CORNELL UNIVERSITY. THE Goswell P. Flower Library THE GIFT OF ROSWELL P. FLOWER FOR THE USE OF THE N. Y. STATE VETERINARY COLLEGE. 1897 ornell University Library An introduction to the study of mammals PRINTED |IN U.S.A Cornell University The original of this book is in the Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924001022684 AN INTRODUCTION TO THE STUDY OF MAMMALS AN INTRODUCTION TO THE STUDY OF MAMMAL LIVING AND EXTINCT BY WILLIAM HENRY FLOWER C.B., F.R.S., D.C.L., LL.D., P.Z.S8., F.LS., F.GS., &e. DIRECTOR OF THE NATURAL HISTORY DEPARTMENTS, BRITISH MUSEUM AND RICHARD LYDEKKER B.A., ¥.G.S§,,, F-2:8.,, &c. THE WOOLLY OPOSSUM LONDON: ADAM AND CHARLES BLACK MDCCCXCI PREFACE ONE of the greatest difficulties experienced by all who undertake a work of this nature, not professing to be an exhaustive treatise on the subject with which it deals, is to determine the amount of detail desirable to be introduced to meet the requirements of the ordinary student, without rendering it too bulky or costly for general use. The experience of those who endeavour to profit by the book can alone decide how far the authors have succeeded in this respect. It will be observed that in many instances certain better-known or more interesting members of the class have been described at considerable length, while it has been necessary to treat others with much greater brevity. . . With regard to the references to the literature of the various groups treated of, it has been the endeavour of the authors to make a selection of such memoirs and works as are likely to prove most valuable to the student for the amount of original informa- tion which they contain, and more especially of those giving full bibliographical data up to the time of their publication, the repetition of which has been considered unnecessary. In a few instances new generic terms have been introduced to vi PREFACE replace some which were already occupied ; these have been pro- posed by Mr. Lydekker, and should be quoted as‘his. The work is based largely upon the article “Mammalia,” to- gether with forty shorter articles, written by the senior of the two authors for the ninth edition of the Encyclopedia Britannica. The account of the orders Rodentia, Insectivora, and Chiroptera con- tributed to the article “Mammalia” by Dr. G. E. Dobson, F.R.S., as well as the articles “Mole,” “Shrew,” and “ Vampyre,” by the same writer, the articles “ Marmot,” “Mouse,” “Opossum,” “ Phal- anger,” “Rat,” “Squirrel,” “Stoat,” “Vole,” and others, by Mr. Oldfield Thomas, and likewise the article “Ape,” by Dr. St. G. Mivart, F.R.S., have also been made use of to a greater or less extent. The best thanks of the authors are due to these three gentlemen for freely permitting the incorporation of their own work in the present volume. Mr. Lydekker undertook the task of arranging the various articles in their proper sequence, selecting from these such portions as seemed suitable, fillmg up the gaps, and adding new matter where necessary ; a large amount of this new matter treating of the extinct forms, and also of the group Artiodactyla. The subsequent revision, both before being sent to the printers, and also when passing through the press, has been made by both authors, who are thus jointly responsible for the whole work. The illustrations are to a great extent those prepared for the various articles in the Encyclopaedia, but many have been added —some drawn expressly for the work, and some borrowed from other publications. For most of the latter the authors take this opportunity of expressing their thanks to the Publication Com- PREFACE i mittee of the Zoological Society of London, as well as to the individual writers.in whose works they first appeared. The authors have further much pleasure in acknowledging the ready and obliging way in which Mr. Oldfield Thomas has, throughout the progress of the work, placed his extensive know- ledge of the group of animals of which it treats at their disposal. Lonvon, Jfarch 1891. CoRRIGENDA. Page 280, for Cheropsis read Cheeropsis. Page 292, for Cheropotamide and Cheropotamus read Cheeropotamide and Chceropotamus. Page 590, for Piecilogale read Precilogale. CONTENTS CHAPTER I IntRoDUCTORY REMARKS Use of term mammals, 1; Characters of mammals, 2; De- velopment of young, 3; Size of mammals, 4; Uses and products of mammals, 4 CHAPTER II GENERAL ANATOMICAL CHARACTERS I. II. III. Iv. Tegumentary Structures Hair, 7; Colour, 8; Scales, etc., 11; Nails, claws, and hoofs, 12; Odour-secreting glands, 12. Dental System. Teeth, 13 ; Structure of ‘eta 13 ; Darcie aaat of uth, 15 ; Forms of ‘teeth, 17 ; Succession of teeth, 19; Arrangement, homologies, and notation of teeth, 21; Dewtal formule, 25 ; Modifications of teeth in relation to function, 28; Taxonomy, 30; Trituberculism, 30. The Skeleton . Definition, 33; Axial ‘Seeion, 34 ; skull, 34; Poul column, 39; Cerviewl vertebree, 41; Dorsal vertebns, 42; Lumbar yankee, 42; Sacral westebnx, 43; Caudal vertebra, 43; Sternum, 44; Ribs, 44; Appendicular skeleton, 46; Anterior limb, 46; Shoulder-girdle, 46 ; Brachium and Ante- brachium, 47 ; Manus, 48; Carpus, 48; Metacarpus and Phal- anges, 49; Posterior limb, 50; Pelvic aide 50; Thigh and eg, 51; Poe, 52. The Digestive System . General considerations, 53; Mouth, 54; Salivary glands, 55; Stomach, 57 ; Intestinal canal, 59; Liver, 60. . Circulatory, Absorbent, Respiratory, and Urinary Systems Blood, 63; Heart, 63; Lymphatic vessels, 65; Ductless glands, 65 ; Nostrils, 66; Trachea, 67; Larynx, 67; Diaphragm, 67 ; Lungs, 68 ; Air-sacs, 68; Urinary Organs, 69; Bladder, 69. PAGE 13 33 53 63 x CONTENTS VI. Nervous System and Organs of Sense Brain, 69; Nerves, 71; Sense of touch, 72; Taste and smell, 72; Sight, 72; Hearing, 73. / VII. Reproductive Organs Testes, 74; Penis, 74; Ovaries and oviduct, 75 ; Mammary glands, 75 ; Secondary sexual characters, 76 ; Placenta, 76. CHAPTER III ORIGIN AND CLASSIFICATION OF THE MAMMALIA Origin, 82; Classification, 84; Table of orders and families, 88. CHAPTER IV GEOGRAPHICAL AND GEOLOGICAL DISTRIBUTION I. Geographical Distribution : F . Zoological regions, 96 ; Palearctic region, 97; Ethiopian region, 98; Oriental region, 100; Celebes, 102; Nearctic region, 102; Neotropical region, 108; Aquatic mammals, 104. II. Geological Distribution . : Sequence of strata, 107; Mesozoic mammals, 109; Multi- tuberculata, 109 ; Polyprotodont types, 113; Tertiary mammals, 115. CHAPTER V THE Supciass PROTOTHERIA OR ORNITHODELPHIA General characters, 117. Family ORNITHORHYNCHIDS, 119; Ornithorhynchus, 119. Family Ecuipnips, 124; Echidna, 125 ; Proechidna, 126 ; Fossil species, 127. CHAPTER VI Tue Svuscuass METaTHERIA OR DIDELPHIA General characters, 128; Distribution, 131; Classification, 181. Suborder POLYPROTODONTIA A : ; : Family DIDELPHYID&, 133; Chironectes, 1384; Didelphys, 135. Family DasyurIDE, 136 ; Subfamily Dasyurine, 186 ; Thylacinus, 136 ; Sarcophilus, 1387 ; Dasyurus, 188 ; Phascolo- gale, 139 ; Sminthopsis, 139; Antechinomys, 1389; Subfamily Myrmecobiine, 140; Myrmecobius, 140. Family PERAMELIDS, 141; Perameles, 142; Peragale, 148; Chewropus, 148. PAGE 69 74 82 93 93 107 128 133 CONTENTS xi Suborder DIPROTODONTIA Family PHAscoLoMYID, 144; Phascolomys, 145; Phascol- onus, 146. Family PHALANGERIDA, 147; Subfamily Tarsipedine, 148; Tarsipes, 148; Subfamily Phalangerine, 149; Phalanger, 149; Trichoswrus, 150; Pseudochirus, 151; Petauroides, 152 ; Dactylopsila, 152; Petuurus, 158; Gymmnobelideus, 154; Dromicia, 154; Distceechurus, 155 ; Acrobates, 155 ; Subfamily Phascolaretine, 155; Phascolarctus, 156. Extinct Puat- ANGEROIDS, 157; Thylacoleo, 157. Family Macropopipa, 158 ; Subfamily Hypsiprymnodontine, 162; Hypsiprymnodon, 162; T'riclis, 162; Subfamily Potoroine, 162; Potorous, 163 ; Bettongia, 163 ; Caloprymnus, 164; pyprymnus, 164; Sudb- family Macropodine, 164; Lagostrophus, 165; Dendrolagus, 165; Dorcopsis, 166; Lagorchestes, 166; Onychogale, 166 ; Petrogale, 167 ; Macropus, 167 ; Extinct genera, 170. Extincr Famittss, 171; Diprotodon, 171; Nototherium, 171. CHAPTER VII THE Susciass EvTHERIA AND THE ORDER EDENTATA . General characters and classification of Eutheria, 173. OrpER EpENTATA : : F : Family Brapypopips, 179; Bradypus, 181; Cholepus, 182; Nothropus, 183. Family MErcATHERiIIDz, 183; Mega- theriwm, 185; Scelidotherium and Mylodon, 188; Promega- therium, 189. Family MyRMECOPHAGIDA, 190; Myrmecophaga, 190; Tamandua, 192; Cycloturus, 198. Family DasyPoDIDA, 194; Subfamily Chlamydophorine, 196 ; Chlamydophorus, 196 ; Subfamily Dasypodine, 197; Dasypus, 197; Xenwrus, 198 ; Priodon, 198; Tolypeutes, 199; Subfamily Tatusiine, 200 ; Tatusia, 200; Extinct genera, 201. Family GLYPTODONTIDA, 202. Family Manipm, 204; Manis, 204; Palcomanis, 208. Family ORYCTEROPODIDE, 208; Orycteropus, 208. Biblio- graphy, 211. CHAPTER VIII THE ORDERS SIRENIA AND CETACEA ORDER SIRENIA : : ; : : Family MANatTipe, 215; Manatus, 215. Family Hatrt- CORIDZ, 220; Halicore, 220. Family Ruyrinipm, 221; LRhytina, 221. Exrincr SIRENIANS, 222; Halitheriwm, 222 ; Other forms, 223. Bibliography, 224. ORDER CETACEA Suborder MysTacoceti . : : : ‘ : Family BALENIDA, 234; Balena, 236; Neobalena, 241; Rhachianectes, 241; Megaptera, 241; Balenoptera, 242; Extinct genera, 245, 176 212 212 225 234 xii CONTENTS Suborder ARCHHOCETI Family ZEUGLODONTIDA, 246 ; Zeuglodon, 246. Suborder ODONTOCETI Family PHYSETERIDS, 247; Subfamily Physeterine, 248 ; | Physeter, 248; Cogia, 250; Extinct physeteroids, 251; Swub- JSamily Ziphiine, 251; Hyperoédon, 252; Ziphius, 254; Meso- plodon, 254; Berardius, 256; Choneziphius, 257. Family SQuaLODONTIDS, 257; Sgwalodon, 257. Family PLATANISTIDS, 257 ; Platanista,258 ; Inia, 259; Pontoporia, 259; Fossil forms, 259. Family DELPHINIDA, 260; Monodon, 260; Delphinapterus, 262; Phoceena, 263 ; Cephalorhynchus, 266 ; Orcella, 267; Orca, 267; Pseudorea, 268; Globicephalus, 268; Grampus, 270; Feresia, 270 ; Lagenorhynchus, 270; Delphinus, 271 ; Tursiops, 271; Prodelphinus, 271; Steno, 271; Sotalia, 272. Biblio- graphy, 272. CHAPTER Ix THE OrpER UNGULATA Uneunata VERA Suborder ARTIODACTYLA Suina, 278. Family HippoporaMipas, 278 ; Hippopotamus, 278. Family Surpm, 281; Sus, 281; Babirwsa, 287 ; Phaco- cherus, 288. Family Dicoryitipm, 289; Dicotyles, 289; Hyotheriwm, ete.,291. EXTINcT TRANSITIONAL ARTIODACTYLES, 292; Cheropotamide, 292; Anthracotheriide, 292; Meryco- potamus, 293 ; Cotylopide, 293 ; Anoplotheriide, 293 ; Ceno- theriide, 294; Dichodontide, 294. Tynopopa, 295. Family CAMELIDm, 295; Camelus, 296; Auchenia, 298; Extinct Cameloids, 303. TracuLina, 305. Family TRAGULIDA, 305; Tragulus, 305 ; Dorcatherium, 306 ; Extinct Traguloids, 306. PrEcora, 307; Antlers, 308; Horns, 310; Teeth, 310; Stomach, 312. Family CERvipm, 313; Subfamily Moschine, 314; Moschus, 314; Subfamily Cervine, 316; Plesiometacarpalia, 316; Cervulus, 316 ; Elaphodus, 318 ; Cervus, 819; Telemeta- carpalia, 323; Rangifer, 324; Alces, 326; Cervalces, 327 ; Capreolus, 327 ; Hydropotes, 328 ; Cariacus, 329 ; Pudua, 330 ; Extinct genera, 330. Family GIRAFFIDZ, 330; Giraffa, 331; Allied extinct types, 332. Family ANTILOCAPRIDS, 333 ; Antilocapra, 333. Family Bovipm, 334; Alcelaphus, 334; Connocheetes, 336; Cephalophus, 338; Tetraceros, 338; Neo- tragus, 338 ; Nanotragus, 339 ; Pelea, 389 ; Cobus, 339; Cervi- capra, 840; Antilope, 340; pyceros, 341; Saiga, 341; Pantholops, 341; Gazella, 341; Hippotragus, 343 ; Oryx, 348 ; Addax, 345; Boselaphus, 345; Tragelaphus, 346; Strepsiceros, 347 ; Oreas, 348; Extinct types, 348; Rupicapra, 349; Nemo- rhedus, 350; Haploceros, 351; Budorcas, 351; Capra, 352 ; Ovis, 354; Ovibos, 357; Bos, 360. PAGE 246 Q47 273 275 275 CONTENTS xiii Suborder PERISSODACTYLA Family Tarrripa, 370; Tapirus, 370; Paleotapirus, 373. Family LOPHIODONTIDA, 373. Family PALHOTHERIIDA, 375. Family Equipe, 376; Protohippus, 380; Hipparion, 380; Equus, 381. Family RHINOCEROTIDA, 402; Rhinoceros, 402; Extinct types, 411. Families LAMBDOTHERIID#, CHALICO- THERIID#, and TITANOTHERIIDS, 412. Family Macrav- CHENIIDA, 414. Family PROTEROTHERIIDA, 414. SUBUNGULATA Suborder HYRACOIDEA : Family Hynacips, 415 ; Hyrax, 417 ; een 418. Suborder PROBOSCIDEA Family ELEPHANTIDE, 423 ; Elephas, 424 ; ans 431. Family DINOTHERIIDE, 435 ; Dinotharint, 435. Suborder AMBLYPODA Vintatherium, 436 ; Bioelpinarn, 437. Suborder CONDYLARTHRA Suborder ToxopontTIa : ‘ : Nesodon, 439 ; Toxodon, 439 ; Typotherium, 440. Group TILLODONTIA Bibliography of Ungulates CHAPTER X Tae ORDER RODENTIA Suborder SImPLICIDENTATA Section SCIUROMORPHA, 448. Family ANOMALURIDA, 449 ; Anomalurus, 449. Family Scruripz, 450; Scturus, 450 ; Rhithrosciurus, 452; Xerus, 452; Tamias, 452; Pteromys and Scturopterus, 453; Hupetaurus, 454; Extinct genera, 454; Arctomys, 454; Cynomys, 455; Spermophilus, 456; Extinct genera, 457. Family HapLopontips, 457; Haplodon, 457. Family Castorips, 457; Castor, 457. Section MyomorrHa, 459. Family Myoxipm, 459; Myoxrus, 459; Eliomys, 459 ; Graphiurus, 459 ; Claviglis, 460; Muscardinus, 460. Family LopHiomyID#&, 460; Lophiomys, 460. Family Muripa, 461 ; Hydromys, 461; Xeromys, 461; Platacanthomys, 462; Gerbillus, 462; Pachyuromys, 462; Mystromys, 462; Otomys and Dasymys, 462; Malacomys, 462; Phiewomys, 462; Dendromys, 463 ; Cricetus, 463; Holochilus, 464; Sigmodon, 464; Rhithrodon and Ochetodon, 464; Neotoma, 464; Hypogeomys, 465 ; Nesomys, 465; Brachytarsomys, 465; Hallomys, 465; Eliwrus, 465 ; Phenacomys, 466; Arvicola, 466; Synaptomys, 467 ; Myodes, 467; Cuniculus, 470; Fiber, 470; Neofiber, 472; LEllobius, PAGE 368 ‘414 415 418 436 438 439 441 442 443 448 xiv CONTENTS PAGE 472; Siphneus, 472; Deomys, 473; Mus, 473; Nesocia, 475; Golunda, 476; Uvromys, 476; Chiruromys, 476; Hapalotis, 476 ; Mastacomys, 476; Acanthomys, 476; Echinothria, 477 ; Typhiomys, 477 ; Cricetomys and Saccostomus, 477 ; Pithechirus, 477. Family Spatactpa, 477; Spalax, 477 ; Rhizomys, 477 ; Bathyergus, 478; Georychus and Myoscalops, 478; Hetero- cephalus, 478. Family Gromyipm, 478; Geomys, 478; Thomomys, 478 ; Dipodomys, 479 ; Perognathus and Heteromys, 479. Family Divopipm, 479; Sminthus, 479; Zapus, 480; Dipus, 480; Alactaga, 480; Platycercomys, 480 ; Pedetes, 480. Section Hystricomorpua, 480. Family OcTopontTips, 480. Ctenodactylus, 481 ; Pectinator, 481 ; Octodon, 481 ; Habrocoma, 482; Schizodon, 482; Ctenomys, 482; Spalacopus, 482 ; Petromys, 482 ; Myopotamus, 482 ; Capromys, 482; Aulacodus, 483 ; Plagiodon, 483 ; Loncheres and Echinomys, 483 ; Mesomys, 483 ; Dactylomys, 483; Cercomys, 483; Carterodon, 484; Fossil forms, 484. Family THERIDoMYIDmH, 484. Family Hystricipa, 484 ; Erethizon, 484 ; Synetheres, 485 ; Chetomys, 486 ; Hystriz, 486; Atherura, 487; Trichys, 487. Family CHINCHILLIDA, 487 ; Chinchilla, 487 ; Lagidiwm and Lagosto- mus, 488; Extinct genera, 488. Family Castororpipa, 488 ; Castoroides, 488. Family DasyprocTips, 488; Dasyprocta, 488 ; Ceelogenys, 489. Family Dixomytpa, 489; Dinomys, 489. Family Cavipx, 489; Cavia, 489; Dolichotis, 490; Hydro- cherus, 490 ; Extinct genera, 491. Suborder DUPLICIDENTATA ‘ é : 491 Family Lacomyipm, 491; Lagomys, 491. Family Lrpo- RIDE, 492; Lepus, 492. CHAPTER XI THE ORDER CARNIVORA : F : : : 496 Suborder CARNIVORA VERA , 497 Section AALUROIDEA, 501. Family Fetipm, 502; Felis, 502 ; Cyncelurus, 523; Extinct genera, 523. Family Viver- RIDE, 525; Cryptoprocta, 525; Viverra, 526; Fossa, 527; Genetta, 528; Prionodon, 530; Poiana, 531; Paradoxurus, 532 ; Arctogale, 583; Hemigale, 533 ; Arctictis, 584; Nandinia, 534; Cynogale, 534; Herpestes, 585 ; Helogale, 537 ; Bdeogale, 537 ; Cyntetis, 537 ; Rhinogale, 537 ; Crossarchus, 5387 ; Suricata, 538; Galidictis, Galidea, and Hemigalidea, 5388; Eupleres, 538; Extinct genera, 5389. Family PRoreLnrpa, 539 ; Proteles, 539. Family Hymnipa, 540; Hyena,540. Section CyNorpDEA, 544. Family CAnipm, 5443; Canis, 546; Lycaon, 553; Icticyon, 553; Otocyon, 554; Extinct genera, 555. Section ARCTOIDEA, 556. Family Urstpa, 557; Ursus, 557 ; Ale7ursus, 560 ; Aluropus, 560; Extinct genera, 561. amily Procyonrpse, 562; Elurus, 562 ; Procyon, 564 ; Bassaris, 566 ; Bassaricyon, 566 ; Nasua, 566; Cercoleptes, 567. Family MusTEuips, 567 ; CONTENTS xv PAGE Lutra, 567 ; Extinct Otters, 570; Latax, 570; Mephitis, 572 ; Conepatus, 574; Arctonyx, 574; Mydaus, 575; Meles, 575 ; Tasxidea, 576; Mellivora, 576; Helictis, 578; Ictonyx, 579; Galictis, 579 ; Mustela, 579; Extinct Mustelines, 590 ; Pacilo- gale, 590 ; Lyncodon, 590; Gulo, 591. Suborder PINNIPEDIA . P 4 : : é 592 Family Oranipsz, 593; Otaria, 593. Family TrRicuE- CHIDE, 596; Trichechus, 597. Family PHoctpx, 600; Halicherus, 601; Phoca, 601; Monachus, 604; Ogmorhinus, 605 ; Lobodon, 605; Pacilophoca, 605; Ommatophoca, 605 ; Cystophora, 605; Macrorhinus, 606 ; Extinct seals, 606 Suborder CREODONTA 606 Hyenodontide, 608 ; es 608 ; Apt sats on Mesonychide, 609. CHAPTER XII THE ORDER INSECTIVORA ‘ ‘ . : ‘ 610 Suborder DERMOPTERA . : & : . 614 Family Giaanoatuawetom: 614; Galeopithecus, 614. Suborder INSECTIVORA VERA. 4 : 616 Family Tupattps, 617 ; Tupaia, 617; Secs 618 ; Ex- tinct genera, 618. Family MAcROSCELIDIDA, 618 ; Iacroscel- ides, 618; Rhynchocyon, 618. Family ErinacEipa, 619; Gymnura, 619 ; Hrinaceus, 620; Extinct genera, 621; Fumily Soricip#&, 621; Sorex, 622; Soriculus, 624; Notiosorex, 624 ; Blarina, 624 ; Crossopus, 625 ; Myosorex, 625 ; Crocidura, 626 ; Diplomesodon, 626; Anurosorex, 626; Chimarrogale, 626 ; Necto- gale, 627; Fossil Soricide, 627. Family Tauripm, 628; Myogale, 628 ; Urotrichus, 629 ; Uropsilus, 629 ; Scalops, 630 ; Scapanus, 630; Condylura, 630; Scaptonyx, 630; Talpa, 680 ; Extinct genera, 634. Family ADAPISORICIDE, 634. Family PoTAMOGALIDA, 634; Potamogale, 635; Geogale, 635. Family SOLENODONTIDH, 635; Solenodon, 636; Centetes, 637 ; Hemi- centetes, 637 ; Ericulus, 688 ; Microgale, 688 ; Oryzorictes, 638 ; Chrysochioris, 689. Exrincr typrs, 640. Bibliography, 640. CHAPTER XIII THE ORDER CHIROPTERA : : ‘ ‘ - 641 Suborder MEGACHIROPTERA : ea 2 650 Family Prernopopips#, 650 ; ree us, 650 ; High 651; Xantharpyia, 652 ; Binet: 653; Cynopterus, 653 ; Harpyia, 653; Cephalotes, 653; Pleralopex, 654; Notopteris, 654 ; Rongeurs, 654 ; Coipanyelerts, and Afelonycterts, 654 ; Nawengicters 655 ; Calltneaycterts, 655 ; Trygenycteris, 655. Xvi CONTENTS Suborder MIcROCHIROPTERA Section VESPERTILIONINA, 655. Family RHINOLOPHID&, 656; Rhinolophus, 656; Hipposiderus, 657; Anthops, 657 ; Rhinonycteris and Tricnops, 658; Coelops, 658; Megaderma, 658. Family VESPERTILIONIDS, 660 ; Plecotus, 660 ; Synotus, 661; Otonycteris, 661; Nyctophilus, 661; Antrozous, 661 ; Vesperugo, 661 ; Chalinolobus, 662 ; Scotophilus, 662; Nyctice- jus, 663; Atalapha, 663; Harpyiocephalus, 663; Vespertilio, 663 ; Cerivoula, 664 ; Natalus, 664 ; Miniopterus, 664 ; Thyrop- tera, 665; Myxopoda, 665; Fossil Vespertilionide, 665. Section EMBALLONURINA, 666. Family EMBALLONURIDA, 666 ; Furipterus and Amorphochilus,666; Emballonura, 667; Coléwra, 667 ; Rhynchonycteris, 667 ; Saccopteryx, 667 ; Taphozous, 667 ; Diclidurus, 668 ; Noctilio, 668 ; Rhinopoma, 669 ; Chiromeles, 669 ; Molossus, 670 ; Nyctinomus, 670 ; Mystacops, 671. Family PHYLLOSTOMATIDH, 672; Chilonycteris, 672; Mormops, 672; Lonchorhina, Otopterus, and Dolichophyllum, 673 ; Vampyrus, ete., 673; Desmodus, 677 ; Diphylia, 678, CHAPTER XIV THE ORDER PRIMATES . | Suborder LEMUROIDEA Family Lemuripz, 683; Indris, 684; Propithecus, 684 ; Avahis, 686; Lemur, 687; Hapalemur, 689; Lepidolemur, 689 ; Chirogaleus, 689; Galago, 690; Nycticebus, 691; Loris, 692; Perodicticus, 693. Family Tarsiipm, 694; Tarsius, 694. Family CHIROMYIDE, 694; Chiromys, 695. EXTINCT LEMUROIDS, 696. Suborder ANTHROPOIDEA Family HaPavipeg, 709; Hapale, 710; Midas, 710. Family CEBIDmH, 711; MMycetes, 711; Pithecta, 712; Uacaria, 712; Callithrix, 713 ; Chrysothrix, 714; Nyctipithecus, 714; Ateles, 715; Eriodes, 715; Lagothriz, 716; Cebus, 717. Family CERCOPITHECID, 718 ; Cynocephalus, 719 ; Theropithecus, 722 ; Cynopithecus, 722; Macacus, 722; Cercocebus, 723; Cerco- pithecus, 724; Nasalis, 725; Semnopithecus, 726; Colobus, 727; Extinct genera, 727. Family Simtpz, 728 ; Hylobates, 728; Simia, 731; Gorilla, 734; Anthropopithecus, 736. Family HomInipa, 739; Homo, 740. Classification of the varieties of Man, 748. PAGE 655 680 682 699 AN INTRODUCTION TO THE STUDY OF MAMMALS LIVING AND EXTINCT CHAPTER I INTRODUCTORY REMARKS Mamata (French, Mammiferes; German, Séiugethierc) is the name invented by Linnzus (from the Latin mamma), and now commonly used by zoologists, for one of the five great classes of vertebrated animals, which, though the best known and undoubtedly the most important group of the animal kingdom, has never received any generally accepted vernacular designation in our language. The unity of structure of the animals composing this class, and their definite demarcation from other vertebrates, were not recognised until comparatively modern times, and hence no word was thought of to designate what zoologists now term a mammal. The nearest equivalents in common use are “beast” and ‘quadruped,” both of which, however, cover a different ground, since they are often used to include the larger four-footed reptiles, and to exclude certain un- doubted mammals, as Man, Bats, and Whales. The limits of the class as now understood by zoologists are perfectly well defined, and, although certain forms still existing on the earth (but not those mentioned above as excluded by the popular idea) are of exceedingly aberrant structure, and exhibit several well- marked characters connecting them with the lower vertebrated groups, common consent retains them in the class with which the great proportion of their characters ally them, and hitherto no traces of any species showing still more divergent or