + _Choloepus_ _43_; + ribs 491; + shifting of pelvis 451; + skull =458=; + sternum 489; + thoraco-lumbar vertebrae 447; + _C. hoffmanni_ cervical vertebrae 443 + + Chondrocranium, salmon 87 + + Chondroid tissue, _Balanoglossus_ 50 + + Chondrostei _32_; + fins 105; + general characters 67; + teeth 110; + see Cartilaginous ganoids + + Chordal sheath, _Amphioxus_ 52 + + Chrysochloridae _49_ + + _Chrysochloris_ _49_; + auditory ossicles 488; + claws 418; + teeth 440 + + _Ciconia_ _41_; + see Stork + + Ciconiiformes _41_ + + Cingulum 376 + + Civet _48,_ _369_; + teeth 437 + + _Cladoselache_ _31,_ _63_; fin 129 + + Clasper 132; + dogfish 82 + + Clavicle 25; + birds 338; + cod 102; + duck 322; + dog 405; + fish 126; + frog 163; + mammals 494 f.; + reptiles 289 + + Claws 3; + birds 330; + crocodile 237; + dog 374; + duck 302; + mammals 417; + turtle 215 + + Clupeidae, _33_ + + _Clupeus_, _33_ + + Cnemial crest, dog, 412; + duck, 326 + + _Coccosteus_, _34_, _70_ + + Coccyx, man, 454 + + Cochliodontidae, _31_ + + _Cochliodus_, _31_; + dental plates, 109 + + Cod, _33_; + appendicular skeleton, 101 f.; + cranium, 96; + mandibular and hyoid arches, =99=; + median fins, 86; + pectoral girdle and fin, =102=; + ribs, 86; + skull, 96 f.; + vertebral column, 83 f. + + _Coelogenys_, _48_; + zygomatic arch, 477 + + _Coenolestes_, _43_, 424 + + Coffer-fish, _33_; + see, _Ostracion_ + + _Colobus_, _49_; + pollex, 512 + + Colubridae, _38_ + + Columbae, _42_ + + Columbidae, _42_ + + Columella, crocodile, 251; + duck, 320; + frog, 157; + turtle, 228 + + Columella cranii, 200 n; + see epipterygoid + + Colymbi, _40_ + + Colymbiformes, _40_ + + Compsognathidae, _38_ + + _Compsognathus_, _38_, 208 + + Condylar ridge, duck, 326 + + Condyle of humerus, dog, 406; + of mandible, dog, 398 + + Condylarthra, _47_; + femur, 519; + general characters, 361; + manus, 509; + skull, 472; + teeth, 432 + + Contour feather, duck, 303 + + Copula, 23 + + _Coracias_, _42_; + see Roller + + Coraciae, _42_ + + Coraciiformes, _42_ + + Coracoid, 25; + cod, 103; + crocodile, 263; + duck, 322; + frog, 163; + Monotremata, 493; + newt, 147; + reptiles, 288; + turtle, 232 + + Coracoid groove, duck, 321 + + Cormorant, _41_; + foot, 342; + skull, 335 + + Cornu, see hyoid + + Cornua trabeculae, 18 + + _Coryphodon_, _47_; + femur, 519; + manus, 510; + pes, 525; + skull, 473; + teeth, 433; + _C. hamatus_, manus, =510= + + Coryphodontidae, _47_ + + Costal plate, turtle, 215; + -- process, duck, 321; + -- shield, turtle, 214 + + Cotyloid bone, 25, 513; + see Acetabular bone + + Cotylopidae, _45_ + + _Cotylops_, _45_; + pollex, 506; + skull, 468 + + Coverts, 306, 328 + + Cranium, 18; + cod, 96 f.; + crocodile, 244 f.; + development of, 16 f.; + dog, 384 f.; + dogfish, 73 f.; + duck, 314; + frog, 154 f.; + newt, 140 f.; + turtle, 222 f. + + Cranio-facial axis, dog, 384 + + Creodonta, _48_; + carpus, 512; + femur, 520; + general characters, 368; + skull, 479; + teeth, 439 + + Cribriform plate, dog, 388, 400 + + Crocodile, _210_, _212_; + anterior limb, 263; + exoskeleton, 237; + pectoral girdle, 262; + pelvic girdle, 266; + posterior limb, 268; + ribs and sternum, 259; + skeleton, 237 f.; + skull, 243 f.; + tarsus, 293; + teeth, 238; + vacuities in surface of cranium, 256; + vertebral column, 239 + + Crocodilia, _39_; + general characters of, 210; + palate, 281; + skull, 277 f.; + succession of teeth, 274; + teeth, 273 + + Crocodilidae, _39_ + + _Crocodilus_, _39_; + _C. palustris_, sternum and associated bones, =261=; + late thoracic and first sacral vertebrae, =242=; + _C. vulgaris_, cervical vertebrae, =239= + + Crossopterygii, _33_; + general characters, 68 + + Crotalidae, _38_ + + _Crotalus_, _38_; + jaws 280; + see Rattlesnake + + Crows, _42_ + + Crura of stapes, dog, 393 + + Cruro-tarsal, ankle joint, 345 + + Crus, 26; + crocodile, 268; + dog, 412; + duck, 326; + frog, 166; + newt, 149; + turtle, 235 + + Crusta petrosa, 5 + + _Cryptobranchus_, _35_, 135; + skull, 175; + _C. lateralis_, sacral vertebrae, 171 + + Cryptodira, _37_; + characters, 194 + + Ctenoid scales, 8, 60, 105 + + Cubitals, 303 f. + + Cuboid, 27; + dog, 415 + + Cuckoo, foot, 342 + + Cuculi, _42_ + + Cuculiformes, _42_ + + Cuneiform bones, 27; + dog, 414 f. + + _Cyclodus_, _38_; + see _Tiliqua_ + + Cycloid scales, 8, 60, 105; + cod, 83 + + _Cyclopidius_, _45_; + skull, 468 + + Cyclospondyli, 114 + + Cyclostomata, _31_; + general characters, 53 + + _Cycloturus_, _44_; + hallux, 522; + manus, 505 + + _Cygnus_, _41_; + see Swan + + _Cynocephalus_, _49_; + cervical vertebrae, 446; + skull, 482 + + Cynoidea, _48_, _369_; + dental formula, 437 + + _Cynognathus_, _36_; + occipital condyle, 277; + teeth, 273 + + Cyprinidae, _33_ + + _Cyprinus_, _33_; + see Carp + + Cypseli, _42_ + + Cypselidae, _42_; + see Swifts + + Cystignathidae, _36_ + + + _Dactylopterus_, _34_; + pectoral fins, 131 + + Dasypodidae, _44_ + + _Dasyprocta_, _48_; + auditory ossicles, 488; + thoraco-lumbar vertebrae, 450 + + Dasyproctidae, _48_ + + _Dasypus_, _44_; + manus, 505; + skull, 459; + stapes, 487; + teeth, 424 + + Dasyuridae, _43_, _350_; + dentition, 423; + pes, 521; + skull, 456 + + Deer, _359_; + manus, 507; + pes, 523; + Chinese water --, _46_, + see _Hydropotes_; + Musk --, _46_, + see _Moschus_; + Red -- fibula, 519 + + Delphinidae, _45_ + + _Delphinus_, _45_, 357; + lumbar vertebrae, 448; + skull, 462 f. + + Deltoid ridge, crocodile, 263; + dog, 406; + frog, 164 + + _Dendrohyrax_, 363 + + Dental formula, regular, 344, 422; + Anthropoidea, 441; + _Astrapotherium_, 432; + _Babirussa_, 428; + Camel, 428; + _Chiromys_, 441; + Chiroptera (many), 440; + Cynoidea, 437; + _Dinotherium_, 434; + Dog, 376; + Duplicidentata, 435; + _Elephas_, 434; + _Erinaceus_, 440; + _Felis_, 437; + _Galeopithecus_, 440; + _Hippopotamus_, 427; + Horse, 430; + _Hydromys_, 436; + Hyracoidea, 362; + Macropodidae, 423; + _Manatus_, 425; + _Notoryctes_, 423; + _Otaria_, 439; + _Procavia_, 432; + _Pteropus_, 441; + Rodentia (most), 435; + Ruminantia, 429; + _Squalodon_, 427; + _Sus_, 428; + Tapiridae, 429; + _Thylacinus_, 423; + _Uintatherium_, 433; + _Ursus_, 439; + _Zeuglodon_, 426 + + Dentary, 22; + crocodile, 258; + cod, 100; + duck, 320; + frog, 161; + newt, 144; + salmon, 94; + turtle, 230 + + Dentine, 5 + + Derbian Screamer, spurs, 330 + + Dermal exoskeleton, crocodile, 237; + fish, 105; + mammals, 419; + reptiles, 271; + turtle, 215 + + Dermo-supra-occipital, Labyrinthodontia, 177; + _Polypterus_, 122 + + Dermochelydidae, _37_ + + _Dermochelys_, _37_, _194_, 214, 270; + carapace and plastron, 272 + + Dermoptera, _48_; + general characters, 370 + + Derotremata, _35_ + + _Desmodus_, _49_; + teeth, 441 + + Desmognathous, 319, 335 + + Development of bone, 10; + of cranium, 16; + of teeth, 7 + + _Dicynodon_, _36_, _192_; + beak, 271; + supratemporal fossa, 283; + teeth, 273 + + Didelphia, _43_; + general characters, 349 + + Didelphyidae, _43_, _350_; + auditory ossicles, 486; + pes, 521; + teeth, 423 + + _Didelphys_, _43_; + atlas, 443; + teeth, 422 + + _Didus_, _42_; + see Dodo + + Digitigrade, defined, 358 n. + + Digits, _26_; + see Manus and Pes + + _Dimetrodon_, _36_; + thoracic vertebrae, 276 + + _Dinichthys_, _34_, _70_ + + Dinocerata, 364; + see Uintatheriidae + + Dinornithes, _40_, _299_; + see Moas + + Dinosauria, _38_; + general characters, 204; + humerus, 290; + ischium, 291; + pectoral girdle, 288; + pes, 293; + pre-orbital vacuity, 284; + ribs, 285; + vertebrae, 275 f. + + Dinotheriidae, _47_ + + _Dinotherium_, _47_, 365; + dental formula, 434; + teeth, 345 + + _Diodon_, _33_; + beaks, 111; + _D. hystrix_, scales, 105 + + Diphycercal tail, 60, 116 + + Diphyodont, defined, 7, 344 + + _Diplacanthus_, _32_, _64_ + + Dipneumona, _34_ + + Dipnoi, _34_; + general characters, 69; + pelvic fins, 131; + skull, 124; + spinal column, 113; + tail, 116; + teeth, 111 + + Dipodidae, _47_ + + Diprotodont, 423 + + Diprotodontia, _43_; + characters, 350 + + Dipteridae, _34_, _70_; + cranium, 124; + tail, 117; + teeth, 111 + + _Dipus_, _47_; + cervical vertebrae, 446; + pes, 526 + + Discoglossidae, _36_ + + _Discoglossus_, _36_; + ribs, 182; + vertebrae, 172 + + Distal, defined, 23 n. + + Divers, _40_; + thoracic vertebrae, 332 + + _Docidophryne_, _36_; + shoulder girdle and sternum, =183= + + Dodo, _42_; + wing, 338 + + Dog, _48_; + arm bones, =407=; + anterior limb, 405; + atlas and axis, =379=; + cranium, 384, =389=, =396=; + dentition, =375=; + innominate bone, =410=; + leg bones, =411=; + manus, 408, =413=, 511; + pectoral girdle, 404; + pelvic girdle, 409; + pes, =413=, 414; + posterior limb, 412; + ribs, 402; + second lumbar vertebra, =382=; + second thoracic vertebra, =382=; + skull, 383, =387=; + sternum, =403=, 404; + vertebral column, 378 + + Dogfish, _64_; + cranium, 73; + exoskeleton, 71; + median fins, 79; + pectoral girdle and fin, 79; + pelvic girdle, 81; + pelvic fin, 81; + ribs, 73; + skull, 73, =75=; + vertebral column, 72; + visceral skeleton, 77; + Spotted and Spiny --, _32_ + + Dolphin, _45_, _357_; + lumbar vertebrae, 448; + Gangetic --, _45_, + see _Platanista_ + + Donkey, skull, =431= + + _Dorcatherium_, _45_; + manus, 507 + + Dorsal vertebra, 16 + + Dorsal shield, crocodile, 238 + + Down feathers, 306 + + _Draco_, _38_; + ribs, 286 + + Dromaeognathous, 335 + + _Dromaeus_, _40_, _299_; + see Emeu + + Duck, _41_, 334; + beak, 329; + claws, 330; + cranium, =313=; + exoskeleton, 302; + pectoral girdle, 321; + pelvic girdle, 324, =311=, =325=; + pes, 327; + posterior limb, 326; + ribs, 320; + skull, 312, =312=, =313=; + sternum, 321; + vertebral column, 307; + wing, 322, =304=, =305= + + Duckbill, _43_; + see _Ornithorhynchus_ + + Dugong, _44_; + humerus, 501; + pelvis, 514; + thoraco-lumbar vertebrae, 448; + see _Halicore_ + + Duplicidentata, _48_, 366; + dental formula, 435; + skull, 478 + + + Eagles, 335 + + Eared Seals, _369_; + scapula, 498; + see Otariidae + + _Echidna_, _43_; + caudal vertebrae, 453; + manus, 504; + pelvis, 513; + sacral vertebrae, 451; + shoulder-girdle and sternum, =494=; + skull, 455; + spines, 417; + spur, 418; + thoraco-lumbar vertebrae, 447 + + Echidnidae, _43_; + pes, 521 + + Ectethmoid, 21 n. + + Ectocondylar ridge, dog, 406 + + Edentata, _43_; + auditory ossicles, 487; + arm bones, 500; + caudal vertebrae, 453; + cervical vertebrae, 443; + manus, 504; + pectoral girdle, 495; + pes, 522; + pelvis, 513; + ribs, 491; + sacrum, 452; + skull, 457; + sternum, 489; + teeth, 424; + thigh