transitional characters have been discovered. There is thus an interval, not bridged over by any known forms, between mammals and other 1 2 INTRODUCTORY vertebrates ; although recent discoveries have shown evidence of a more or less marked affinity between the most generalised mammals and a peculiar group of extinct reptiles known as the Anomodontia (or Theromora), which are themselves nearly related to the equally extinct Labyrinthodont amphibians of the Paleozoic and Mesozoic epochs. In the gradual order of evolution of living beings, mammals, taken altogether, are certainly the highest in organisation, as, with the possible exception of birds, they were the last to appear on the earth’s surface. But, as in speaking of all other large and greatly differentiated groups, this expression must not be understood in too limited a sense. The tendency to gradual perfection for their particular station in life, which all groups manifest, leads to various lines of specialisation, or divergence from the common or general type, which may or may not take the direction of elevation. A too complex and sensitive condition of organisation may in some circumstances of life be disadvantageous, and modifi- cation may then take place in a retrograde direction. Thus in mammals, as in other classes, there are low as well as high forms, but by any tests that can be applied—especially those based on the state of development of the central nervous system—it will be seen that the average exceeds that of any other class; that the class contains many species far excelling those of any other in perfection of structure, and especially one form which is un- questionably the culminating point yet arrived at amongst organised beings. With regard to the time of the first appearance of mammals upon the earth, the geological record is provokingly imperfect. At the commencement of the Tertiary period they were abundant, and already modified into most of the leading types at present existing, It was at one time thought that they first came into being at this date, but the discovery of more or less fragmentary remains of numerous and generally small species has revealed the existence of some forms of the class at various periods throughout almost the whole of the age of the deposition of the Secondary or Mesozoic rocks. This subject will be reverted to later on. It hardly need be said that mammals are vertebrated animals, and possess all the characteristics common to the members of that division of the animal kingdom. They are separated from the Ichthyopsida (fishes and amphibians), and agree with the Sauropsida (reptiles and birds) in the possession during their development of an amnion and allantois, and in never having external branchie or gills. They differ from reptiles and resemble birds in being warm- blooded, and having a heart with four cavities and a complete double circulation. They differ from both birds and reptiles in the red corpuscles of the blood being non-nucleated and, with very few INTRODUCTORY 3 exceptions, circular in outline ; in the lungs being freely suspended in a thoracic cavity, separated from the abdomen by a complete muscular partition—the diaphragm—which is the principal agent in inflating the lungs in respiration ; in having but one aortic arch, which curves over the left bronchus ; in the skin being more or less clothed with hair; in the greater perfection of the commissural system of the cerebral hemispheres, which has either a complete corpus callosum, or an incomplete one associated with a very large anterior commissure; in having no syrinx or inferior vocal organ, but a complete larynx at the upper end of the trachea; in having a mandible of which each ramus (except in very early developmental conditions) consists of a single bone on each side, articulating to the squamosal without the intervention of a quad- rate bone; in having a pair of laterally placed occipital condyles instead of one median one; and in the very obvious character of the female being provided with mammary glands, by the secretion of which the young (usually produced alive, although in the lowest forms by means of externally hatched eggs) are nourished for some time after birth. In common with all vertebrated animals, mammals never have more than two pairs of limbs ; as the larger number live ordinarily on the surface of the earth, in the great majority of the class both pairs are well-developed and functional, and adapted for terres- trial progression. Mammals are, however, by no means limited to this situation. Thus some species spend the greater part of their lives beneath the surface, their fore limbs being specially modified for burrowing; others, again, are habitually arboreal, their limbs being fitted for climbing or hanging to boughs of trees; some are as aerial as birds, the fore limbs being developed into wings of a special character ; while in others which are as aquatic as fishes, the limbs assume the form of fins or paddles. In many of the latter the hinder extremities are either completely suppressed, or present only in a rudimentary state. In no known mammal are the fore limbs absent. The hinder extremity of the axis of the body is usually prolonged into a tail, which may be a mere pendent appendage, or may be modified to perform various functions, as grasping boughs in climbing, or even gathering food, in the case of the prehensile- tailed Monkeys and Opossums, swimming in the Cetacea, and acting as a flap to drive away troublesome insects from the skin in the Ungulata. The state of development of the young at the time of birth varies greatly in the different groups. Thus among the Marsupials where there is no connection during intra-uterine life between the circulatory systems of the parent and the foetus, the young are born in an exceedingly imperfectly developed condition. For their 4 INTRODUCTORY protection the mother, in a large number of cases, has a special pouch enclosing the mamme, into which the young are transferred at birth, and in which they remain till they are well developed. Among the higher, or Placental types, however, where a connection exists between the maternal and foetal circulations previous to birth, the young are always born in a much more highly developed state than among the Marsupials, although we meet with great variations in this respect. In those forms which habitually live in holes, like many Rodents, the young are always very helpless at birth; and the same is also true of many of the Carnivora, which are well able to defend their young from attack. In the great order of Ungulate, or Hoofed Mammals, where in the majority of cases defence from foes depends upon fleetness of foot, or upon huge corporeal bulk, the young are born in a very highly developed condition, and are able almost at once to run by the side of the parent. This state of relative maturity at birth reaches its highest development in the Cetacea, where it is evidently associated with the peculiar conditions under which these animals pass their existence. In the Primates, however, we again find the young produced in a more or less helpless condition, and requiring a long period before they attain their full development, this being more especially the case with those higher forms which approximate in structure to man. In point of size mammals vary to a greater extent than the existing members of any one class of animals, and include the largest living inhabitants of the earth. The extremes of size are marked on the one hand by the whale known as Sibbald’s Rorqual, which attains a length of eighty feet and a weight of nearly as many tons, and on the other by the Pigmy-Shrew and the minute Harvest- mouse, which can climb a stem of wheat. Of all the living creatures inhabiting our globe, mammals are by far the most important in their economic uses, since, in addition to being the only animals capable of labour for human benefit, they furnish the greater portion of the animal food of many races of man, and likewise a large amount of their clothing. In these respects the Ungulates hold the first place. As regards employment for labour, with the exception of the Dogs used for sleighing by the Esquimaux, and those which among some European nations draw light carts, all the mammals in general use are Ungulates. Of the first importance are the Horses and Asses, which are employed as beasts of draught or burden over nearly the whole globe. Among many nations, however, cattle, as represented by the true Oxen, the Buffalos, and the Yaks of Tibet, occupy a still more important position, while in the highlands of Tibet Sheep are largely used for carrying burdens. In other regions, again, the place of the Horse and the Ass is taken by the Camels, INTRODUCTORY 5 which are peculiarly fitted for traversing parched and arid deserts, while in the Andes we find the Llamas serving the same office. In Lapland and other parts of the northern regions the Reindeer is the main agent employed in draught. Lastly, we must not omit to mention the Indian Elephant, which, from its vast strength, is so useful in transport through the wilder parts of its native country. As regards food, we again find the Ungulates, and more especially the Artiodactyle division, taking the foremost place; and in this connection we have only to mention, among animals kept in a domestic condition, Swine, Cattle, Sheep, and Goats—the three latter affording not only their flesh, but also milk and its resulting cheese and butter. To many races, however, Mares and Camels are the chief milk producers, while the Laps make use of the milk of the Reindeer. The Rodents, as represented by Hares and Rabbits, occupy a minor position as furnishers of food. In relation to clothing, the Ungulates are likewise of paramount importance, as exemplified by the wool of the Sheep, which is so valuable on account of its peculiar property of felting. Furs, however, are mostly yielded by mammals of other orders, among which the Fur-seals are perhaps the most important at the present day. Many other Carnivores yield valuable furs, among which may be mentioned Bears, Foxes, Racoons, Skunks, Minks, Otters, and Ermines. Of less importance are certain Rodents, such as the Squirrels, Rabbits, Hares, etc., while the hair of the Beaver was formerly much sought after for the manufacture of hats. Returning to the Ungulates, we may notice the importance of horse-hair, the employment of camel’s hair for brushes, and the many uses of the bristles of the pig. Some of the Monkeys yield fur which has been extensively used. Leather, again, is almost exclusively supplied by mammals, and mainly by the Ungulates. Three other important products, namely horn, buck’s-horn, and ivory, are likewise obtained solely from the same great order. Horn, as we shall notice in the sequel, is the sheath covering the bony horn-cores of the Oxen, while buck’s-horn is the commercial term applied to the antlers of the Deer, which are largely used for knife-handles and other purposes. True ivory is the product of the two species of Elephant; but other kinds of ivory are obtained from the teeth of the Sperm Whale and the tusks of the Walrus and Hippopotamus, the latter kind having been extensively employed some years ago for artificial teeth. For many purposes the place of ivory is taken by bone, this being mostly obtained from Ungulates. The bones of Camels are of an especially firm texture and good colour, and are largely employed in India for inlaying. Other important uses of bones are in the form of bone-dust as manure, and also as a source of phosphoric acid. The horns of the African Rhinoceros and the hide of the Hippopotamus are occasionally 6 INTRODUCTORY manufactured into small canes or whips. Horns and hoofs are also largely employed in the manufacture of glue. Formerly the so-called whalebone, or more properly baleen, was much used, especially to form the ribs of umbrellas and in stiffening ladies’ apparel, but the gradual destruction of the Right Whales, its only source of supply, has largely restricted its use of late years. The Cetacea are also of great economical importance from the abundance of oil yielded by the thick layer of blubber underlying the skin. Large quantities of valuable oil are also furnished by the Walrus and the Seals. Spermaceti, which was at one time extensively used in the manufacture of candles, is obtained from a large cavity in the head of the Sperm Whale or Cachalot, and also from the Hyperoddon or Bottle-nosed Whale. The nature of ambergris, a peculiar substance found floating on the surface of the sea and employed in perfumery, was long a matter of controversy ; but it appears to be an intestinal concretion of the Sperm Whale. Other substances of more importance to the perfumer are musk, the product of the Musk-Deer of the Himalaya, and civet, which is obtained from the so-called Civet Cat and other allied Carnivores. A secretion of the Beaver has also been used in perfumery and in medicine. CHAPTER II GENERAL ANATOMICAL CHARACTERS I. TEGUMENTARY STRUCTURES Hair.—The external surface of the greater number of members of the class is thickly clothed with a peculiarly modified form of epidermis, commonly called hair. This consists of hard, elongated, slender, cylindrical or tapering, filiform, unbranched masses of epidermic material, growing from a short papilla sunk at the bottom of a follicle in the derm or true skin. Such hairs upon different parts of the same animal, or upon different animals, assume various forms, and are of various sizes and degrees of rigidity,—as seen in the delicate soft velvety fur of the Mole, the stiff bristles of the Pig, and the spines of the Hedgehog and Porcupine, all modifications of the same structures. Each hair is composed usually of a cellular pithy internal portion, containing much air, and a denser or more horny cortical part. In some animals, as Deer, the substance of the hair is almost entirely composed of the medullary or cellular substance, and it is consequently very easily broken ; in others the horny part prevails almost exclusively, as in the bristles of the Wild Boar. In the Three-toed Sloth (Bradypus) the hairs have a central horny axis and a pithy exterior. Though generally nearly smooth, or but slightly scaly, the surface of some hairs is strongly imbricated, notably so in some Bats; while in the Two-toed Sloth (Cholepus) the hairs are longitudinally grooved or fluted. Though usually more or less cylindrical or circular in section, hairs are often elliptical or flattened, as in the curly-haired races of men, the terminal portion of the hair of Moles and Shrews, and conspicuously in the spines of the Rodents Xerus and Platacantho- mys. Hair having a property of mutual cohesion or “felting,” which depends upon a roughened scaly surface and a tendency to curl, as in domestic Sheep (in which animal this property has been especially cultivated by selective breeding), is called “wool.” 8 GENERAL ANATOMICAL CHARACTERS In a large number of mammals hairs of one kind only are scattered pretty evenly over the surface ; but in many there are two kinds, one longer, stiffer, and alone appearing on the surface, and the other shorter, finer, and softer, constituting the under fur, analogous to the down of birds. This under fur, or pashm as it is called by the natives of Kashmir, is especially abundant in the mammals inhabiting the cold plateau of Tibet and the adjacent regions. In many cases hairs of a different character from those of the general surface grow in special regions, forming ridges or tufts on the median dorsal or ventral surface or elsewhere. The tail is very often completed in this way by variously disposed elongated hairs. The margins of the eyelids are almost always furnished with a special row of stiffish hairs, called cilia or eyelashes ; and in most mammals specially modified hairs, constituting the vibrisse or whiskers, and endowed, through the abundant nerve supply of their basal papillae, with special tactile powers, grow from the lips and cheeks. In some mammals the hairy covering is partial and limited to particular regions ; in others, as the Hippopotamus and the Sirenia, though scattered over the whole surface, it is extremely short and scanty ; but in none is it reduced to so great an extent as in the Cetacea, in which it is limited to a few small bristles confined to the neighbourhood of the lips and nostrils, and often only present in the young or even foetal condition. Some kinds of hairs, as those of the mane and tail of the Horse, appear to persist throughout the life-time of the animal; but more generally, as in the case of the body hair of the same animal, they are shed and renewed periodically, generally annually. Many mammals have a longer hairy coat in winter, which is shed as summer comes on; and some few, which inhabit countries covered in winter with snow, as the Arctic Fox, Variable Hare, and Ermine, undergo a complete change of colour in the two seasons, being white in winter, and gray or brown in summer. The several species of Cape Mole (Chrysochloris), the Desmans or Water Moles (Myogale), and Potamogale velot, are remarkable as being the only mammals whose hair reflects those iridescent tints so common in the feathers of tropical birds. The principal and most obvious purpose of the hairy covering is to protect the skin against external influences, especially cold and damp. Its function in the hairless Cetacea is supplied by the specially modified and thickened layer of adipose tissue beneath the skin, called “ blubber.” Colour.—From the consideration of hair we are easily led to that of colour. As a general rule, bright and primary colours are absent in the class ; but among the Baboons we find brilliant patches of scarlet or blue on some of the bare portions of the body, and one of the South American Monkeys (Brachyurus) has its whole face of TEGUMENTARY STRUCTURES 9 a bright crimson. The most general colours are various shades of gray, brown, and tawny, with a frequent tendency to whiteness of the ventral surface of the body ; but among the Squirrels, and more especially those provided with a parachute for flying, we find brilliant russets, passing into orange and red. Dark brown or black is also “not very uncommon, as in the Bears and the Sable Antelope of South Africa. Entirely white mammals are rare, and mostly characteristic of the polar regions, or of countries having a long and snowy winter. An entirely white Bat (Diclidwrus albus) occurs, however, in South America. In the large majority of mammals that exhibit a varied coloration, the upper and most exposed parts of the surface present the richest and darkest colours, the under parts being pale or often quite white. The Ratels, Gluttons, 4lurus, Hamsters, and some others are exceptions to this rule. A large number of mammals having a ground colour of gray, tawny, or dun are marked by stripes or spots, which are generally of a darker hue than the ground colour, as in many Carnivora, but more rarely are lighter, as in the Fallow and Axis Deer and several species of Ante- lope. These stripes very generally run transversely to the axis of the body, as in the Tasmanian Thylacine, the Tiger, and the Zebra; but they may be longitudinal, as in several of the Civet family. There has been considerable discussion as to whether the striped or the spotted is the more primitive type of coloration; but no very conclusive arguments have been brought forward in favour of either view. It is, however, manifest that in several groups of mammals there is a tendency to lose the spots, and more rarely the stripes, and to assume a uniform colour. Thus the young of nearly all the species of Deer are spotted, whereas the adults of only the Fallow and Axis Deer are so marked. The same is true of most of the Pigs ; and the young of the Malayan and American Tapirs are marked by light-coloured stripes and spots on a dark ground. In like manner the young of the Lion and the Puma exhibit distinct spots which disappear with advancing age. In most of our domestic horses of various shades of bay and brown we may detect “‘ dappling ” on the under hair when the outer coat has been removed, which is not apparent on the surface of the latter. Many varieties of the Ass and the Horse also exhibit a tendency to the presence of stripes on the legs, which would seem to indicate a descent from a striped Zebra-like type. A peculiar feature, which is, however, common to many other groups of animals, is the tendency to what is known as melanism, or the production of black or dark individuals or races of particular species, due to an excess of pigment in the skin and hair. Thus we may have black Leopards and Jaguars, black Wolves, and black Rabbits. The opposite to melanism, and of more frequent occurrence, is 10 GENERAL ANATOMICAL CHARACTERS albinism—a condition in which the pigment or colouring matter usually present in the tissues constituting the external coverings of the body, and which gives them their characteristic hue, is absent. When it occurs the hair is of an opaque white, the claws, hoofs, etc., of a pale horn-colour, and the skin and eyes pink, in consequence of the colour of the blood which circulates through them being no longer concealed by the stronger hues of the pigments. An animal in this condition is called an albino. In complete albinism there is a total absence of pigment throughout the system. This condition occurs occasionally as an individual peculiarity among wild animals of many kinds; but it has never been perpetuated among them in dis- tinct races or species. The disadvantage of absence of pigment in the eye, causing a certain amount of intolerance of light, is probably sufficient