and shin, 517; + thoraco-lumbar vertebrae, 447 + + Eel, _33_; + scales, 105 + + Elasmobranchii, _31_; + cranium, 118 f.; + clasper, 132; + general characters, 61; + pelvic fins, 131; + ribs, 125; + teeth, 109; + vertebral column, 113 f.; + visceral arches, 120 + + _Elasmotherium_, _46_; + mesethmoid, 470 + + Elephant, _47_; + auditory ossicles, 487; + caudal vertebrae, 453; + ribs, 491; + skull, 473 f., =474= and =475=; + tusks, 420; + see also Proboscidea + + Elephantidae, _47_ + + _Elephas_, _47_, _364_; + dental formula, 434; + _E. planifrons_, 435; + see Elephant + + _Elginia_, _36_; + skull, 191, 283 + + Embolomerous, 172 + + Emeu, _40_, _299_; + aftershaft, 328; + claws, 330 + + Enamel, 4; + -- cap, 7; + -- organ, 7 + + Endochondral ossification, 11 + + Endoskeleton, Amphibia, 170; + birds, 331 f.; + cod, 83 f.; + crocodile 239, f.; + dog, 377 f.; + dogfish, 71 f.; + duck, 306 f.; + fish, 112 f.; + frog, 151 f.; + mammals, 442 f.; + newt, 138 f.; + reptiles, 275 f.; + turtle, 218 f. + + Engystomatidae, _36_ + + Entoplastron, turtle, 217 + + Epanorthidae, _43_, _350_ + + Epi-branchial, cod, 101; + dogfish, 78; + salmon, 94 + + Epicoracoid, 25; + frog, 163; + turtle, 232; + Monotremes, 493; + vestiges of in Rodentia, 497 + + _Epicrium_, _35_; + orbit, 179 + + Epidermal exoskeleton, birds, 328; + crocodile, 237; + dog, 374; + duck, 302; + mammals, 416; + reptiles, 270; + turtle, 214 + + Epi-hyal, cod, 100; + dog, 399; + salmon, 94 + + Epi-otic, 20; + cod, 96; + crocodile, 250; + Labyrinthodontia, 177; + reptiles, 278; + salmon, 89; + turtle, 227 + + Epiphysis, 11 + + Epiplastron, turtle, 217 + + Epiprecoracoid, Amphibia, 184; + turtle, 232 + + Epipterygoid, Lacertilia, 200; + reptiles, 278 + + Epipubis, crocodile, 267; + newt, 149; + turtle, 235 + + Episternum, 217; + frog, 163 + + Equidae, _46_; + mane, 416; + scapula, 496; + skull, 471 + + _Equus_, _46_; + see Horse + + Erinaceidae, _49_ + + _Erinaceus_, _49_; + dental formula, 440; + pelvic symphysis, 515; + presternum, 490; + see Hedgehog + + Esocidae, _33_ + + Esox, _33_; + attachment of teeth, 107 + + Ethmoid, 394; + see median ethmoid + + Ethmoidal plane, 390; + -- region, 21 + + Ethmo-palatine ligament, dogfish, 77 + + Ethmo-turbinal, dog, 395 + + _Euchirosaurus_, _35_; + vertebrae, 171 + + Eustachian canal, see Canal + + Eusuchia, _39_; + general characters, 212 + + Eutheria, _43_; + general characters, 351 + + _Exocaetus_, _33_; + pectoral fins, 131 + + Exoccipital, 19; + cod, 97; + crocodile, 246; + dog, 386; + duck, 314; + frog, 154; + newt, 141; + salmon, 89; + turtle, 224 + + Exoskeleton, 2; + Amphibia, 168; + birds, 328; + crocodile, 237; + dog, 374; + dogfish, 71; + duck, 302; + fish, 104; + ganoids, 66; + mammals, 442; + reptiles, 270; + turtle, 214 + + Extensor side, defined, 29 + + Extra-branchial, dogfish, 79 + + Extra-columella, crocodile, 251; + turtle, 228 + + + Fabella, dog, 412 + + _Falco_, _41_ + + Falcon _41_, 335 + + Falconiformes, _41_ + + Feathers, 3, 328; + duck, 302 + + Felidae, _48_; + claws, 418 + + _Felis_, _48_; + dental formula, 437; + thoraco-lumbar vertebrae, 450 + + Femoral shield, turtle, 215 + + Femur, 26; + crocodile, 268; + dog, 412; + duck, 326; + frog, 166; + mammals, 517 f.; + newt, 149; + ox and rhinoceros, =518=; + turtle, 235 + + Fenestra ovalis, crocodile, 250 f.; + dog, 392; + duck, 316; + frog, 157; + turtle, 228; + -- rotunda, dog, 392; + duck, 316 + + Fenestral recess, duck, 316 + + Fibula, 26; + crocodile, 268; + dog, 412; + duck, 327; + frog, 166; + newt, 149; + turtle, 235 + + Fibulare, 27; + see Tarsus + + File-fish, _33_; + see _Balistes_ + + Filoplume, 306 + + Finches, _42_ + + Fins, fish, 115; + caudal --, Cetacea, 453; + cod, 87; + fish, 116; + Ichthyosauria, 195; + median --, cod, 86; + dogfish, 79; + pectoral --, cod, 103; + dogfish, 79; + pelvic --, cod, 103; + dogfish, 81 + + Fin-rays, 105, 115; + cod, 83, 103; + dogfish, 79 + + Firmisternia, _36_, 185 + + Fish, appendicular skeleton, 126; + endoskeleton, 112 f.; + exoskeleton, 104; + general characters, 60 f.; + paired fins, 127 f.; + ribs, 125 f.; + skull, 117 f.; + spinal column, 112 f.; + teeth, 106 f. + + Fissipedia, _48_; + general characters, 368 + + Flamingo, 335 + + Flexor side, defined, 29 + + Floating ribs, dog, 402; + mammals, 490 + + Flower, Sir W.H., on succession of teeth in elephants, 434 + + Flying-fish, _33_; + -- fox, _49_, _371_, + skull, 481, + see _Pteropus_; + -- gurnard, _34_, + see _Dactylopterus_; + -- lemur, _48_, + see _Galeopithecus_; + -- lizard, _38_, + see _Draco_ + + Fontanelle, salmon, 89; + anterior --, dogfish, 74; + frog, 154; + posterior --, frog, 154 + + Foot, crocodile, 269; + dog, 414; + frog, 167; + newt, 149 f.; + turtle, 236 + + Foramen: + anterior palatine --, dog, 401; + condylar --, dog, 401; + -- cordiforme, reptiles, 292; + ect-epicondylar --, _Sphenodon_, 290; + ent-epicondylar --, 191 n.; + Carnivora vera, 502; + Cebidae, 503; + _Choloepus_, 500; + Condylarthra, 362, 502; + Creodonta, 368; + Insectivora, 503; + Lemurs, 503; + Marsupials, 500; + reptiles, 290; + external mandibular --, crocodile, 258; + inferior dental --, dog, 399; + infra-orbital --, dog, 401; + Rodents, 477; + ilio-sciatic --, duck, 325; + internal mandibular --, crocodile, 258; + internal orbital --, dog, 401; + interparietal --, Labyrinthodontia, 173, 177; + reptiles, 277; + -- lacerum anterius, dog, 388, 400; + -- lacerum medium, dog, 402; + -- lacerum posterius, dog, 392, 401; + lachrymal --, dog, 394, 401; + -- magnum, cod, 97; + crocodile, 257; + dog, 386, 402; + dogfish, 76; + duck, 314; + frog, 154; + newt, 141; + salmon, 89; + turtle, 224; + mental --, dog, 399; + obturator --, duck, 326; + dog, 410; + ophthalmic --, dogfish, 74; + optic --, dog, 400; + dogfish, 74; + orbitonasal --, dogfish, 74; + -- ovale, crocodile, 249; + dog, 400; + pneumatic --, duck, 323; + pneumogastric --, dogfish, 76; + posterior palatine --, dog, 401; + postglenoid --, dog, 402; + pre-acetabular --, Chiroptera, 515; + -- rotundum, dog, 400; + stylomastoid --, dog, 392 f., 400; + thyroid --, dog, 410; + trigeminal --, duck, 316; + -- triosseum, duck, 322 + + Fore-arm, 26; + crocodile, 265; + dog, 406; + duck, 323; + frog, 164; + newt, 147; + turtle, 233 + + Fossa: + cerebellar --, dog, 392; + cerebral --, dog, 392; + digital --, dog, 412; + floccular --, dog, 392; + infratemporal --, see lateral temporal --; + lachrymal --, Ruminants, 469; + lateral temporal --, crocodile, 257; + _Sphenodon_, 283; + olecranon --, dog, 406; + prescapular --, dog, 405; + postscapular --, dog, 405; + post-temporal --, _Sphenodon_, 283; + pterygoid --, crocodile, 257; + suborbital --, Ruminants, 469; + supra-acetabular --, Ruminants, 514; + supratemporal --, crocodile, 249, 256; + reptiles, 283; + supra-trochlear --, dog, 406; + temporal --, dog, 398; + trochanteric --, dog, 412 + + Fowl, _41_, 335; + claws, 330; + skeleton, =301= + + Fox, _48_ + + Frigate bird, _41_; + clavicles, 338 + + Frog, anterior limb, 164; + cranium, =155=, =157=; + hyoid apparatus, 161; + pelvic girdle, 165; + posterior limb, 166; + shoulder-girdle and sternum, =183=; + skull, 154 f., =155=, =159=; + teeth, 151; + vertebral column, 152; + Common --, Edible --, Fire-bellied --, Green-tree --, + Horned --, Midwife --, Painted -- and Toad --, _36_; + + Frontal, 19; + cod, 96; + crocodile, 249; + dog, 388; + duck, 314; + newt, 141; + salmon, 91; + turtle, 225; + -- segment, crocodile, 249; + dog, 388; + turtle, 225 + + Fronto-parietal, frog, 156 + + Frugivorous bats, manus 512; + see Pteropidae + + Fulcra 67; + _Polypterus_ 106 + + Furcula 296; duck 322 + + + Gadidae _33_ + + _Gadus_ _33_; + see Cod + + Galeopithecidae _48_ + + _Galeopithecus_ _48_, _370_; + dental formula 440; + inter centra 450; + pelvic symphysis 515; + skull 480 + + _Galesaurus_ _36_, _192_; teeth 273 + + _Galeus_, _32_; + occipital joint 118 + + Galli _41_ + + Galliformes _41_ + + _Gallus_ _41_; + _G. bankiva_ skeleton =301= + + Gannet, _41_; wing =339= + + Ganoid scales 8, 60, 104 + + Ganoidei _32_; + general characters and distribution 66; + pectoral girdle 126; + pelvic fins 132; + teeth 110; + skull 121 f.; + spinal column 112 and 114 + + Garialidae _39_ + + _Garialis_ _39_, _212_ + + Garial 210 + + Gar pike 33; + see _Lepidosteus_ + + _Gavialis_ _39_ + + _Gazella_ _46_ + + Gazelle _46_; + skull 468 + + Geckonidae _37_; + see Gecko + + Gecko _37_; + epipubis 293; + parietals 277; + supratemporal fossa 283; + vertebrae 275 + + Gibbon _49_; + ribs 493; + skull 482 + + Gill-rays, dogfish 78; + salmon 95 + + _Giraffa_ _46_ + + Giraffe _46_, _359_; + cervical vertebrae 445; + manus 507; + pes 523; + ulna 501 + + Giraffidae _46_; skull 469 + + Girdle bone, frog 156 + + Glenoid cavity 25; + crocodile 263; + dog 405; + duck 322; + frog 162; + newt 146; + turtle 232; + --fossa, dog 394 + + Globe-fish _33_ + + _Globicephalus_ _45_; + cervical vertebrae 354; + manus 506; + skull 463 + + Gluteal surface of ilium, dog 410 + + _Glyptodon_ _44_; + carapace 419; + cervical vertebrae 443; + caudal vertebrae 453; + manus 505; + pelvis 513; + pes 522; + teeth 425; + thoraco-lumbar vertebrae 447 + + Glyptodontidae _44_, _352_; + skull 459; + see also _Glyptodon_ + + Gnathostomata _31_, _59_ + + Golden mole _49_; + see Chrysochloris + + Goniopholidae _39_ + + _Goniopholis_ _39_; + vertebrae 275 + + Goose 334; + beak 329; + Spur-winged--41 + + _Gorilla_ _49_; + carpus 512; + ribs 493; + scapula 499; + skull =483=; + thoraco-lumbar vertebrae 450 + + Gruidae _41_; + see Cranes + + Gruiformes _41_ + + Guinea-pig _48_; + tail 454 + + Gular shield, turtle 215 + + Gulls _42_, 335; + aftershaft 328 + + Gymnodontidae _33_; + beaks 111 + + Gymnophiona _35_; + branchial arches 180; + general characters 136; + ribs 182; + scales 168; + skull 177; + teeth 169; + vertebrae 172 + + _Gymnura_ _49_; + teeth 440; + zygomatic arch 481 + + _Gypogeranus_ _41_; + claws 330 + + _Gyrinophilus_ _35_; + vertebral column 171 + + + Haddock _33_ + + Hadrosauridae _39_ + + _Hadrosaurus_ _39_; + skull 284 + + Hag or hag-fish _31_, 54 f. + + Hair 3; + dog 374; + mammals 416 + + _Halicore_ _44_, _352_; + manus 505; + skull 460; + teeth 425; + see Dugong + + Halicoridae _44_ + + Halitheriidae _44_ + + _Halitherium_ _44_, 352; + femur 518; + pelvis 514; + teeth 425 + + Hallux 26; + dog 415; + duck 327; + frog 167 + + Hamular process, dog 397 + + Hand, crocodile 266; + dog 408 f.; + duck 324; + frog 165; + newt 147; + turtle 233 + + _Hapale_ _49_ + + Hapalidae _49_, 372 f.; + teeth 441 + + Hare _48_, 366; + acetabular bone 515; + dental formula 435; + femur 520; + humerus 502; + pelvis 515; + scapula 497; + skull 476; + tail 454; + thoraco-lumbar vertebrae 449; + Cape jumping --, _47_; + see _Pedetes_ + + _Harpagus_, _41_; + serrated beak, 334 + + _Harriotta_, _32_, _66_ + + _Hatteria_, _37_, _197_; + see _Sphenodon_ + + Haversian canals, 10; + -- system, 10 + + Hawks, beak of, 330 + + Hedgehog, _49_, _370_; + auditory ossicles, 488; + humerus, 503; + presternum, 490; + skull, 480; + spines, 417; + see _Erinaceus_ + + Hemichordata, _30_, _50_ + + _Heptanchus_, _31_; + branchial arches, 63, =120=; + vertebrae, 114 + + Herbivorous dentition, 427, 430 + + Heron, _41_, 335; + interorbital septum, 333; + powder down feathers, 329 + + Herring, _33_ + + _Hesperornis_, _40_, _299_; + caudal vertebrae, 333; + clavicles, 338; + teeth, 330; + wing, 338 + + Heterocercal tail, 60, 116 + + Heterodont, 7 + + Heterostraci, _31_; + general character, 54 + + _Hexanchus_, _31_; + branchial arches, 63, 121 + + Hinge joint, 13 + + _Hipparion_, _46_; + manus, 508; + pes, =524= + + Hippopotamidae, _45_ + + _Hippopotamus_, _45_, _359_; + dental formula, 427; + hair, 416; + mandible, 467; + manus, 506; + pes, 523; + scapula, 496; + skull, 467; + teeth, 345 + + Hoatzin, _41_; + see _Opisthocomus_ + + Holocephali, _32_, 65, 104; + clasper, 132; + spinal column, 113; + tail, 116; + teeth, 109 + + Holoptychiidae, _33_ + + _Holoptychius_, _33_; + scales, 105 + + Holostei, _33_; + general characters, 68; + teeth, 110; + see Bony Ganoids + + Hominidae, _49_, _373_ + + _Homo_, _49_; + see Man + + Homocercal tail, 60, 69, 117; + codfish, 87 + + Homodont, defined, 7 + + Hoofs, 3, 418 + + Hoopoe, _42_, 335 + + _Hoplopterus_, spur, 330 + + Hornbill, _42_, 331; + bony crest, 334; + interorbital septum, 333 + + Horns, 3, 417 + + Horny plates on palate, 418; + -- teeth, Lampreys, 4; + Myxinoids, 57; + _Ornithorhynchus_, 4 + + Horse, _46_, _360_; + fibula, 519; + malleus, 487; + manus, 507; + pes, =524=; + skull, 471; + teeth, 345, 430; + ulna, 501 + + Howling monkey, _49_; + see _Mycetes_ + + Humerals, duck, 303 f. + + Humeral shield, turtle, 215 + + Humerus, 26; + crocodile, 263; + dog, 405; + duck, 323; + frog, 164; + newt, 147; + turtle, 232; + wombat, =500= + + Humming-birds, _42_, 335 + + Humpbacked whale, _44_, _357_ + + _Hyaena_, _48_, _369_; + hallux, 526; + pollex, 511; + sacral vertebrae, 452; + teeth, 437 + + Hyaenidae, _48_; + humerus, 502 + + _Hyaenodon_, _48_, _368_ + + Hyaenodontidae, _48_ + + Hyaline cartilage, 10 + + _Hydrochaerus_, _48_; + teeth, 437; + see Capybara + + _Hydromys_, _47_; + dental formula, 436 + + Hydrophidae, _38_; + scales, 270 + + _Hydropotes_, _46_; + canines, 429 + + _Hyla_, _36_; + fronto-parietal fontanelle, 179; + sternum, 184 + + Hylidae, _36_ + + _Hylobates_, _49_; + ribs, 493; + skull, 482 + + Hyoid, 21; + alligator, =285=; + Amphibia, 180; + birds, 336; + cod, 100; + crocodile, 259; + dogfish, 77; + dog, 399; + duck, 320; + frog, 161; + newt, 144; + reptiles, 284; + salmon, 94; + turtle, 231, =285= + + Hyomandibular, 23; + cod, 100; + dogfish, 78; + salmon, 94 + + _Hyomoschus_, _45_ + + Hyoplastron, turtle, 217 + + Hyostylic, 61, 119 + + _Hyotherium_, _45_; + teeth, 427 + + _Hyperodapedon_, _37_, _198_; + premaxillae, 284 + + _Hyperoödon_, _44_; + skull, 464; + sternum, 489; + thoracic vertebrae, 448 + + Hypo-branchial, cod, 101; + dogfish, 78 + + Hypo-hyal, cod, 100; + salmon, 95 + + Hypo-ischium, Lacertilia, 292 + + Hypoplastron, turtle, 217 + + Hyporachis, 328 + + _Hypsilophodon_, _39_; + predentary bone, 284 + + Hypsodont, defined, 345, 429 + + Hypural bone, cod, 85 + + Hyracidae, _47_ + + Hyracoidea, _47_; + femur, 519; + general characters, 362; + manus, 510; + nails, 418; + skull, 472; + teeth, 432 + + _Hyracotherium_, _46_; + manus, 508; + scapula, 496 + + _Hyrax_, _47_, _363_; + see _Procavia_ + + Hystricidae, _47_ + + Hystricomorpha, _47_; + auditory ossicles, 488 + + _Hystrix_, _47_; + auditory ossicles, 488; + see Porcupine + + + Ichthyodorulites, 106 + + Ichthyoidea, _35_; + general characters, 134 + + Ichthyopsida, _31_; + general characters, 59 + + Ichthyopterygium, 130 + + _Ichthyornis_, _40_; + mandible, 335; + pelvis, 341; + teeth, 330; + vertebrae, 332 + + Ichthyornithiformes, _40_, 300 + + Ichthyosauria, _37_; + general characters, 195; + ribs, 285 + + Ichthyosauridae, _37_ + + _Ichthyosaurus_, _37_, _197_; + limbs, 290; + palatines, 281; + pectoral girdle, 288; + position of limbs, 28; + skull, =196=; + teeth, 273; + vertebral column, 275 + + Ichthyotomi, _31_; + general characters, 62; + fins, 130 f.; + tail, 116 + + _Iguana_, _38_; + teeth, 273 + + Iguanidae, _38_; + zygosphenes, 200, 276 + + _Iguanodon_, _39_, 208 f.; + jaws, 292; + predentary, 284; + sternum, 288; + teeth, 272 f.; + vertebrae, 275 + + Iguanodontidae, _39_ + + Iliac surface of ilium, dog, 410 + + Ilium, 25; + crocodile, 266; + dog, 409; + duck, 325; + frog, 165; + mammals, 513 f.; + newt, 149; + reptiles, 291; + turtle, 235 + + Incisors, dog, 376 f.; + mammals, 344 + + Incus, dog, 393; + man, dog and rabbit, =485= + + Infra-marginal shield, turtle, 215 + + Infra-pharyngeal bone, cod, 101 + + _Inia_, _45_; + cervical vertebrae, 444; + lumbar vertebrae, 448 + + Innominate bone, dog, 409; + mammals, 513 + + Insectivora, _48_; + arm bones, 503; + auditory ossicles, 488; + cervical vertebrae, 446; + general characters, 369 f.; + manus, 512; + pelvis, 515; + pes, 527; + sacrum, 452; + shoulder-girdle, 499; + skull, 480; + sternum, 490; + tail, 454; + teeth, 440; + thigh and shin, 520; + thoraco-lumbar vertebrae, 450 + + Insectivora vera, _49_; + general characters, 370 + + inter centra, 15; + _Galeopithecus_, 370; + Ichthyosauria, 195; + Labyrinthodontia, 172; + _Sphenodon_, 198; + _Talpa_, 450 + + Interclavicle, 25; + crocodile, 263; + Monotremata, 494; + reptiles, 289 + + Intercondylar notch, dog, 412 + + Intergular shield, turtle, 215 + + Interhyal, cod, 100 + + Intermedium, 27; + see Carpus and Tarsus + + Intermuscular bones, cod, 86 + + Internasal septum, dogfish, 76 + + Interorbital septum, birds, 333; + crocodile, 247; + duck, 317; + reptiles, 277 + + Interspinous bones, cod, 86 + + Intertarsal ankle joint, 190 + + Intervertebral discs, 15, 378 + + Ischial tuberosity, dog, 411 + + Ischium, 25; + crocodile, 266; + dog, 410; + duck, 325; + frog, 165; + newt, 149; + turtle, 235 + + _Ischyodus_, _32_, _66_ + + Ivory, 5 + + + Jacana, _42_; + see _Parra_ + + _Jacare_, _39_; + scutes, 271 + + Jaws, 21; + cod, 98 f.; + crocodile, 252 f.; + dog, 395 f.; + dogfish, 77; + duck, 317 f.; + frog, 158 f.; + newt, 143 f.; + salmon, 93 f.; + turtle, 229 f. + + Jerboa, _47_; + cervical vertebrae, 446; + pes, 526 + + Joints, kinds of, 13 + + Jugal, 22; + crocodile, 255; + dog, 398; + duck, 318; + turtle, 229 + + Jugulares, 132 + + + Kangaroo, _43_; + dental formula, 423; + lumbar vertebrae, 447; + pectineal process, 513; + pes, 522; + tail, 453; + teeth, 345 + + Kestrel, claws, 330 + + Killer, _45_; + see _Orca_ + + Kiwi, _40_; + see _Apteryx_ + + Knee-cap, see patella + + Koala, _43_; + lumbar vertebrae, 447; + pes, 522; + tail, 453 + + Kükenthal, W., on teeth of Cetacea, 426; + on teeth of Marsupials, 422 + + + Labial cartilage, dogfish, 77; + _Squatina_, 119 + + Labridae, _33_ + + _Labrus_, _33_; + see Wrasse + + Labyrinthodontia, _35_; + buckler, 168; + general characters, 135; + interparietal foramen, 173; + pelvis, 187; + ribs, 182; + skull, 176; + teeth, 169 + + Lacertilia, _37_; + general characters, 199; + pectoral girdle, 288; + skull, 277; + vertebrae, 275 + + Lachrymal, 20; + cod, 97; + crocodile, 251; + dog, 394; + duck, 317; + salmon, 93 + + Lacunae, 10 + + _Lagenorhynchus_, _45_; + skull, 462 + + _Lagostomus_, _47_; + maxillae, 477 + + Lambdoidal crest, duck, 315 + + Lamella of malleus, dog, 393 + + Lamnidae, _32_ + + Lamprey, _31_, 55 f. + + Lancelet, _30_; + see _Amphioxus_ + + Laridae, _42_; + see Gulls + + Larks, _42_ + + Larvacea, _30_; + notochord, 51 + + _Latax_, _48_; + pes, 526 + + Lateral ethmoid, 21; + cod, 97; + salmon, 89 f. + + Leathery turtle, _37_; + see _Dermochelys_ + + Lemuroidea, _49_; + caudal vertebrae, 454; + general characters, 372; + nails, 418; + ribs, 493; + sacrum, 452; + skull, 482; + thoraco-lumbar vertebrae, 450; + see Lemurs + + Lemurs, carpus, 512; + pes, 527; + teeth, 441; + see Lemuroidea + + Lenticular, 485; + dog, 393 + + _Lepidosiren_, _34_, _70_; + branchial arches, 125; + fins, 130 + + Lepidosteidae, _33_ + + _Lepidosteus_, _33_; + attachment of teeth, 108; + distribution, 66; + pectoral fins, 131; + scales, 67, 104; + skull, 123; + tail, 117; + vertebrae, 68 + + _Lepidotus_, _33_; + teeth, 110 + + Leporidae, _48_ + + Lepospondyli, _35_ + + _Lepus_, _48_; + see Hare + + _Leptoptilus_, _41_; + see Adjutant + + _Lialis_, _37_, 289 + + Limbs, general account, 26; + modifications in position of, 28; + reptiles, 289 + + Llama, _45_, 359; + cervical vertebrae, 445; + skeleton, =496=; + teeth, 428 + + Limicolae, _42_ + + Lingual apparatus, lampreys, 58; + myxinoids, 57 + + Lion, _48_ + + _Loemanctus longipes_, shoulder girdle and sternum, =287= + + Loggerhead turtle, carapace, =216= + + _Lophiodon_, _46_; + teeth, 345, 429 + + Lophiodontidae, _46_ + + Lophiomyidae, _47_ + + _Lophiomys_, _47_; + pes, 526; + skull, 476 + + _Lophius_, attachment of teeth, 107 + + Lower jaw, see Mandible + + Lumbar vertebrae, 16; + crocodile, 242; + dog, 378 f.; + duck, 311 + + Lunar, 27; + dog, 408 + + + _Macacus_, _49_; + cervical vertebrae, 446 + + _Machaerodus_, _48_; + upper canines, 437 + + _Macrauchenia_, _46_, 358; + calcaneum, 360; + cervical vertebrae, 445; + fibula, 519; + tarsus, 523; + ulna, 501 + + Macraucheniidae, _46_, 509 + + Macropodidae, _43_, _350_; + dental formula, 423; + pes, 522 + + _Macropus_, _43_; + see Kangaroo + + Macroscelidae, _49_ + + _Macroscelides_, _49_; + skull, 480 + + Magnum, 27; + see Carpus + + Malar, 22; + see jugal + + Malleus, dog, 393; + man, dog and rabbit, =485= + + Mammalia, _42_; + auditory ossicles, 485; + cervical vertebrae, 442; + exoskeleton, 416; + general characters, 343; + manus, 503; + Mesozoic --, 348; + pectoral girdle, 493; + pelvic girdle, 512; + pes, 521; + ribs, 490; + sacral and caudal vertebrae, 451; + skull, 455; + sternum, 489; + thigh and shin, 517; + thoraco-lumbar vertebrae, 447 + + Man, _49_; + arm bones, 503; + auditory ossicles, 