to account for this. Several races of true albinos, as White Ferrets, Rabbits, Rats, and Mice, have, however, been established under the protection of man, and in them this ab- normal condition is propagated from generation to generation. Partial albinism—a condition in which the absence of pigment is limited to portions of the surface, or, at all events, does not extend to the eyes—is much more common as an individual variation both in domestic and in wild animals. It is possible that the artificial conditions incident to domestication increase the tendency to its occurrence ; but, whether this be so or not, it certainly becomes perpetuated more frequently among domesticated than among wild animals. This may be accounted for partly by its proving of no disadvantage to them, and partly by the frequent selection by man of animals of such colour in preference to others. The result is that there is no completely domestic animal of which white races do not exist. On the other hand, to most wild animals even partial albinism seems to be a disadvantage in the struggle for existence, since, except in the case of species inhabiting lands continually covered with snow, it renders them more conspicuous objects both to their enemies and their prey, and hence it is rarely perpetuated. In northern regions, however, a large proportion of species are regularly and normally of a white colour, either, as the Polar Bear, all the year through, or, as the Ermine or Stoat, Arctic Fox, and Alpine Hare, during the winter season. The coloration in these cases is obviously protective, as it is also to a great extent in many other instances throughout the class. Among conspicuously coloured mammals, it has been observed that the vertical black and tawny stripes of the Tiger harmonise so well with the brown and green grasses of its native jungle as to render the animal almost invisible when lying among them ; while the dappled hide of the Giraffe is said to agree equally well with the chequered splashes of light and shade in the clumps of tall mimosas among which it feeds) The uniformly tawny hue of the TEGUMENTARY STRUCTURES II Lion accords well with the prevailing tint of its native desert ; and any one who has seen an Elephant or Buffalo in the deep shades of an Indian forest will realise how perfectly adapted is their dull, slaty colour to concealment in such a spot. The dun colour of the Wild Ass of India is equally well suited to the sandy deserts of Kutch ; it is also stated that the brilliant stripes of the Zebras of Africa are arranged in such proportion as exactly to match the pale tint which arid ground possesses when seen by moonlight.1 The most remarkable instance of protective coloration is, however, to be found in the Sloths of South America, in which the coarse gray hairs so closely resemble a mass of lichenous growth that it is almost impossible to distinguish these animals when at rest from the gnarled and lichen-clad boughs from which they suspend them- selves. This resemblance is increased by the fact that the hairs actually develop a growth of lichens upon themselves. That the sombre coloration of these animals has been produced to harmonise with their present surroundings seems to be evident by the circum- stance that when the long hair is plucked off the under fur is seen to present a bold alternation of black and yellow stripes, which may probably be regarded as the original primitive coloration of this group. Scales, etc.—True scales, or flat imbricated plates of horny material, covering the greater part of the body, so frequently occurring in reptiles, are found only in one family of mammals, the Mande or Pangolins; but these are also associated with hairs growing from the intervals between the scales, or on the parts of the skin not covered by them. Similarly, imbricated epidermic productions form the covering of the under surface of the tail of the flying Rodents of the genus Anomalurus ; and flat scutes, with the edges in apposition, and not overlaid, clothe both surfaces of the tail of the Beaver, Rats, and others of the same order, and also of some Insectivores and Marsupials. The Armadillos alone have an ossified exoskeleton, composed of plates of true bony tissue, developed in the derm or corium, and covered with scutes of horny epidermis. Other epidermic appendages are the horns of Ruminants and Rhinoceroses,—the former being elongated, tapering, hollow caps of hardened epidermis of fibrillated structure, fitting on and growing from conical projections of the frontal bone, and always arranged in pairs, while the latter are of similar structure, but solid and without any internal bony support, and (in all existing species) situated in the median line. Callosities, or bare patches covered with hardened and thickened epidermis, are found covering the pads under the soles of the feet and undersurfaces of the toes of nearly all mammals, upon the ischial tuberosities of many Apes, the sternum of Camels, on the inner side of the limbs of the 1 Galton’s South Africa, p. 187. 12 GENERAL ANATOMICAL CHARACTERS Equide, the grasping under surface of the tail of the prehensile-tailed Monkeys, etc. The greater part of the skin of both species of one-horned Asiatic Rhinoceros is immensely thickened and stiffened by increase of the tissue both of the derm and epiderm, con- stituting the well-known jointed “armour-plated” hide of those animals. Nails, Claws, and Hoofs.—With very few exceptions, the terminal extremities of the digits of both limbs are more or less protected or armed by epidermic plates or sheaths, constituting the various forms of nails, claws, or hoofs. These are wanting in the Cetacea alone. A perforated spur, with a special secreting gland in connection with it, is found attached to the hind leg of the males of the three genera of Monotremata, Ornithorhynchus, Proechidna, and Echidna, Odour - secreting Glands.—Besides the universally distributed sebaceous glands connected with the pilose system, most mammals have special glands situated in modified portions of the integument, often involuted to form a shallow recess or a deep sac with a narrow opening, situated in various parts of the surface of the body, and secreting odorous substances, by the aid of which individuals appear to recognise one another, and probably affording the princi- pal means by which wild animals are able to become aware of the presence of other members of the species, even at great dis- tances. Although the commencement of the modifications of portions of the external covering for the formation of special secretions may be at present difficult to understand, the principle of natural selection will readily explain how such organs become fixed and gradually increase in development in any species, especi- ally as there would probably be a corresponding modification and increased sensibility of the olfactory organs. Such individuals as by the intensity and peculiarity of their scent had greater power of attracting the opposite sex would certainly be those most likely to leave descendants to inherit and in their turn propagate the modi- fication. To this group of structures belong the suborbital gland or “crumen” of Antelopes and Deer, the frontal gland of the Muntjak and of Bats of the genus Hipposiderus, the submental gland of the Chevrotains and of Taphozous and some other Bats, the post-auditory follicle of the Chamois, the temporal gland of the Elephant, the lateral glands of the Musk-Shrew, the dorsal gland of the Peccary, the inguinal glands of Antelopes, the preputial glands of the Musk- Deer and Beaver (already alluded to in connection with the use made of their powerfully odorous secretion in medicine and per- fumery) and also of the Swine and Hare, the anal glands of Carni- vora, the perineal gland of the Civet (also of commercial value), the caudal glands of the Fox and Goat, the gland on the humeral membrane of Bats of the genus Saccopteryx, the post-digital gland of DENTAL SYSTEM 13 the Rhinoceros, the inter-digital glands of the Sheep and many Ruminants, and numerous others. In some of these cases the glands are peculiar to, or more largely developed in, the male; in others they are found equally developed in both sexes. Il, DENTAL SYSTEM The dental system of mammals may be considered rather more in detail than space permits for some other portions of their structure, not only on account of the important part it plays in the economy of the animals of this class, but also for its interest to zoologists as an aid in the classification and identification of species. Owing to the imperishable nature of their tissues, teeth are preserved for an. indefinite time, and in the case of extinct species frequently offer the only indications available from which to derive an idea of the characters, affinities, and habits of the animals to which they once belonged. Hence even their smallest modifications have received great attention from comparative anatomists, and they have formed the subject of many special monographs.! Teeth are present in nearly all mammals, and are applied to various purposes. They are, however, mainly subservient to the function of alimentation, being used either in procuring food, by seizing and killing living prey or gathering and biting off portions of vegetable material, and more indirectly in tearing or cutting through the hard protective coverings of food sub- stances, as the husks and shells of nuts, or in pounding, crushing, or otherwise mechanically dividing the solid materials before swallowing, so as to prepare them for digestion in the stomach. Certain teeth are also in many animals most efficient weapons of offence and defence, and for this purpose alone, quite irrespective of subserviency to the digestive process, are they developed in the male sex of many herbivorous animals, in the females of which they are absent or rudimentary. Teeth belong essentially to the tegumentary or dermal system of organs, and, as is well seen in the lower vertebrates, pass by almost insensible gradations into the hardened spines and scutes formed upon the integument covering the outer surface of the body ; but in mammals they are more specialised in structure and limited in locality. In this class they are developed only in the 1L. F. E. Rousseau, Anatomie comparée du Systeme dentaire chez ? Homme et chez les principaux Animaux, 2d ed., 1839; F. Cuvier, Des Dents des Mammiferes considérées comme caractéres zoologiques, 1822-25; R. Owen, Odontography, 1840-45 ; C. G. Giebel, Odontographic, 1855; C. S. Tomes, Manual of Dental Anatomy, Human and Comparative, 3d ed., 1889. 14 GENERAL ANATOMICAL CHARACTERS gums or fibro-mucous membrane covering the alveolar borders of the upper and lower jaws, or, in other words, the premaxillary and maxillary bones and the mandible. In the process of develop- ment, for the purpose of giving them that support which is needful for the performance of their functions, they almost always become implanted in the bone,—the osseous tissue growing up and mould- ing itself around the lengthening root of the tooth, so that ultimately they become apparently parts of the skeleton. In no mammal, however, does ankylosis or bony union between the tooth and jaw normally take place, as in many fishes and reptiles, —a vascular layer of connective tissue, the alveolo-dental mem- brane, always intervening! The presence of two or more roots, frequently met with in the cheek-teeth of mammals, implanted in corresponding distinct sockets of the jaw, is now peculiar to animals of this class.” : Structure-—The greater number of mammalian teeth when fully formed are not simple and homogeneous in structure, but are com- posed of several distinct tissues, which are enumerated below. The pulp, a soft substance, consisting of a very delicate gelatinous connective tissue, in which numerous cells are imbedded, and abundantly supplied with blood-vessels and nerves, constitutes the central axis of all the basal part of the tooth, and affords the means by which the vitality of the whole is preserved. The nerves which pass into the pulp and endow the tooth with sensibility are branches of the fifth pair of cranial nerves. The pulp occupies a larger relative space, and performs a more important purpose, in the young growing tooth than afterwards, as by the calcification and conversion of its outer layers the principal hard constituent of the tooth, the dentine, is formed. In teeth which have ceased to grow the pulp occupies a comparatively small space, which in the dried tooth is called the pulp-cavity. This communi- cates with the external surface of the tooth by a small aperture at the apex of the root, through which the branches of the blood- vessels and nerves, by which the tooth receives its nutrition and sensitiveness, pass in to be distributed in the pulp. In growing teeth the pulp-cavity is widely open, while in advanced age it often becomes obliterated, and the pulp itself entirely converted into bone-like material. The dentine or ivory forms the principal constituent of the greater number of teeth. When developed in its most character- istic form, it is a very hard but elastic substance, white, with a yellowish tinge, and slightly translucent. It consists of an organic 1 The lower incisors of some species of Shrews are, however, said to become ankylosed to the jaw in adult age. * The teeth of the extinct Dinosaurian reptile Triceratops have two distinct roots, placed transversely to the axis of the jaws. DENTAL SYSTEM 15 matrix, something like, but not identical with, that of bone, richly impregnated with calcareous salts (chiefly calcium phosphate), these constituting in a fresh human tooth 72 per cent of its weight. When subjected to microscopical examination it is seen to be every- where permeated by nearly parallel branching tubes which run, in a slightly curving or wavy manner, in a general direction from the centre towards the free surface of the tooth. These tubes com- municate by open mouths with the pulp-cavity, and usually ter- minate near the periphery of the dentine by closed ends or loops, though in Marsupials and certain other mammals they penetrate into the enamel. They are occupied in the living tooth by soft gelatinous fibrils connected with the cells of the pulp. A variety of dentine, permeated by canals containing blood-vessels, met with commonly in fishes and in some few mammals, as the Megatheriwm, is called vaso-dentine. Other modifications of this tissue occasionally met with are called osteo-dentine and secondary dentine,—the latter being a dentine of irregular structure which often fills up the pulp-cavity of old animals. The enamel constitutes a thin investing layer, complete or partial, of the outer or exposed and working surface of the dentine of the crown of the teeth of most mammals. This is the hardest tissue met with in the animal body, containing from 95 to 97 per cent of mineral substances (chiefly calcium phosphate and some carbonate, with traces of fluoride). Its ultimate structure consists of prismatic fibres, placed generally with their long axes at right angles to the free surface of the tooth. Enamel is easily distin- guished from dentine with the naked eye by its clear, bluish-white, translucent appearance. The cement or crusta petrosa is always the most externally placed of the hard tissues of which teeth are composed, as will be under- stood when the mode of development of these organs is considered. It is often only found as a thin layer upon the surface of the root; but sometimes, as in the complex-crowned molar teeth of the Horse and Elephant, it is a structure which plays a very important part, covering and filling in the interstices between the folds of the enamel. In appearance, histological structure, and chemical com- position it is closely allied to osseous tissue, containing lacune and canaliculi, though only when it is of considerable thickness are Haversian canals present in it. Development.—The two principal constituents of the teeth, the dentine and the enamel, are developed from the two layers of the mucous membrane of the jaw—the dentine from the deeper or vas- cular, the enamel from the superficial or epithelial layer. The latter dips down into the substance of the gum, and forms the enamel-organ or germ, the first rudiment of the future tooth, which is constantly present even in those animals in which the enamel is not found as a -“ 16 GENERAL ANATOMICAL CHARACTERS constituent of the perfectly-formed tooth. Below the mass of epi- thelial cells thus embedded in the substance of the gum, and remaining connected by a narrow neck of similar structure with the epithelium of the surface, a portion of the vascular areolar tissue becomes gradually separated and defined from that which surrounds it, and assumes a distinct form, which is that of the crown of the future tooth,—a single cone in the case of simple teeth, or with two or more eminences in the complex forms. This is called the dental papilla or dentine germ, and by the gradual conversion of its tissue into dentine the bulk of the future tooth is formed, the uncalcified central portion remaining as the pulp. The conversion of the papilla into hard tissue commences at the outer surface of the apex, and gradually proceeds downwards and inwards, so that the form of the papilla exactly determines the form of the future dentine, and no alteration either in shape or size of this portion of the tooth, when once calcified, can take place by addition to its outer surface. In the meanwhile, calcification of a portion of the cells of the enamel- organ, which adapts itself like a cap round the top of the dentinal papilla, and has assumed a somewhat complex structure, results in the formation of the enamel-coating of the crown of the tooth. While these changes are taking place the tissues immediately sur- rounding the tooth-germ become condensed and differentiated into a capsule, which appears to grow up from the base of the dental papilla, and encloses both this and the enamel-germ, constituting the follicle or tooth-sac. By the ossification of the inner layer of this follicle the cement is formed. This substance, therefore, unlike the dentine, increases from within outwards, and its growth may accordingly be the cause of considerable modification of form and enlargement, especially of the roots, of certain teeth, as those of Seals and some Cetacea. The delicate homogeneous layer coating the enamel surface of newly-formed teeth, in which cement is not found in the adult state, and known as Nasmyth’s membrane, is considered by Tomes as probably a film of this substance, too thin to exhibit its characteristic structure, though by others it is believed to be derived from the external layer of the enamel-organ. The homology of the teeth with the dermal appendages, hairs, scales, and claws, has already been alluded to, and it will now be seen that in both cases two of the primary embryonic layers are concerned in their develop- ment—the mesoblast and epiblast—although in very different pro- portions respectively. Thus in the hair or nail the part derived from the epiblast forms the principal bulk of the organ, the mesoblast only constituting the papilla or matrix. But in the tooth the epi- blastic portion is limited to the enamel, and is always of relatively small bulk and often absent, while the dentine (the principal con- stituent of the tooth) and the cement are formed from the mesoblast, When more than one set of teeth occur in mammals, those of DENTAL SYSTEM 17 the second set are developed in a precisely similar manner to the: first, but the enamel-germ, instead of being derived directly from an independent part of the oral epithelium, is formed from a budding out of the neck of the germ of the tooth succeeded. In the case of the true molars, which have no predecessors, the germ of the first has an independent origin, but that of the others is derived from the neck of the germ of the tooth preceding it in the series. The foundations of the permanent teeth are thus laid as it were almost simultaneously with those of their predecessors, although they remain in many cases for years before they are developed into functional activity. Although the commencement of their formation takes place at an early period of embryonic life, teeth are in nearly all mam- mals still concealed beneath the gum at the time of birth. The period of eruption, or “cutting” of the teeth as it is called, that is, their piercing through and rising above the surface of the mucous membrane, varies much in different species. In some, as Seals, the whole series of teeth appears almost simultaneously; but more often there are considerable intervals between the appearance of the individual teeth, the front ones usually coming into place first, and those at the back of the mouth at a later period. Forms of Teeth.