488; + caudal vertebrae, 454; + cervical vertebrae 446; + pelvis, 515; + pes, 527; + ribs, 493; + scapula, 499; + skull, 483; + sternum, 490; + teeth, 441 + + Manatee, _44_; + see _Manatus_ + + Manatidae, _44_ + + _Manatus_, _44_; + cervical vertebrae, 444; + dental formula, 425; + humerus, 501; + manus, 505; + pelvis, 514; + skull, 460; + sternum, 489; + teeth, 345; + thoraco-lumbar vertebrae, 448 + + Mandible, birds, 335; + cod, 100; + crocodile, 258; + dog, 398; + duck, 319; + frog, 160; + Hippopotamus, =467=; + Isabelline bear, =438=; + newt, 144; + salmon, 94; + turtle, 230 + + Manidae, _44_; + see _Manis_ + + _Manis_, _44_; + auditory ossicles, 487; + manus, 504; + scales, 3, 417; + skull, 459; + _M. macrura_ xiphisternum, 489; + see Pangolin + + Manubrium of malleus 486; + dog, 393; + -- sterni, dog 404 + + Manus, 26; + crocodile, 265; + dog, 408, =413=; + duck, 323; + frog, 164; + mammalia, 503; + newt, 147; + Perissodactyles, =508=; + turtle, 233 + + Marginal plate, turtle 216; + -- ray, 131; + -- shield, turtle, 214 + + Marmoset, _49_, 372 f. + + Marmot, frontals, 476 + + Marsipobranchii, _31_, _53_; + spinal column, 56 + + Marsupial bones, 513 + + Marsupial mole, 43; + see _Notoryctes_ + + Marsupialia, _43_; + arm bones, 499; + auditory ossicles, 486; + cervical vertebrae, 443; + caudal vertebrae, 453; + general characters, 349; + manus, 504; + pectoral girdle, 494; + pelvis, 513; + pes, 521; + ribs, 491; + sacral vertebrae, 451; + skull, 456; + teeth, 422; + thigh and shin, 517; + thoraco-lumbar vertebrae, 447 + + _Mastodon_, _47_, _365_; + teeth, 434 + + _Mastodonsaurus_, _35_, _136_; + pelvis, 187 + + Mastoid portion of periotic, dog, 391 + + Maxilla, 22; + cod, 98; + crocodile, 254; + dog, 397; + duck, 318; + frog, 159; + newt, 144; + turtle, 229 + + Maxillo-mandibular arch, 21 + + Maxillo-palatine, duck, 318 + + Maxillo-turbinal, dog, 395 + + Meatus, external auditory --, crocodile, 250; + dog, 393; + turtle, 228; + internal auditory --, crocodile, 251; + dog, 392, 400; + turtle, 228 + + Meckel's cartilage, 22; + cod, 100; + dogfish, 77; + salmon, 94 + + Median ethmoid, 21; + cod, 98; + Gymnophiona, 179; + salmon, 91; + -- fin, Amphibia, 52; + cod, 86; + dogfish, 79 + + Megachiroptera, _49_; + general characters, 371 + + _Megalobatrachus_, _35_, _135_; + carpus, 186; + skull, 175 + + Megalosauridae, _38_ + + _Megalosaurus_, _38_, _208_ + + _Megapodius_, spur, 330 + + _Megaptera_, _44_, _357_ + + Megatheriidae, _44_, 352; + humerus, 501; + leg bones, 517; + pelvis, 513; + sacrum, 452; + teeth, 424; + thoraco-lumbar vertebrae, 447 + + _Megatherium_, _44_; + femur, 517; + manus, 505; + pectoral girdle, 495; + pes, 522; + skull, 458 + + Megistanes, _40_, _299_ + + Membranous cranium, 17 + + Menobranchidae, _35_ + + _Menobranchus_, _35_, _135_; + carpus, 185; + pes, 188; + skull, 174; + teeth, 169 + + _Menopoma_, _35_; + see _Cryptobranchus_ + + Mento-meckelian, 22; + frog, 161; + reptiles, 284 + + Merganser, _41_; + beak, 329 + + _Mergus_, _41_ + + Merrythought, duck, 322 + + Mesethmoid, 20; + dog, 390; + duck, 317 + + _Mesoplodon_, _44_; + teeth, 427 + + Mesopterygium, 79 + + Mesosauridae, _37_ + + _Mesosaurus_, _37_ + + Mesosternum, dog, 404 + + Metacarpal quill, duck, 303 + + Metacarpo-digital, duck, 303 + + Metacarpus, 26; + see Manus + + Metacromion, hares and rabbits, 497 + + Meta-pterygium, 79 + + Metatarsus 26; + see Pes + + Metatheria, _43_; + general characters, 349 + + _Metriorhynchus_, _39_, 278 + + Microchiroptera, _49_; + general characters, 371 + + _Microgale_, _49_; + caudal vertebrae, 454 + + Mid-digital quill, duck, 303 + + Milk-teeth, 344; + dog, 377; + horse, 430 + + Moa, _40_, _299_; + aftershaft, 328; + pectoral girdle, 336; + wing, 338 + + Molar teeth, 344; + dog, 376 f. + + Mole, _49_, _370_; + auditory ossicles, 488; + cervical vertebrae, 446; + humerus, 503; + manus, 512; + presternum, 490; + shoulder girdle, 499; + skull, 481; + teeth, 440; + Golden --, _49_; + see _Chrysochloris_; + Marsupial --, _43_; + see _Notoryctes_ + + _Molge_, _35_, _135_; + see Newt + + Monitor, _38_; + see _Varanus_ + + Monkey, _49_, _373_; + see under Primates + + Monodelphia, _43_; + characters of, 351 + + _Monodon_, _45_, _357_; + see Narwhal + + Monophyodont, defined, 7, 344 + + Monopneumona, _34_ + + Monotremata, _42_; + arm bones, 499; + auditory ossicles, 486; + caudal vertebrae, 453; + cervical vertebrae, 443; + general characters, 346; + manus, 504; + pectoral girdle, 493; + pelvis, 513; + pes, 521; + ribs, 490; + sacral vertebrae, 451; + skull, 455; + sternum, 489; + teeth, 422; + thigh and shin, 517; + thoraco-lumbar vertebrae, 447 + + _Morosaurus_, _38_, _207_; + pes, 294 + + _Mosasaurus_, _38_, _204_ + + _Moschus_, _46_; + canines, 429 + + Mouse, _47_; + teeth, 437; + see _Mus_ + + Mud-fish, _34_ + + Multituberculata, _43_, _348_ + + Muntjac, _46_; + see _Cervulus_ + + Muraenidae, _33_ + + Muridae _47_ + + _Mus_, _47_; + _M. musculus_, teeth, 437; + _M. sylvaticus_, sternum and shoulder girdle, 498 + + Musk deer, _46_; + canines, 429 + + Mustelidae, _48_; + teeth 439 + + _Mycetes_, 49; + hyoid, 485; + mandible, 484; + skull, 482 + + Myliobatidae, _32_; + teeth, 109 + + Myomorpha, _47_ + + _Myrmecobius_, teeth, 423 + + _Myrmecophaga_, _44_; + manus 505; + pectoral girdle, 495; + skull, 458 + + Myrmecophagidae, _44_, 424; + see Anteaters + + Mystacoceti, _44_; + general characters, 356; + hind limb, 518; + manus, 505; + pectoral girdle, 495; + skull, 461; + teeth, 426 + + _Myxine_, _31_, 55; + fins, 115; + notochordal sheath, 9 + + Myxinoidei, _31_, 55 + + + Nails, 3; + Amphibia, 168; + mammals, 417 + + Nares: + anterior --, crocodile, 252, 257; + dog, 401; + duck, 317; + newt, 143; + turtle, 225, 229; + posterior --, crocodile, 257; + dog, 402; + duck, 318; + frog, 158; + newt, 143; + turtle, 230 + + Narial cavity, salmon, 89; + -- passage, crocodile, 254; + dog, 395; + -- septum, dog, 401 + + Narwhal, _45_, _357_; + teeth, 427 + + Nasal 21; + crocodile, 252; + dog, 394; + duck, 317; + frog, 158; + newt, 143; + turtle, 228; + -- capsule, 20; + cod, 97; + crocodile, 252; + dog, 394; + dogfish, 74; + frog, 158; + newt, 143; + turtle, 228; + -- cavity, dog, 388; + -- fossae, salmon, 89; + -- horns, rhinoceros, 3 + + Navicular, 27; + dog, 414 + + Neornithes, _40_; + general characters, 298 + + _Nesodon_, _46_, _361_; + pes, 525; + teeth, 432 + + Nesodontidae, _46_ + + Neural arch, 14; + -- plate, turtle, 215; + -- spine, 14 + + Neuromere, defined, 112 + + Newt, _35_; + anterior limb, 147; + hyoid apparatus or visceral arches, 144, =181=; + pelvic girdle, 149; + ribs, 145; + shoulder girdle, =146=; + skull, 140; + sternum, 145; + vertebral column, 138 + + Notidanidae, _31_; + calcification of vertebrae, 114; + pectoral fins, 130; + vertebral column, 113; + visceral arches, 63, 119 f. + + Notochord, _Amphioxus_ 52; + Balanoglossus, 50; + dogfish, 72; + Tunicates, 51 + + _Nothosaurus_, _37_, _193_; + supratemporal fossae, 283 + + Nothosauridae, _37_ + + _Notoryctes_, _43_; + arm bones, 500; + caudal vertebrae, 453; + cervical vertebrae, 443; + claws, 418; + dental formula, 423; + manus, 504; + pelvis, 513; + pectoral girdle, 494; + pes, 521; + ribs, 491; + sacrum, 452; + skull, 457; + sternum, 489; + thigh and shin, 517 + + Notoryctidae, _43_, _350_ + + Nuchal plate, turtle, 215; + -- shield, crocodile, 238; + turtle, 214 + + _Nyrania_, _35_; + palatines, 177 + + + Occipital condyle, crocodile, 246; + dog, 386; + duck, 315; + frog, 154; + turtle, 224; + -- crest, dog, 386; + -- segment, crocodile, 246; + dog, 384; + turtle, 224 + + _Odontaspis_, _32_; + succession of teeth, =107= + + Odontoblast, 7 + + Odontoceti, _44_; + general characters, 357; + manus, 505; + pectoral girdle, 495; + skull, 462; + sternum, 489; + teeth, 426 + + Odontolcae, _40_; + general characters, 299 + + _Odontopteryx_, _40_; + jaws, 334 + + _Ogmorhinus_, _48_; + mandibular ramus, =439= + + Olecranon process, dog, 406; + duck, 323; + frog, 164 + + Olfactory capsule, see nasal capsule; + -- cavity, dog, 388; + -- chamber, dog, 395; + -- fossa, dog, 390 + + Olm, _35_ + + _Omosaurus_, exoskeleton, 272 + + Omosternum, frog, 163 + + _Onychodactylus_, _35_; + nails, 168 + + Operculum, cod, 101; + salmon, 95 + + Ophidia, _38_; + general characters, 202; + jaw bones, 280; + scales, 270; + skull, 277 f.; + vertebral column, 275 + + _Ophisaurus_, _38_; + limbs, 289; + pectoral girdle, 289 + + Opisthocoelous, defined, 14 + + _Opisthocomus_, 41; + skull, 334 + + Opisthotic, 20; + cod, 96; + crocodile, 250; + salmon, 89 f.; + turtle, 227 + + Opossum, _43_; + caudal vertebrae, 453; + teeth, 423 + + Optic capsule, 20; + crocodile, 251; + dog, 394; + turtle, 228 + + Orang, _49_; + carpus, 512; + ribs, 493; + thoraco-lumbar vertebrae, 450 + + Orbit, crocodile, 257; + dogfish, 74; + duck, 317 + + Orbital ring, cod, 97; + salmon, 93 + + Orbitosphenoid, 19; + dog, 388; + duck, 317; + newt, 141 + + _Orca_, _45_; + teeth, 427 + + _Oreodon_, _45_; + see _Cotylops_ + + Ornithodelphia, _42_; + general characters, 346 + + Ornithosauria, _212_ + + Ornithorhynchidae, _43_ + + _Ornithorhynchus_, _43_; + beak, 3, 418; + caudal vertebrae, 453; + manus, 504; + pelvis, 513; + sacral vertebrae, 451; + shoulder girdle, =347=; + skull, 455; + spur, 418; + tarsus, 27 n.; + teeth, 4, 346, 422; + thoraco-lumbar vertebrae, 447 + + Ornithopoda, _39_; + general characters, 209 + + Orthopoda, _39_; + general characters, 208; + pubes, 292 + + Orycteropodidae, _44_; + teeth 425 + + _Orycteropus_, _44_; + hair, 416; + manus, 505; + pectoral girdle, 495; + pelvis, 513; + skull, 459; + see Aard Vark + + Osborn, H.F., on Mesozoic Mammals, 348 + + Os entoglossum, duck, 320 + + Osteoblast, 11 + + Osteoclast, 11 + + Osteodentine, 108 + + Osteostraci, _31_; + general characters, 54 + + _Ostracion_, _33_, 69; + plates, 105 + + Ostracionidae, _33_ + + Ostracodermi, _31_; + general characters, 54 + + Ostrich, _40_, _299_; + aftershaft, 329; + cervical vertebrae, =331=; + claws, 330; + foot, 342; + manus, 338; + pelvic girdle and sacrum, =340=; + pubis, 341; + tibio-tarsus, 341; + wing, =339= + + _Otaria_, _48_; + dentition, 439; + tympanic bulla, 480 + + Otariidae, _48_, _369_; + auditory ossicles, 488; + scapula, 498; + skull, 480 + + Owen's apteryx, pelvic girdle and sacrum, =340= + + Owen's chameleon, epidermal horns, 271 + + Owls, _42_, 335; + aftershaft, 329; + foot, 342 + + Owl-parrot, _42_; + see _Stringops_ + + Ox, _46_, 359; + atlas