—The simplest form of tooth may be exemplified on a large scale by the tusk of the Elephant (Fig. 1,1) Itisa hard mass almost entirely composed of dentine, of a conical shape at first, but during growth becoming more and more cylindrical or uniform in width. The enamel-covering, present on the apex in its earliest condition, soon disappears, but a thin layer of cement covers the circumference of the tooth throughout life. In section it will be seen that the basal portion is hollow, and contains a large conical pulp, as broad at the base as the tooth itself, and deeply imbedded in the bottom of a recess, or socket, in the maxillary bone. This pulp continues to grow during the lifetime of the animal, and at the same time is converted at its surface into dentine. The tooth therefore continually elongates, but the use to which the animal subjects it in its natural state causes the apex to wear away, at a rate generally proportionate to the growth at the base, other- wise it would become of inconvenient length and weight. Such teeth of indefinite growth are said to be “rootless,” or to have “persistent pulps.” One of the corresponding front teeth of man (Fig. 2, II. and III.) may be taken as an example of a very different condition. After its crown is fully formed by calcification of the germ, the pulp, though continuing to elongate, begins to contract in diameter; a neck or slight constriction is formed ; and the remainder of the pulp is con- verted into the root (often, but incorrectly, called “fang”), a taper- ing conical process imbedded in the alveolar cavity of the bone, and 2 18 GENERAL ANATOMICAL CHARACTERS having at its extremity a minute perforation, through which the vessels and nerves required to maintain the vitality of the tooth enter Fia. 1.—Diagrammiatic Sections of various forms of Teeth. 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 huinan 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 the Ox, with the enamel covering the crown deeply folded, and the depressions filled up 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. tooth. the pulp-cavity, which is very different from the widely open cavity at the base of the growing tooth, When the crown of the tooth is broad and complex in character, in- stead of having a single root, it may be supported by two or more roots, each of which is implanted in a distinct alveolar recess or socket, and to the apex of which a branch of the com- mon pulp-cavity is continued (Fig. 1,1V.) Such teeth are called “rooted teeth.” When they have once attained their position in the jaw, with the neck a little way above the level of the free margin of the alveolus, and embraced by the gum or tough fibro- vascular membrane covering the alveolar border, and hav- ing the root fully formed, they can never increase in length or alter their posi- tion ; if they appear to do so in old age, it being only in consequence of absorption and retrocession of the sur- rounding alveolar margins. Tf, as often happens, their surface Wears away in mas- tication, it is never renewed. The open cavity at the base of the imperfectly developed tooth (Fig. 1, IL.) causes it to resemble the persistent condition of the rootless The latter is therefore a more primitive condition, the formation of the root being a completion of the process of tooth development. Functionally it is, however, difficult to say that the DENTAL SYSTEM 19 one is a higher form than the other, since they both serve important and different purposes in the animal economy. As is almost always the case in nature, intermediate conditions between these two forms of teeth are met with. Thus some teeth, as the molars of the Horse, and of many Rodents, are for a time rootless, and have growing pulps producing very long crowns with parallel sides, the summits of which may be in use and beginning to wear away while the bases are still growing; but ultimately the pulp contracts, forms a neck and distinct roots, and ceases to grow. The canine tusks of the Musk Deer and of the Walrus have persistent pulps, and are open at their base until the animal is of advanced age, when they close, and the pulp ceases to be renewed. The same sometimes happens in the tusks of very old Boars. The simplest form of the crown of a tooth is that of a cone; but this may be variously modified. Thus it may be flattened, with its edges sharp and cutting, and pointed at the apex, as in the laterally compressed premolars of most Carnivora; or it may be chisel- or awl-shaped, with a straight truncated edge, as in the human incisors ; or it may be broad, with a flat or rounded upper surface. Very often there is a more or less prominent ridge encircling the whole or part of the base of the crown just above the neck, called the cingu- lum, which serves as a protection to the edge of the gum in masti- cating, and is most developed in flesh-eating and insectivorous animals, in which the gums are liable to be injured by splinters of bone or other hard fragments of their food. The form of the crown is frequently rendered complex by the development upon its surface of elevations or tubercules called cusps or cones, or by ridges usually transverse, but sometimes variously curved or folded. When the crown is broad and the ridges are greatly developed, as in the molars of the Elephant, Horse, and Ox (Fig. 1, V.), the inter- spaces between them are filled with cement, which supports them and makes a solid compact mass of the whole tooth. When such a tooth wears away at the surface by friction against the opposed tooth of the other jaw, the different density of the layers of the substances of which it is composed—enamel, dentine, and cement—arranged in characteristic patterns, causes them to wear unequally, the hard enamel ridges projecting beyond the others, and thus giving rise to a grinding surface of great mechanical advantage. Succession.—The dentition of all mammals consists of a definite set of teeth, almost always of constant and determinate number, form, and situation, and, with few exceptions, persisting in a functional condition throughout the natural term of the animal’s life. In many species these are the only teeth which the animal ever possesses, —the set which is first formed being permanent, or, if accidentally lost, or decaying in extreme old age, not being replaced 20 GENERAL ANATOMICAL CHARACTERS by others. These animals are called Monophyodont. But in the larger number of mammals, certain of the teeth are preceded by others, which may be only of a very transient, rudimentary, and functionless character (being in the Seals, for example, shed either before or within a few days after birth), or may be considerably developed, and functionally occupy the place of the permanent teeth for a somewhat lengthened period, during the growth and develop- ment of the latter and of the jaws. In all cases these teeth disappear (by the absorption of their roots and shedding of the crowns) before the frame of the animal has acquired complete maturity, as evidenced by the coalescence of the epiphyses of the osseous system. As these teeth are, as a general rule, present during the period in which the animal is nourished by the milk of the mother, the name of “milk-teeth” (French dents de lait, German milchzéhne) has been commonly accorded to them, although it must be understood that the epoch of their presence is by no means necessarily synchronous with that of lactation. Animals possessing such teeth are called Diphyodont. No mammal is known to have more than two sets of teeth ; and the definite and orderly replacement of certain members of the series is a process of quite a different nature from the indefinite succession which takes place in all the teeth continuously throughout the lifetime of the lower vertebrates. When the milk-teeth are well developed, and continue in place during the greater part of the animal’s growth, as is especially the case with the Ungulata, and, though to a less degree, with the Primates and Carnivora, their use is obvious, since taken all together they form structurally a complete epitome on a small scale of the more numerous and larger permanent set (see Fig. 3), and, con- sequently, are able to perform the same functions, while time is allowed for the gradual maturation of the latter, and especially while the jaws of the growing animal are acquiring the size and strength sufficient to support the permanent teeth. Those animals, therefore, that have a well-developed and tolerably persistent set of milk-teeth may be considered to be in a higher state of development, as regards their dentition, than those that have the milk-teeth absent or rudimentary. It is a very general rule that individual teeth of the milk and permanent set have a close relationship to one another, being originally formed,as mentioned above, in exceedingly near proximity, and with, at all events so far as the enamel-germ is concerned, a direct connection. Moreover, since the latter ultimately come to occupy the position in the alveolar border temporarily held by the former, they are spoken of respectively as the predecessors or suc- cessors of each other. But it must be understood that milk-teeth may be present which have no successors in the permanent series, DENTAL SYSTEM 21 and, what is far more general, permanent teeth may have no pre- decessors in the milk series. The complete series of permanent teeth of most mammals forms a complex machine, with its several parts adapted for different functions,—the most obvious structural modification for this purpose being an increased complexity of the individual components of the series from the anterior towards the posterior extremity of such series. Since, as has just been said, the complete series of the milk teeth often presents structurally and functionally a similar machine, but composed of fewer individual members, and the anterior of which are as simple, and the posterior as complex as those occupying corresponding positions in the permanent series,—and since the milk-teeth are only developed in relation to the anterior or lateral, never to the most posterior of the permanent series,—it follows that the hinder milk-teeth are usually more complex than the teeth of which they are the predecessors in the permanent series, and represent functionally, not their immediate successors, but those more posterior permanent teeth which have no direct predecessors. This character is clearly seen in those animals in which the various members of the molar series are well differentiated from each other in form, as the Carnivora, and also in Man. In animals which have two sets of teeth the number of those of the permanent series which are preceded by milk-teeth varies greatly, being sometimes, as in Marsupials and some Rodents, as few as one on each side of each jaw, and sometimes including the larger portion of the series. Although there are difficulties in some cases in arriving at a satisfactory solution of the question, it is, on the whole, safest to assume that when only one set of teeth is present, this corresponds to the permanent teeth of the Diphyodonts. When this one set is completely developed, and remains in use throughout the animal’s life, there can be no question on this subject. When, on the other hand, the teeth are rudimentary and transient, as in the Whalebone Whales, it is possible to consider them as representing the milk series; but there are weighty reasons in favour of the opposite conclusion.! Arrangement, Homologies, and Notation of Teeth.