and axis, =445=; + three cervical vertebrae, =15=; + femur, =518=; + manus, 507; + teeth, 345; + two thoracic vertebrae, =449= + + + Paca, _48_ + + Paired fins, 127 + + Palaeoniscidae, _32_ + + _Palaeoniscus_, _32_; + scales, 67 + + Palaeospondylidae, _31_ + + _Palaeospondylus_, _31_, 58 + + _Palaeosyops_, _46_; + teeth, 432 + + Palaeotheriidae, _46_ + + _Palaeotherium_, 46; + skull, 471; + teeth, 430 + + _Palamedea_, _41_; + spur, 330, 338 + + Palamedeae, _41_ + + Palate, reptiles, 280 f. + + Palatine, cod, 98; + crocodile, 254; + dog, 397; + duck, 318; + frog, 160; + salmon, 93; + turtle, 230 + + Palato-pterygo-quadrate bar, 22; + dogfish, 77; + fish, 120 f.; + salmon, 93 + + Palm civet, _48_ + + Pangolin, _44_; + pectoral girdle, 495; + pelvis, 513; + caudal vertebrae, 453; + see _Manis_ + + Parachordals, 17 + + _Paradoxurus_, _48_; + tail, 454 + + Parasphenoid, 21; + cod, 97; + frog, 156; + newt, 141; + reptiles, 278; + salmon, 93 + + Parasuchia, _39_; + general characters, 211 + + Parethmoid, 21 n. + + Pariasauria, _36_ + + _Pariasaurus_, _36_, _192_; + pectoral girdle, 289; + pelvis, 292; + supratemporal fossa, 283; + teeth, 273 + + Parietal, 19; + cod, 96; + crocodile, 247; + dog, 386; + duck, 314; + newt, 141; + salmon, 91; + turtle, 225; + -- segment, crocodile, 247; + dog, 386; + turtle, 225 + + Paroccipital process, dog, 386 + + _Parra_, _42_; + spur, 330 + + Parrots, 335; + aftershaft, 328; + beak, 330; + epiphyses of centra, 332; + foot, 342; + powder-down feathers, 329; + skull, 334 + + Parrot fish, _33_; + see _Scarus_ + + Passeres, aftershaft, 328 + + Passeriformes, _42_ + + Patella, dog, 412; + duck, 327 + + _Pavo_, _41_; + _P. cristatus_, shoulder girdle and sternum, =337= + + Peacock, _41_; + see _Pavo_ + + Peccary, pes, 523 + + Pecora, _46_, _359_; + teeth, 429 + + Pectinated incisors, _Galeopithecus_, 370, 440; + _Procavia_, 362 + + Pectineal process, duck, 326 + + Pectoral fins, cod, 103; + dogfish, 79; + -- girdle, 24; + Amphibia, 184; + birds, 336; + cod, 101; + crocodile, 262; + dog, 404; + dogfish, 79; + duck, 321; + fish, 126; + frog, 162; + mammalia, 493; + newt, 145; + reptiles, 288; + turtle, 231; + -- shield, turtle, 215 + + _Pedetes_, _47_; + manus, 511; + tail, 454 + + Pelican, _41_, 335; + clavicles, 338 + + _Pelicanus_, _41_; + _P. conspicillatus_ shoulder girdle and sternum, =337= + + _Pelobates_, _36_; + vertebrae, 172; + _P. cultripes_ teeth, 169 + + Pelobatidae, _36_ + + Pelvic fins, cod, 103; + dogfish, 82; + fish, 131; + -- girdle, 25; + Amphibia, 187; + birds, 339; + crocodile, 266; + dog, 409; + dogfish, 81; + duck, 324; + fish, 127; + frog, 165; + mammals, 512; + newt, 149; + Ratitae, =340=; + Reptilia, 291; + turtle, 235 + + Penguin, _40_; + distribution of feathers, 328; + fibula, 341; + foot, 342; + manus, 338; + metatarsus, 342; + pneumaticity of skeleton, 331; + skull, 333; + sternum, 336; + thoracic vertebrae, 332; + wing, 329, =339= + + Penna, duck, 303 + + Pentedactylate, defined, 26 + + _Perameles_, _43_; + atlas, 443; + pectoral girdle, 494 + + Peramelidae, _43_, _350_; + auditory ossicles, 486; + pes, 522 + + _Perca_, _34_ + + Perch, _34_; + pelvic fin, 132; + urostyle, 117 + + Percidae, _34_ + + Perennibranchiata, _35_; + characters, 135 + + Perichondrium, 10 + + Perichordal sheath, 16 + + Periosteal ossification, 10 + + Periosteum, 10 + + Periotic, dog, 390; + -- capsule, see Auditory capsule + + Perissodactyla, _46_; + cervical vertebrae, 445; + general characters, 359; + manus, 507; + pes, 523; + ribs, 491; + scapula, 496; + skull, 470; + teeth, 429; + thoraco-lumbar vertebrae, 448 + + Persistent pulps, 5 + + Pes, 26; + crocodile, 268; + dog, =413=, 414; + duck, 327; + frog, 166; + mammals, 521; + reptiles, 293; + turtle, 236; + of Tapir, Rhinoceros, _Hipparion_ and Horse, =524= + + _Petromyzon_, _31_, 55 f.; + notochordal sheath, 9 + + Petromyzontidae, _31_, 55 + + Petrous portion of periotic, dog, 391 + + _Pezophaps_, _42_; + see Solitaire + + _Phacochaerus_, _45_; + teeth, 428 + + _Phaëthon_, _41_; + metatarsals, 342 + + _Phalacrocorax_, _41_ + + Phalangeridae, _43_, 350 + + Phalanges, 26; + see Manus and Pes + + Phaneroglossa, _36_ + + Pharyngo-branchial, cod, 101; + dogfish, 78; + salmon, 95 + + Pharyngognathi, _33_ + + _Phascolarctus_, _43_; + see Koala + + Phascolomyidae, _43_, _350_ + + _Phascolomys_, _43_, 349; + see Wombat + + _Phascolotherium_, _43_, 348 + + Phenacodontidae, _47_ + + _Phenacodus_, _47_, _362_; + caudal vertebrae, 454; + manus, =510=; + pes, 525; + scapula, 497; + skull, 472; + thoraco-lumbar vertebrae, 449 + + _Phocaena_, _45_, _357_; + skull, 462; + thoraco-lumbar vertebrae, 448; + _P. phocaenoides_, ossicles, 420 + + Phocidae, _48_, _369_; + scapula, 497; + tympanic bulla, 480 + + _Phoronis_, _30_, 50 f. + + _Phororhacos_, _41_; + anterior nares, 333; + ischia, 341 + + _Physeter_, _44_; + cervical vertebrae, 444; + manus, 505; + skull, 464; + teeth, 426 + + Physeteridae, _44_; + ribs, 491; + thoraco-lumbar vertebrae, 448 + + _Physodon_, _44_; + teeth, 426 + + Physodontidae, _44_ + + Physostomi, _33_ + + Phytosauridae, _39_ + + _Phytosaurus_, _39_; + see _Belodon_ + + Pici, _42_ + + _Picus_, _42_; + see Woodpecker + + Pig, _45_, _359_; + skull, 465 f., =466=; + teeth, 345, 427 + + Pigeons, _42_, 334 f.; + aftershaft, 329; + pneumaticity of skeleton, 331 + + Pike, _33_; + pelvic fin, 132; + teeth, 107, 110 + + Pinnipedia, _48_; + arm bones, 502; + auditory ossicles, 488; + general characters, 369; + manus, 511; + pelvis, 515; + pes, 526; + skull, 480; + teeth, 439; + thigh and shin, 520; + thoraco-lumbar vertebrae, 450 + + _Pipa_, _36_; + hyoid apparatus, 182; + jaws, 169; + skull, 180; + sternum, 184; + vertebrae, 172 + + Pipidae, _36_ + + Pisces, _31_; + general characters, 60 + + Piscivorous dentition, 426, 440 + + Pisiform, 345, 504; + crocodile, 265; + dog, 408; + turtle, 233 + + Pituitary fossa, crocodile, 247; + -- space, 17 + + Placodontia, _36_ + + _Placodus_, _36_, _192_; + teeth 273 + + Placoid scale, 4, 60, 104 + + Plantigrade, defined, 358 n. + + Plastron, _Dermochelys_, 272; + _Chelone midas_, 217, =218=, 271 + + Platanistidae, _45_ + + _Platanista_, _45_; + cervical vertebrae, 444; + skull, 464 + + Plectognathi, _33_; + vertebrae, 115 + + _Plectropterus_, _41_; + _P. gambensis_, spur, 330 + + Plesiosauridae, _37_; + limbs, 193; + parasphenoid, 192; + skull, 278 + + _Plesiosaurus_, _37_, _193_; + position of limbs, 28 + + Pleuracanthidae, _63_; + fins, 115 + + Pleurodira, _37_; + general characters, 195 + + Pleurodont, 159, 199, 273 + + Pleuronectidae, _33_ + + Pleuropterygii, _31_, 63 + + _Pliosaurus_, _37_, _193_ + + Plovers, _42_, 334; + thoracic vertebrae, 332 + + Pneumaticity of bird's skeleton, 331 + + _Polacanthus_, _39_; + exoskeleton, 272 + + Pollex, 26; + see Manus + + _Polyodon_, _32_, 104; + distribution, 66; + pectoral fins, 131; + skull, 122; + spinal column, 112; + teeth, 110 + + Polyodontidae, _32_ + + _Polyonax_, _39_, _209_; + beak, 271; + frontals, 277; + jaw, 274; + predentary, 284 + + Polyprotodont, 423 + + Polyprotodontia, _43_; + general characters, 350 + + Polypteridae, _33_ + + _Polypterus_, _33_, _68_; + distribution, 66; + exoskeleton, 67; + pectoral fins, 131; + pelvic fins, 132; + pelvis, 127; + scales, 104; + skull, 122; + tail, 116 + + _Pontoporia_, _45_; + cervical vertebrae, 444; + teeth, 426 + + Porcupine, _47_; + pes, 526; + skull, 476, =477=; + spines, 417 + + Porpoise, _45_, _357_; + thoraco-lumbar vertebrae, 448 + + Postaxial, 28 + + Posterior cornu, duck, 320; + turtle, 231; + -- limb, 26; + Amphibia, 188; + birds, 341; + dog, 412; + duck, 326; + frog, 166; + newt, 149, =148=; + reptiles, 293; + turtle, 235, =234= + + Postfrontal, 21; + crocodile, 250; + turtle, 225 + + Postorbital bar, crocodile, 250, 255 f.; + _Hatteria_, 283; + -- groove, dogfish, 76 + + Post-temporal, cod, 102; + reptiles, 283; + -- bar, crocodile, 256; + _Hatteria_, 283 + + _Potamogale_, _49_, 367, 370; + shoulder girdle, 499; + teeth, 440 + + Potamogalidae, _49_ + + Powder-down feathers, 329 + + Pre-axial, 28 + + Precoracoid, 25; + frog, 163; + newt, 147; + reptiles, 288; + turtle, 232 + + Predentary, reptiles, 284 + + Predigital quill, duck, 303 + + Prefrontal, 21; + crocodile, 249; + reptiles, 278; + turtle, 225 + + Prefronto-lachrymal, newt, 141 + + Prehallux, frog, 167 f. + + Premaxilla, 22; + cod, 98; + crocodile, 252; + dog, 398; + duck, 314, 318; + frog, 158; + newt, 143; + salmon, 94; + turtle, 230 + + Premolar, dog, 370, 377; + mammals, 344 + + Prenasal process, frog, 158 + + Pre-orbital vacuity, reptiles, 283 + + Presphenoid, 19; + dog, 388 + + Prespiracular ligament, dogfish, 77 + + Presternum, dog, 404 + + Primaries, duck, 303 + + Primates, _49_; + arm bones, 503; + auditory ossicles, 488; + cervical vertebrae, 446; + general characters, 372; + manus, 512; + pelvis, 515; + pes, 527; + ribs, 493; + sacrum, 452; + shoulder girdle, 499; + skull, 482 f.; + sternum, 490; + tail, 454; + teeth, 441; + thigh and shin, 520; + thoraco-lumbar vertebrae, 450 + + _Priodon_, _44_; + caudal vertebrae, 453; + manus, 505; + stapes, 487; + sternum, 489; + teeth, 424 + + Pristidae, _32_ + + _Pristis_, _32_; + snout or rostrum, 109, 119 + + Proboscidea, _47_; + arm bones, 502; + cervical vertebrae, 445; + general characters, 364; + femur, 519; + manus, 511; + pelvis, 514; + pes, 526; + scapula, 497; + skull, 473; + teeth, 433; + thoraco-lumbar vertebrae, 449 + + _Procavia_, _47_, _363_; + auditory ossicles, 487; + caudal vertebrae, 453; + dental formula, 432; + humerus, 502; + manus, 510; + pelvis, 514; + pes, 525; + ribs, 491; + scapula, 497; + skull, 433, 472; + tarsus, 27; + thoraco-lumbar vertebrae, 449 + + Process, alinasal --, frog, 158; + basi-pterygoid --, birds, 334; + coracoid --, dog, 405; + coronoid -- (of mandible), dog, 398; + duck, 319; + coronoid -- (of ulna), dog, 408; + pectineal --, duck, 326; + postfrontal --, duck, 316; + postglenoid --, dog, 394; + postorbital -- (of frontal), dog, 388; + postorbital -- (of jugal), dog, 398; + posterior articular --, duck, 319; + zygomatic --, dog, 394 + + Processus brevis, 486; + -- gracilis, 486; + -- longus, 486; + dog, 393 + + Procoelous, defined, 14 + + _Prodelphinus_, _45_; + skull, 462 + + Proganosauria, _37_ + + Prone position, 29 + + Prongbuck, _46_; + horns, 417 + + Pro-otic, 20; + frog, 157; + turtle, 227 + + Pro-pterygium, dogfish, 79 + + Proteidae, _35_ + + _Proteles_, _48_; + teeth, 437 + + Protelidae, _48_ + + Proterosauridae, _37_ + + _Proterosaurus_, _37_; + teeth, 198, 274; + vertebrae, 197 + + _Proteus_, _35_, 135, 182; + branchial arches, 180; + digits, 187; + pes, 188; + skull, 174 + + _Protopterus_, _34_, _70_, 117; + branchial arches, 121, 124; + fins, 130; + skull, 124; + vestigial gill on pectoral girdle, 129 + + Prototheria, _42_; + general characters, 346 + + Proximal, defined, 23 n. + + _Psephurus_, distribution, 66 + + _Pseudopus_, _38_; + limbs, 289 + + Psittaci, _42_; + see Parrots + + _Pteranodon_, _39_, _274_; + pectoral girdle, 289 + + Pteranodontidae, _39_ + + _Pteraspis_, _31_, _54_ + + _Pterichthys_, _31_, _55_ + + _Pterocles_, _42_; + see Sandgrouse + + Pteroclidae, _42_ + + Pterodactylidae, _39_ + + _Pterodactylus_, _39_, 213 + + Pteropidae, _49_; + skull, 481 + + _Pteropus_, _49_; + dental formula, 441; + tail, 454 + + Pterosauria, _39_; + general characters, 212; + ischia, 292; + limbs, 291; + pre-orbital vacuity, 284; + ribs, 285; + sternum, 287; + vertebrae, 275 f. + + Pterotic, 20; + cod, 96; + salmon, 90 f. + + Pterygoid, cod, 98; + crocodile, 255; + dog, 397; + duck, 318; + frog, 160; + newt, 144; + salmon, 93; + turtle, 230; + -- fossa, crocodile, 255; + -- plate, dog, 388 + + Pterylae, 328 + + Pubis, 25; + crocodile, 266 f.; + duck, 325; + dog, 411; + frog, 165; + newt, 149; + reptiles, 292; + turtle, 235 + + Pygal plate, turtle, 217; + -- shield, turtle, 214 + + Pygopodidae, _37_ + + Pygostyle, duck, 307, 312 + + _Python_, _38_; + ischio-pubis, 292; + jaws, 280; + vestiges of limbs, 289, 293 + + Pythonomorpha, _38_; + general characters, 204; + limbs, 290; + teeth, 273 + + + Quadrate, 22; + cod, 98; + crocodile, 255; + duck, 319; + frog, 160; + newt, 144; + salmon, 93; + turtle, 229 + + Quadratojugal, 22; + crocodile, 255; + duck, 318; + frog, 160; + turtle, 229 + + Quill, duck, 302 + + + Rabbit, _48_, 366; + pollex, 511 + + Raccoon, 369 + + Rachis, duck, 302 + + Rachitomous, defined, 171 + + Radiale, 27; + see Carpus + + Radialia, 115; + dogfish, 79 f. + + Radio-ulna, frog, 164 + + Radius, 26; + crocodile, 265; + dog, 406; + duck, 323; + newt, 147; + turtle, 233 + + _Raia_, _32_; + calcification of vertebrae, 114 + + Raiidae, _32_ + + _Rana_, _36_; + see Frog + + Ranidae, _36_; + shoulder girdle, 185 + + Rat, pes, 526 + + Ratitae, 40; + caudal vertebrae, 333; + clavicles, 338; + foot, 342; + general characters, 298; + skull, 333; + sternum, 336; + vomers, 334; + wing, 338 + + Rattlesnake, _38_; + rattle, 3, 270 + + Ray, pectoral fin, 130; + Eagle --, Electric -- and Sting --, _32_ + + Rectrices, 303, 329 + + Reed-fish, _33_ + + Reindeer, antlers, 469 + + Remicle, duck, 304 + + Remiges, 303, 329 + + Reptiles, anterior limb, 290; + exoskeleton, 270; + fossae in skull, 281; + pectoral girdle, 288; + pelvic girdle, 291; + posterior limb, 293; + ribs, 285; + skull, 276; + sternum, 287; + teeth, 272; + vertebral column, 275 + + Reptilia, _36_; + general characters, 190; + see Reptiles + + _Rhabdopleura_, _30_, 50 + + _Rhamphastos_, _42_; + see Toucan + + Rhamphorhynchidae, _39_ + + _Rhamphorhynchus_, _39_, 213, 274 + + _Rhea_, _40_; + aftershaft, 329; + claws, 330; + ischia, 341; + manus, 338; + _R. macrorhyncha_, pelvic girdle and sacrum, =340= + + Rheornithes, _40_ + + _Rhina_, _32_; + see _Squatina_ + + Rhinal process, frog, 158 + + _Rhinoceros_, _46_, 360, 419; + femur, =518=; + fibula, 519; + malleus, 487; + manus, 508; + nasal horns, 3, 417; + pes, 525; + skull, =421=, 470; + teeth, 430; + ulna, 501; + _R. antiquitatis_, 470 + + Rhinocerotidae, _46_ + + Rhinolophidae, _49_ + + Rhiptoglossa, _38_ + + Rhizodontidae, _33_ + + _Rhizodus_, _33_; + teeth, 110 + + Rhynchocephalia, _37_; + general characters, 197; + humerus, 290; + teeth, 273 f.; + vertebrae, 275 + + Rhynchosauridae, _37_; + maxillae, 198 + + _Rhytina_, _44_, 352, 425; + humerus, 501; + skull, =460= + + Rhytinidae, _44_ + + Ribs, 23; + Amphibia, 182; + birds, 336; + cod, 86; + crocodile, 259; + dog, 402; + dogfish, 73; + duck, 320; + fish, 125; + frog, 153; + mammalia, 490; + newt, 145; + reptiles, 285 + + Ridge, supra-orbital and suborbital, dogfish, 74 + + Rodentia, _47_; + auditory ossicles, 488; + cervical vertebrae, 446; + dental formula, 435; + general characters, 365; + pelvis, 515; + pes, 526; + humerus, 502; + manus, 511; + ribs, 493; + sacrum, 452; + shoulder girdle, 497; + skull, 476; + sternum, 489; + tail, 454; + teeth, 421; + thigh and shin, 520; + thoraco-lumbar vertebrae, 449 + + Roller, _42_, 335 + + Rooted teeth, defined, 5 + + Rorqual, _44_, 357; + cervical vertebrae, 444 + + Rostrum, crocodile, 247; + dogfish, 74; + duck, 316; + _Pristis_, 119; + -- of sternum, duck, 321 + + Röse, C., on teeth of Marsupials, 422 + + Ruminantia, _46_, _359_; + auditory ossicles, 487; + fibula, 519; + horny plates on palate, 418; + hyoid, 470; + manus, 507; + odontoid process, 445; + pes, 523; + scapula, 495; + teeth, 420, 429 + + + Sabre-toothed lion, _48_; + see _Machaerodus_ + + Sacral ribs, crocodile, 243; + -- surface of ilium, dog, 409; + -- vertebrae, 16; + crocodile, 243; + dog, 383; + duck, 312; + frog, 153; + newt, 140; + turtle, 222 + + Sacrum, duck, 310; + see Sacral vertebrae + + Sagittal crest, dog, 386 + + _Saiga_, skull, 468 + + Salamander, _35_ + + _Salamandra_, _35_, _135_; + antibrachium and manus of larva, =186=; + manus of larva, 185; + tarsus, 27 + + Salamandrina, _35_, _135_; + skull, 175; + sternum, 182 + + _Salmo_, _33_ + + Salmon, _33_; + branchial arches, 95; + chondrocranium, 87; + opercular bones, 95; + pectoral fins, 131; + skull, 87 + + Salmonidae, _33_ + + Sandgrouse, _42_, 335 + + _Sarcophilus_, _43_; + teeth, 423 + + Sauropoda, _38_; + general characters, 205; + teeth, 273; + vertebrae, 276 + + Sauropsida, _36_; + general characters, 189 + + Sauropterygia, _37_; + general characters, 192; + limbs, 290; + palate, 281; + pectoral girdle, 288; + vertebrae, 276 + + Saw-fish, _32_; + see _Pristis_ + + Scales, cod, 83; + crocodile, 237; + ctenoid, 8; + cycloid, 8; + duck, 302; + ganoid, 8; + Gymnophiona, 168; + mammals, 417 + + Scale-foot, 37 + + Scalpriform, 366 + + _Scaphirhynchus_, _32_, _104_; + distribution, 66; + exoskeleton, 67; + spinal column, 112 + + Scaphoid, 27; + mammals, 504 f. + + Scapho-lunar, dog, 408 + + Scapula, 25; + cod, 103; + crocodile, 263; + dog, 404; + duck, 322; + frog, 162; + newt, 146; + turtle, 232 + + Scapular shield, armadillo, 419 + + Scapus, duck, 302 + + _Scarus_, _33_; + beaks, 111 + + Scelidosauridae, _39_ + + Schizognathous, defined, 335 + + Scincidae, _38_ + + _Scincus_, _38_; + scutes, 271 + + Sciuromorpha, _47_ + + Sclerotic, turtle, 228 + + Screamer, _41_; + spurs, 330 + + Scutes, armadillos, 419; + crocodile, 237; + reptiles, 271 + + Scylliidae, _32_ + + _Scyllium_, _32_; + calcification of vertebrae, 114; + pectoral fins, 130; + suspensorium, 119; + see Dogfish + + _Scymnus_, _32_, _118_; + calcification of vertebrae, 114; + mandibular arch, 120; + pectoral fins, 130 + + _Scythrops_, _42_; + interorbital septum, 333 + + Sea leopard, _48_; + see _Ogmorhinus_; + -- lion, _48_; + manus, 511; + pes, 526; + position of limbs, 29; + -- otter, _48_; + pes, 526 + + Seal, _369_; + manus, 511; + pes, 526; + sacral vertebrae, 452; + scapula, 497 + + Secondaries, duck, 303 f. + + Secretary-bird, _41_; + claws, 330 + + Selachii, _31_; + general characters, 63; + teeth, 108 + + Selenodont, defined, 345, 428 + + Sella turcica, crocodile, 247; + dog, 386 + + Semionotidae, _33_ + + Semiplumae, 328 + + Sense capsules, see Auditory, Nasal and Optic capsule + + _Seps_, 38; + limbs, 289 + + Shagreen, 61 + + Shaft of feather, 302 + + Shark, 64; + Frill-gilled --, _31_; + see _Chlamydoselache_; + Port Jackson --, _32_; + see _Cestracion_ + + Sheep, _359_; + manus, 507; + teeth, 345 + + Shields of turtle, 214 + + Shin, 26; + see Crus + + Shoulder girdle, see Pectoral girdle + + Shrew, _49_, _370_; + auditory ossicles, 488; + cervical vertebrae, 446; + presternum, 490; + skull, 481; teeth, 440 + + Sigmoid notch, dog, 406 + + Siluridae, _33_; + plates, 105 + + _Simia_, _49_; + ribs, 493; + skull, 484; + thoraco-lumbar vertebrae, 450 + + Simiidae, _49_, _373_ + + Simplicidentata, _47_, 366 + + _Siphonops_, _35_; + _S. annulatus_, skull, =178= + + _Siredon_, _35_; + skull, 175; + teeth, 169; + visceral arches, =181= + + _Siren_, _35_, _135_, 188; + beaks, 168; + branchial arches, 180; + digits, 187; + skull, 174; + teeth, 169 + + Sirenia, _44_, 522; + arm bones, 501; + caudal vertebrae, 453; + cervical vertebrae, 443; + general characters, 352; + hair, 416; + horny plates, 418; + manus, 505; + pectoral girdle, 495; + pelvis, 514; + ribs, 491; + skull, 459; + sternum, 489; + teeth, 425; + thoraco-lumbar vertebrae, 448 + + Sirenidae, _35_ + + Sirenoidei, _34_; + general characters, 70 + + _Sivatherium_, _46_; + skull, 469 + + Skate, _32_ + + Skeletogenous layer, 14, 16; + _Amphioxus_, 52, 112 + + Skeleton, defined, 1; + Cape Buffalo, =492=; + _Ceratodus_, =128=; + cod, 83 f.; + crocodile, 237 f.; + dog, 374 f.; + duck, 302 f.; + frog, 151 f.; + llama, =496=; + newt, 138 f.; + turtle, 214 f. + + Skink, _38_; + see _Tiliqua_ + + Skull, 16 f.; + Amphibia, 173 f.; + Anura, 179 f.; + birds, 333 f.; + cod, 96 f.; + crocodile, 243 f.; + diagram of Mammalian, 385; + Dipnoi, 124; + dog, 383 f.; + dogfish, 73 f.; + donkey, =431=; + duck, 312 f.; + fish, 117; + frog, 154 f., =159=; + _Globicephalus_, =463=; + Gymnophiona, 177; + Indian elephant, =474=; + Mammalia, 455; + Marsipobranchii, 57; + pig, =466=; + _Procavia_, =433=; + reptiles, 276 f.; + Rhinoceros, =421=; + _Rhytina_, =460=; + sloth, =458=; + Tasmanian wolf, =456=; + Teleostei, 124; + turtle, 222 f.; + wombat, =456= + + Sloth, _43_, _352_; + auditory ossicles, 487; + arm bones, 500; + claws, 418; + leg bones, 517; + manus, 505; + pectoral girdle, 495; + pelvis, 513; + pes, 522; + ribs, 491; + sacrum, 452; + skull, 457; + sternum, 489; + teeth, 424; + thoraco-lumbar vertebrae, 447 + + Snake, _38_; + see Ophidia + + Sole, _33_ + + _Solea_, _33_ + + _Solenodon_, _49_; + teeth, 440 + + Solenodontidae, _49_ + + Solitaire, _42_, _330_; + wing, 338; + wrist, 330 + + _Sorex_, _49_; + pelvis, 515; + see Shrew + + Soricidae, _49_; + skull, 481 + + Spalacidae, _47_ + + _Spatularia_, _32_; + distribution, 66 + + _Spelerpes_, _35_; + branchial arches, 180; + ribs, 182; + _S. belli_ teeth, 169 + + Sperm whale, _44_, _357_; + see _Physeter_ + + _Sphargis_, _37_; + see _Dermochelys_ + + Sphenethmoid, frog, 156 + + Sphenisci, _40_; + see Penguins + + Sphenisciformes, _40_ + + _Sphenodon_, _37_, 197 f.; + carpus, 291; + cervical vertebrae, 275; + fossae in skull, 281; + humerus, 290; + interparietal foramen, 277; + ribs, 286; + skull, =282=; + tarsus, 293; + teeth, 274 + + Sphenodontidae, _37_ + + Sphenoidal fissure, dog, 388 + + Sphenotic, 20; + cod, 97; + salmon, 89 + + Spider monkey, _49_; + see _Ateles_ + + Spinacidae, _32_ + + Spinal column, 13; + Dipnoi, 113; + fish, 112; + Holocephali, 113; + Marsipobranchii, 56 + + Spines, Elasmobranchs, 61; + mammals, 417 + + Spiny ant-eater, _43_; + see _Echidna_; + -- mouse, 47; + see _Acanthomys_ + + Splenial, 22; + crocodile, 258; + duck, 320; + turtle, 231 + + Spurs, birds, 330; + Monotremata, 418 + + Spur-winged goose, 330; + -- plover, 330 + + Squalidae _31_, _64_ + + _Squalodon_, _45_, _357_; + dental formula, 427 + + Squalodontidae, _45_ + + Squamata, _37_; + general characters, 198; + position of teeth, 272; + skull, 278 + + Squamosal, 21; + crocodile, 256; + dog, 394; + duck, 316; + frog, 160; + newt, 144; + turtle, 229 + + _Squatina_, _32_; + calcification of vertebrae, 114; + labial cartilages, 119; + tail, 63; + vertebral column, 114 + + Squatinidae, _32_ + + Squirrels, frontals, 476; + pes, 526 + + Stapes, dog, 393; + frog, 157; + man, dog, rabbit, =485=; + newt, 141 + + Steganopodes, _41_ + + Stegosauria, _39_; + general characters, 209 + + Stegosauridae, _39_ + + _Stegosaurus_, _39_, _208_ f.; + exoskeleton, 272 + + Steller's sea-cow, _44_; + see _Rhytina_ + + Stereornithes, _41_ + + Stereospondyli, _35_ + + Sternal ribs, crocodile, 259; + dog, 402; + duck, 320; + mammals, 490 f. + + Sternebra, dog, 404 + + Sternum, 24; + Amphibia, 182; + birds, 336; + crocodile, 260, =261=; + dog, =403=; + duck, 321; + frog, 163; + Mammalia, 489; + newt, 145; + reptiles, 287 + + Stork, 335; + White --, _41_ + + Striges, _42_; + see Owls + + _Stringops_, _42_; + sternum, 336 + + _Struthio_, _40_, _299_; + see Ostrich + + Struthiornithes, _40_, _299_ + + Sturgeon, _32_; + see _Acipenser_ + + Stylo-hyal, dog, 399 + + Suborbital bar, duck, 318; + -- ridge, dogfish, 76 + + Subplantigrade, defined, 358 n. + + Subungulata, _46_; + arm bones, 502; + general characters, 360; + manus, 509; + pelvis, 514; + pes, 525; + shoulder girdle, 497; + skull, 471; + teeth, 432; + thigh and shin, 519 + + Suidae, _45_ + + Suina, _45_, 358 f.; + fibula, 519; + manus, 507; + odontoid process, 445; + pelvis, 514; + pes, 523; + ulna, 501 + + Sula, _41_; + see Gannet + + Supinator ridge, dog, 406 + + Supine position, defined, 29 + + Supra-angular, 22; + crocodile, 258; + duck, 319; + turtle, 230 f. + + Supracaudal shield, turtle, 214 + + Supraclavicle, cod, 102 + + Supra-occipital, 19; + crocodile, 247; + dog, 386; + duck, 315; + turtle, 224 + + Supra-orbital, 20; + crocodile, 251 + + Suprapharyngeal bone, cod, 101 + + Suprascapula, crocodile, 263; + frog, 162 + + Supratemporal arcade, crocodile, 256; + reptiles, 281 + + Surinam toad, _36_; + see _Pipa_ + + Sus, _45_; + dental formula, 428; + see Pig + + Suspensorium, Amphibia, 173; + dogfish, 78; + duck, 319; + frog, 160; + newt, 144; + Pisces, 61 + + Sutures, 12 + + Swan, _41_; + cervical and thoracic vertebrae, 332 + + Swift, _42_, 335; + foot, 342 + + Symplectic, cod, 100; + salmon, 94 + + + Tails, fish, 60 + + Talpa, _49_; + pelvis, 515; + see Mole + + Talpidae, _49_ + + Tapir, _46_, 360; + malleus, 487; + pes, =524=, 525; + teeth, 345; + see _Tapirus_ + + Tapiridae, _46_; + dental formula, 429 + + _Tapirus_, _46_; + fibula, 519; + manus, =508=; + skull, 471; + see Tapir + + Tarsier, _49_ + + Tarsiidae, _49_ + + _Tarsipes_, _43_, 349; + mandible, 457 + + _Tarsius_, _49_, 372; + pes, 527 + + Tarso-metatarsus, duck, 327 + + Tarsus, 26 f.; + crocodile, 268; + dog, 414; + frog, 166; + newt, 150; + turtle, 236 + + Tasmanian devil, _43_; + see _Sarcophilus_; + -- wolf, _43_; + see _Thylacinus_ + + _Tatusia_, _44_; + stapes, 487; + teeth, 424 + + Tectospondyli, 114 + + Tectrices, duck, 306 + + Teeth, =6=; + Amphibia, 169; + birds, 330; + cod, 83; + crocodile, 238; + development, 7; + dog, 374 f.; + fish, 106 f.; + frog, 158 f.; + horses, 5; + mammals, 344, 420 f.; + pharyngeal, 8; + reptiles, 272 f.; + structure, 4; + succession, 7 + + Teleosauridae, _39_ + + _Teleosaurus_, _39_; + palate, 281; + scutes, 271; + vertebrae, 275 + + Teleostei, _33_; + general characters, 69; + ribs, 126; + skull, 124; + tail, 117; + teeth, 110; + vertebral column, 115 + + Temnospondyli, _35_ + + Tenrec, _49_; + see _Centetes_ + + Tentorium, dog, 392 + + Terrapin, _37_ + + _Testudo_, _37_, _194_ + + _Tetraceros_, _46_; + horns, 417 + + _Thalassochelys_, carapace, =216= + + Thecodont, defined, 273 + + Theriodontia, _36_ + + Theromorpha, _36_; + general characters, 191; + humerus, 290; + pectoral girdle, 288; + ribs, 285; + skull, 278; + teeth, 273; + vertebral column, 275 f. + + Theropoda, _38_; + general characters, 207; + teeth, 273 + + Thoracic ribs, crocodile, 259; + see Ribs; + -- vertebrae, 16; + crocodile, 241, =242=; + dog, 381, =382=; + duck, 310; + turtle, 221 + + Thoraco-lumbar vertebrae, mammals, 447 f. + + Thornback skate, 104 + + _Thylacinus_, _43_; + atlas, 443; + dental formula, 423; + pelvis, 513; + pes, 521; + skull, =456= + + _Thylacoleo_, _43_; + skull, 457 + + Thyro-hyal, dog, 399 + + Tibia, 26; + crocodile, 268; + dog, 412; + newt, 149; + turtle, 235 + + Tibiale 27; + see Tarsus + + Tibio-fibula, frog, 166 + + Tibio-tarsus, duck, 326 + + Tichorhine Rhinoceros, 470 + + Tiger, _48_ + + _Tiliqua_, _38_; + scutes, 200, 271 + + Tillodontia _47_, 365; + femur, 520; + manus, 511; + teeth, 435 + + Tinamidae 300; + caudal vertebrae, 333; + vomers, 334 + + Tinamiformes, _41_ + + _Tinamus_, _41_; + ischia, 341 + + Titanotheriidae, _46_; + skull, 470; + teeth, 432 + + _Titanotherium_, _46_; + humerus, 501; + manus, =508=; + pes, 525 + + Toad, _36_; + shoulder girdle, 185 + + Tope, _32_ + + Torpedinidae, _32_ + + _Torpedo_, _32_, 104 + + Tortoise, _37_; + position of limbs, 28 + + _Tortrix_, ischio-pubis, 292; + traces of posterior limb, 293 + + Toucan, _42_; + foot, 342 + + _Toxodon_, _46_, 361; + femur, 519; + pes, 525; + teeth, 432 + + Toxodontia, _46_; + general characters, 361; + manus, 509; + skull, 472; + teeth, 432 + + Toxodontidae, _46_ + + Trabeculae 11; + -- cranii, 17 + + Tragulidae, _45_ + + Tragulina, _45_, 359; + fibula, 519; + manus, 507; + odontoid process, 445; + pes, 523; + skull, 468; + teeth, 429; + ulna, 501 + + Transpalatine, crocodile, 255; + reptiles, 278 + + Trapezium, 27; + dog, 408 + + Trapezoid, 27; + dog, 408 + + Trichechidae, _48_, 369 + + _Trichechus_, _48_; + see Walrus + + Trionychia, _37_; + general characters, 194 + + Trionychidae, _37_ + + _Trionyx_, _37_, 193 f.; + exoskeleton, 214, 270; + skull, 283; + vestiges of teeth, 274 + + _Trissolepis_, _32_; + scales, 104 + + _Tritylodon_, _43_; + teeth, 348 + + Trochanter, dog 412; + duck, 326 + + Trochilidae, _42_; + see Humming-birds + + Trochlea, crocodile, 263; + dog, 405 f.; + duck, 323; + turtle, 232 + + Trogon, _42_; + foot, 342 + + Trogonidae, _42_ + + _Tropidonotus_, _38_; + jaws, 280; + skull, =279= + + Trunk vertebrae, cod, 84; + see thoracic and lumbar vertebrae + + _Trygon_, _32_; + calcification of vertebrae, 114; + caudal spine, 106 + + Trygonidae, _32_ + + Tuberosities of humerus, dog, 405; + of ischium, dog, 411 + + Tunicata, _30_, 51 + + _Tupaia_, skull, 480; + thoraco-lumbar vertebrae, 450 + + Turbinals, dog, 395 + + _Tursiops_, _45_; + skull, 462 + + Turtle, _37_; + anterior limb, 232, =234=; + cranium, 222 f., =226=; + hyoid, 231, =285=; + mandible, 230; + pectoral girdle, 231; + pelvic girdle, 235; + pes, 236; + plastron, 217, =218=; + posterior limb, =234=, 235; + sense capsules, 227; + skull, 222; + vertebral column, 219; + Leathery --, see _Dermochelys_; + Snapping --, see _Trionyx_ + + Tylopoda, _45_, 359; + fibula, 519; + manus, 507; + odontoid process, 445; + pelvis, 514; + pes, 523; + skull, 468; + teeth, 428; + ulna, 502 + + Tympanic, dog, 392; + -- cavity, crocodile, 250; + diagram of mammalian, =391=; + dog, 393; + duck, 315 f.; + turtle, 228; + -- recess, duck, 315 + + Tympano-hyal, dog, 399 + + Typhlopidae, _38_; + scales, 270; + skull, 278 + + _Typhlops_, _38_; + ischio-pubis, 292; + traces of posterior limb, 293 + + Typotheriidae, _46_ + + _Typotherium_, _46_, 358, 361; + clavicle, 495, 497; + femur, 519; + pes, 525; + skull, 472; + teeth, 432 + + + _Udenodon_, _36_, _192_; + beak, 271 + + Uintatheriidae, _47_; + skull, 364 + + _Uintatherium_, _47_; + dental formula, 433; + leg, 519; + limbs and limb girdles, =516=; + manus, 510; + pelvis, 514; + skull, 473 + + Ulna, 26; + crocodile, 265; + dog, 406; + duck, 323; + frog, 164; + newt, 147; + turtle, 233 + + Ulnare, 27; + see Carpus + + Umbilicus, inferior and superior, duck, 303 + + Uncinate process, 190; + crocodile, 259; + duck, 320 + + Unciform, 27, 345, 504; + dog, 408 + + Ungulata, _45_; + auditory ossicles, 487; + caudal vertebrae, 453; + cervical vertebrae, 445; + general characters, 357; + manus, 506; + pectoral girdle, 495; + pes, 522; + ribs, 491; + sacrum, 452; + skull, 464 f.; + sternum, 489; + teeth, 427 f.; + thoraco-lumbar vertebrae, 448 + + Ungulata vera, _45_; + arm bones, 501; + general characters, 358; + manus, 506; + pelvis, 514; + thigh and shin, 519 + + Upper arm, 26; + crocodile, 263; + dog, 405; + duck, 323; + frog, 164; + newt, 147; + turtle, 232 + + _Upupa_, _42_; + see Hoopoe + + Urochordata, _30_, 51 + + Urodela, _35_; + general characters, 134; + pelvis, 187; + ribs, 182; + skull, 174 + + Urohyal, cod, 101; + duck, 320 + + Urostyle, Anura 172; + cod, 85; + frog, 153; + Teleostei, 117 + + Ursidae, _48_; + humerus, 502 + + _Ursus_, _48_; + dental formula, 439; + see Bears + + + Vacuities, anterior palatine --, crocodile, 252, 258; + -- in reptilian skull, 281; + posterior palatine --, crocodile, 254, 257; + pre-orbital --, reptiles, 283 f. + + Vampire, _49_; + teeth, 441 + + Vane, of feather, 303 + + Varanidae, _38_ + + _Varanus_, _38_; + shoulder girdle, =202=; + skull, =201= + + Vasodentine, 108, 272 + + Vertebral column, 14; + Amphibia, 170; + birds, 332; + cod, 83; + crocodile, 239; + dog, 378; + duck, 307; + Elasmobranchs, 113; + frog, 152; + mammals, 442; + newt, 138; + turtle, 219; + -- ribs, crocodile, 259; + dog, 402; + duck, 320; + -- shield, turtle, 214 + + Vertebrata, general characters, 53 + + Vexillum, of feather, 303 + + Vibrissae, dog, 374 + + Viscacha, _47_ + + Visceral skeleton, 21; + dogfish, 77; + Elasmobranchs, 119 f. + + _Viverra_, _48_; + acetabular bone, 515 + + Viverridae, _48_ + + Vomer, 21; + cod, 98; + crocodile, 252; + dog, 395; + duck, 317; + frog, 158; + salmon, 93; + turtle, 229 + + Vomero-palatine, newt, 143 + + _Vultur_, _41_ + + Vulture, _41_; + Black --, shoulder girdle and sternum, =337= + + + Waders, 335 + + Walrus, _48_, 367, 369; + canines, 420; + manus, 511; + pes, 526; + position of limbs, 29; + skull, 480; + teeth, 440 + + Warblers, _42_ + + Wart hog, _45_; + teeth, 428 + + Weasel, _369_ + + Whale, baleen, 3, 418; + Ca'ing --, _45_, + see _Globicephalus_; + Humpbacked --, _44_, 357; + Right --, _44_, 357; + Sperm --, _44_, 357, + see _Physeter_; + True or Whalebone --, 356 + + Whiting, _33_ + + Wild duck, _41_; + see Duck + + Wing, duck, 322; + Gannet, Ostrich, and Penguin, =339= + + Wolf, _48_ + + Wombat, _43_; + atlas, 443; + pes, 521; + sacrum, 451; + skull, =456=; + tail, 453; + teeth, 423 + + Woodpecker, _42_, 335; + foot, 342; + hyoid, 336 + + Wrasse, _33_; + teeth, 111 + + + _Xenacanthus_, _31_; + pectoral fins, 130 + + Xenopidae, _36_ + + _Xenopus_, _36_; + branchial arches, 182; + nails, 168; + pelvis, 188; + ribs, 182 + + Xiphiplastron, turtle, 217 + + Xiphisternal horn, crocodile, 260 + + Xiphisternum, dog, 404; + frog, 163 + + Xiphoid process, duck, 321 + + + _Zeuglodon_, _44_, 353, 356; + dental formula, 426; + dermal plates, 420 + + Zeuglodontidae, _44_ + + Zygantra, defined, 199 n.; + reptiles, 276 + + Zygapophyses, cod, 84; + crocodile, 240 f.; + dog, 379 f.; + duck, 308 f.; + frog, 152 f.; + newt, 139; + turtle, 219 f. + + Zygosphene, defined, 199 n.; + reptiles, 276 + + +CAMBRIDGE: PRINTED BY J. AND C.F. CLAY, AT THE UNIVERSITY PRESS. + + + + +CAMBRIDGE BIOLOGICAL SERIES. + + +General Editor, A.E. SHIPLEY, M.A., F.R.S., Fellow and Tutor of +Christ's College. + +=A Text-Book of Zoogeography.= By FRANK E. BEDDARD, M.A., F.R.S., +Prosector of the Zoological Society of London. With 5 Maps. Crown 8vo. +6_s._ + +=The Elements of Botany.= By FRANCIS DARWIN, M.A., M.B., F.R.S., +Fellow of Christ's College. With 94 Illustrations. Crown 8vo. _Second +Edition._ 4_s._ 6_d._ + + _Journal of Education._ A noteworthy addition to our botanical + literature. + +=Practical Physiology of Plants.= By FRANCIS DARWIN, M.A., F.R.S., and +E. HAMILTON ACTON, M.A. Crown 8vo. With 45 Illustrations. _Second +Edition._ 4_s._ 6_d._ + + _Nature._ The authors are much to be congratulated on their + work, which fills a serious gap in the botanical literature of + this country. + +=Lectures on the History of Physiology= during the Sixteenth, +Seventeenth and Eighteenth Centuries. By Sir M. FOSTER, K.C.B., M.P., +M.D., D.C.L., F.R.S., Fellow of Trinity College. Demy 8vo. With a +Frontispiece. 9_s._ + + _Nature._ There is no more fascinating chapter in the history + of science than that which deals with physiology, but a concise + and at the same time compendious account of the early history + of the subject has never before been presented to the English + reader. Physiologists therefore owe a debt of gratitude to Sir + Michael Foster for supplying a want which was widely felt.... No + higher praise can be given to the book than to say that it is + worthy of the reputation of its author.... It is by no means an + easy task to do adequate justice to the mine of literary and + historic research which the author has laid open to view. + + _Guardian._ We must urge not merely students of physiology, but + all those interested in the history of modern thought, to buy + and read the book.... The present volume has a charm and + fascination which is too often wanting in historical studies. + His portraits live, and live in a live world; they are not mere + dead pictures set in a stereotyped, historical framework. + + _Spectator._ We can recommend this admirable and suggestive + book with confidence to all, laymen or doctors, who wish to + trace the gradual growth of man's knowledge of the physical + basis of his life. + +=The Soluble Ferments and Fermentation.= By J. 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With numerous illustrations. 10_s._ 6_d._ _net_. + +=The Vertebrate Skeleton.= By SIDNEY H. REYNOLDS, M.A. Crown 8vo. With +110 Illustrations. 12_s._ 6_d._ + +=Manual of Practical Morbid Anatomy=, being a Handbook for the +Post-mortem Room. By H.D. ROLLESTON, M.A., M.D., F.R.C.P., and A.A. +KANTHACK, M.D., M.R.C.P. Crown 8vo. 6_s._ + +=Fossil Plants: for students of Botany and Geology.= By A.C. SEWARD, +M.A., F.R.S., Fellow of Emmanuel College. In 2 vols. Demy 8vo. Vol. I. +12_s._ [Vol. II. _In the Press._] + + _Revue Scientifique._ Nous ne pouvons entrer dans le détail; + mais il est évident que M. Seward, praticien distingué + lui-même, est très au courant des travaux des autres, il les + cite et utilise abondamment; et ceci est fait pour inspirer + confiance. Au total, son oeuvre est appuyée sur des bases + solides, et elle restera sans doute longtemps le bréviaire, le + manuel de ceux qui veulent, non pas seulement s'initier à la + paléobotanique, mais retrouver les renseignements qui sont + épars dans des centaines de monographies qu'on a souvent peine + à se procurer. Le livre de M. Seward fait partie des _Cambridge + Natural Science Manuals_, et il est digne de cette collection, + qui est elle-même digne du foyer scientifique universellement + réputé, où il a vu le jour. + + _Guardian._ We have already alluded to the spirit of caution + which characterises the book, and we may add that it promises + to be most helpful to the botanist who would extend his + researches into the past. + + _Natural Science._ A most successful treatment of a difficult + subject. All of importance is brought forward and impartially + discussed.... Such a book has long been a desideratum. Mr + Seward's style is clear and concise, and the many pitfalls into + which beginners are apt to stumble are clearly pointed out. + + +=Zoology. An Elementary Text-Book=. By A.E. SHIPLEY, M.A., F.R.S., and +E.W. MACBRIDE, M.A. (Cantab.), D.Sc. (London), Professor of Zoology in +McGill University, Montreal. Demy 8vo. With numerous Illustrations. +10_s._ 6_d._ _Net_. + + _Pilot._ A very business-like and convenient manual of modern + Zoology. + + _School World._ As a thoroughly trustworthy and instructive + text-book for serious students, the work can be strongly + recommended. Its value is enhanced by the large number of + excellent illustrations, many of which are delightfully fresh. + + _Oxford Magazine._ It is readable, well arranged, well printed, + copiously and admirably illustrated, and it covers the whole + field of zoology. + + _Nature._ There pervades the pages of the work a freshness of + style and unconventionality which render them pleasant reading + and attractive; while, in the frequent allusion to the + commonest occurrences of daily life and human affairs, the + interest of the reader is assured. + + _Pall Mall Gazette._ Precisely the sort of book which, if it + came into a thoughtful boy's hands, would turn him from a + smatterer into a student.... One of the most instructive and + attractive books that could be put into the hands of a young + naturalist. + +=Grasses=: a Handbook for use in the Field and Laboratory. By H. +MARSHALL WARD, Sc.D., F.R.S., Fellow of Sidney Sussex College, +Professor of Botany in the University of Cambridge. With 81 figures. +Crown 8vo. 6_s._ + + _Pilot._ Brimful of matter. + + _Field._ The work is essentially suited to the requirements of + those desirous of studying the grasses commonly grown in this + country, and it can fairly be said that it furnishes an amount + of information seldom obtained in more pretentious volumes. + + _Athenaeum._ Botanists and Agriculturists alike have reason to + thank Prof. Ward for this very serviceable addition to the + literature of grasses. + +=Trees=: A Handbook of Forest Botany for the Woodlands and the +Laboratory. By H. MARSHALL WARD, Sc.D., F.R.S., Fellow of Sidney +Sussex College, Honorary Fellow of Christ's College and Professor of +Botany in the University of Cambridge. In six volumes. 1. Buds and +Twigs, 2. Leaves, 3. Inflorescences and Flowers, 4. Fruits and Seeds, +5. Seedlings, 6. General Characters. Vol. I. Buds and Twigs. Crown +8vo. Illustrated. 4_s._ 6_d._ _net_. + +=A Treatise on the British Freshwater Algae.= By G.S. WEST, M.A., +A.R.C.S., F.L.S., Professor of Natural History at the Royal +Agricultural College, Cirencester. Demy 8vo. 10_s._ 6_d._ _net_. + +=A Manual and Dictionary of the Flowering Plants and Ferns=. By J.C. +WILLIS, M.A., Director of the Royal Botanic Gardens, Ceylon. _Second +Edition._ Complete in one volume. Crown 8vo. 10_s._ 6_d._ + +=Elementary Palaeontology--Invertebrate.= By HENRY WOODS, M.A., F.G.S., +University Lecturer in Palaeozoology. Crown 8vo. _Third Edition._ +Revised and enlarged, with 112 Illustrations. 6_s._ + +=Outlines of Vertebrate Palaeontology for students of Zoology.= By +ARTHUR SMITH WOODWARD, M.A., F.R.S., Keeper of the Department of +Geology in the British Museum. Demy 8vo. With numerous Illustrations. +14_s._ + + _Athenaeum._ The author is to be congratulated on having produced + a work of exceptional value, dealing with a difficult subject in a + thoroughly sound manner. + + +_In preparation._ + +=Morphology and Anthropology.= By W.L.H. DUCKWORTH, M.A., Fellow and +Lecturer of Jesus College, University Lecturer in Physical Anthropology. + +=The Origin and Influence of the Thorough-bred Horse.= By W. RIDGEWAY, +M.A., Disney Professor of Archaeology and Fellow of Gonville and Caius +College. With numerous Illustrations. Demy 8vo. + +=The Morphology of Plants.= By J.C. WILLIS, M.A. + + =London=: C.J. CLAY AND SONS, + CAMBRIDGE UNIVERSITY PRESS WAREHOUSE, + AVE MARIA LANE, + AND + H.K. LEWIS, 136, GOWER STREET, W.C. + + =Glasgow=: 50, WELLINGTON STREET + + * * * * * + + + Transcriber Notes + + Italic text is denoted by _underscores_ and bold text by =equal + signs=. Subscripts are represented using braces, e.g. V{1}, and + superscripts are introduced with a caret, e.g. 2^e + + Obvious punctuation and spelling errors, and inconsistent hyphenation + have been corrected. + + The oe ligature in the text has been replaced with the characters oe. + + + + + +End of Project Gutenberg's The Vertebrate Skeleton, by Sidney H. Reynolds + +*** END OF THE PROJECT GUTENBERG EBOOK 43431 *** |