— The teeth of the two sides of the jaws are always alike in number and character, 1 This and other questions concerning the homologies, notation, and suc- cession of the teeth of mammals are more fully developed in two memoirs by one of the present writers :—‘‘ Remarks on the Homologies and Notation of the Teeth of the Mammalia,” in the Journal of Anatomy and Physiology, vol. iti. p. 262, 1869 ; and ‘“‘ Notes on the First or Milk Dentition of the Mammalia,” in the Trans. Odontological Society of Great Britain, 1871. See also an important memoir by Oldfield Thomas on the ‘‘Homologies and Succession of the teeth in the Dasyuride,” Phil. Trans. 1887, pp. 443-462. 22 GENERAL ANATOMICAL CHARACTERS except in cases of accidental or abnormal variation, and in the one remarkable instance of constant deviation from bilateral symmetry among mammals, the tusks of the Narwhal (Monodon), in which the left is of immense size, and the right rudimentary. In cer- tain mammals, such as the Dolphins and some Armadillos, which have a very large series of similar teeth, not always constant in number in different individuals, there may be differences in the two sides; but, apart from these, in describing the dentition of any mammal, it is quite sufficient to give the number and characters of the teeth of one side only. Since the teeth of the upper and the lower jaws work against each other in masticating, there is a general correspondence or harmony between them, the projections of one series, when the mouth is closed, fitting into corresponding depressions of the other. There is also a general resemblance in the number, characters, and mode of succession of both series, so that, although individual teeth of the upper and lower jaws may not be in any strict sense of the term homologous parts, there is a great con- venience in applying the same descriptive terms to the one as are used for the other. The simplest dentition as a whole is that of many species of Dolphin (Fig. 2), in which the crowns are single-pointed, slightly Fig. 2.—Upper and Lower Teeth of one side of the Mouth of a Dolphin (Lagenorhynchus) as an example of the homodont type of dentition. The bone covering the outer side of the roots of the teeth has been removed to show their simple character. curved cones, and the roots also single and tapering, and all alike in form from the anterior to the posterior end of the series, though it may be with some slight difference in size, those at the two extremities of the series being rather smaller than the others. Such a dentition is called Homodont, and in the case cited, as the teeth are never changed, it is also Monophyodont. Such teeth are adapted only for catching slippery living prey, as fish. In a very large number of mammals the teeth of different parts of the series are more or less differentiated in character, and have different functions to perform. The front teeth are simple and one-rooted, and are adapted for cutting and seizing, They are called “incisors.” The back- or cheek-teeth have broader and more complex crowns, tuberculated or ridged, and are sup- DENTAL SYSTEM 23 ported on two or more roots. They crush or grind the food, and are hence called “molars.” Many animals have, between these two sets, a tooth at each corner of the mouth, longer and more pointed than the others, adapted for tearing or stabbing, or for fixing struggling prey. From the conspicuous development of such teeth in the Carnivora, especially the Dogs, they have received the name of “canines.” A dentition with its component parts so differently formed that these distinctive terms are applicable to them is called Heterodont. In most cases, though by no means invariably, animals with Heterodont dentition are also Diphyodont. This general arrangement is extremely obvious in a considerable number of mammals; and closer examination shows that, under very great modification in detail, there is a remarkable uniformity of essential characters in the dentition of a large number of members of the class belonging to different orders and not otherwise closely allied ; so much so indeed that it has been possible (chiefly through the researches of Sir Richard Owen) to formulate a common plan of dentition from which the others have been derived by the alteration of some and suppression of other members of the series, and occasionally, but very rarely, by addition. The records of paleontology fully confirm this view, as by tracing back many groups now widely separated in dental characters we find a gradual approximation toa common type. In this generalised form of mammalian dentition (which is best exemplified in the genera Anoplotherium and Homalodontotheriwm) the entire number of teeth present is 44, or 11 above and 11 below on each side. Those of each jaw are placed in continuous series without intervals between them; and, although the anterior teeth are simple and _ single- rooted, and the posterior teeth complex and with several roots, the transition between the two kinds is gradual. In dividing and grouping such teeth for the purpose of descrip- tion and comparison, more definite characters are required than those derived merely from form or function. The first step towards a classification has been made by the observation that the upper jaw is composed of two bones, the premaxilla and the maxilla, and that the suture between these bones separates the three anterior teeth from the others. These three teeth, then, which are implanted by their roots in the premaxilla, form a distinct group, to which the name of “incisor” is applied. This distinction is, however, not so important as it appears at first sight, for, as mentioned when speaking of the development of the teeth, their connection with the bone is only of a secondary nature, and, although it happens conveniently for our purpose that in the great majority of cases the segmentation of the bone coincides with the interspace between the third and fourth tooth of the series, still, when it does not happen to do so, as in the case of the Mole, we must not give 24 GENERAL ANATOMICAL CHARACTERS too much weight to this fact, if it contravenes other reasons for determining the homologies of the teeth. The eight remaining teeth of the upper jaw offer a natural division, inasmuch as the posterior three never have milk-predecessors ; and, although some of the anterior teeth may be in the same case, the particular one preceding these three always has such a predecessor. These three then are grouped apart as the “molars,” or, since some of the teeth in front of them often have a molariform character, “true molars.” Of the five teeth between the incisors and molars the most anterior, or that which is usually situated close behind the premaxillary suture, almost always, as soon as any departure takes place from the simplest and most homogeneous type, assumes a lengthened and pointed form, and is the tooth so developed as to constitute the “canine” or “laniary” tooth of the Carnivora, the tusk of the Boar, etc. It is customary therefore to call this tooth, whatever its size or form, the “canine.” The remaining four are the “ pre- molars” or ‘false molars.” This system of nomenclature has been objected to as being artificial, and in many cases not descriptive, the distinction between premolars and canine especially being sometimes not obvious; but the terms are now in such general use, and are so practically convenient—especially if, as it is best to do in all such cases, we forget their original signification and treat them as arbitrary signs—that it is not likely they will be super- seded by any that have been proposed as substitutes for them. With regard to the lower teeth the difficulties are greater, owing to the absence of any suture corresponding to that which defines the incisors above; but since the number of the teeth is the same, the corresponding teeth are preceded by milk-teeth, and in the large majority of cases it is the fourth tooth of the series which is modified in the same way as the canine (or fourth tooth) of the upper jaw, it is quite reasonable to adopt the same divisions as with the upper series, and to call the first three, which are implanted in the part of the mandible opposite to the premazxilla, the incisors, the next the canine, the next four the premolars, and the last three the molars. It may be observed that when the mouth is closed, especially when the opposed surfaces of the teeth present an irregular outline, the corresponding upper and lower teeth are not exactly opposite, otherwise the two series could not fit into one another; but as arule the points of the lower teeth shut into the interspaces in front of the corresponding teeth of the upper jaw. This is seen very distinctly in the canine teeth of the Carnivora, and is a useful guide in determining the homologies of the teeth of the two jaws. Objections have certainly been made to this view, because, in certain rare cases, the tooth which, accord- ing to it, would be called the lower canine has the form and function of an incisor (as in Ruminants and Lemurs), and on the DENTAL SYSTEM 25 other hand (as in Cotylops, an extinct Ungulate from North America) the tooth that would thus be determined as the first premolar has the form of a canine ; but it should not be forgotten that, as in all such cases, definitions derived from form and function alone are quite as open to objection as those derived from position and relation to surrounding parts, or still more so. Dental formule.—For the sake of brevity the complete dentition, arranged according to these principles, is often described by the following formula, the numbers above the line representing the * teeth of the upper, those below the line those of the lower jaw :— - : 1-1 4-4 8-3 11-11 3_3 canines >—, premolars 1 a ee total 44. Since, however, initial letters may be substituted for the names of each group, and it is quite unnecessary to give more than the numbers of the teeth on one side of the mouth, the formula may be conveniently abbreviated into— 1$,ct pt m $= 41; total 44. incisors molars The individual teeth of each group are always enumerated from before backwards, and by such a formula as the following— ; 41,72,73,¢,p1, p 2, p 3, p4, m1, m2, m3 41,72,73,¢,p1, p2, p38, p4,m1,m2,m3 or more briefly— .1,2,3 1 1, 2, 3,4 1, 2, 3 ae Ee ea ee A special numerical designation is thus given by which each one can be indicated. In mentioning any single tooth, such a sign as ™1 will mean the first upper molar, mi the first lower molar, and so on. The use of such signs saves much time and space in description.! It was part of the view of the founder of this system of dental notation that, at least throughout the group of mammals whose dentition is derived from this general type, each tooth has its strict homologue in all species, and that in those cases in which fewer than the typical number are present (as in all existing mammals except the genera Sus, Gymnura, Talpa, and Myogale), the teeth that are missing can be accurately defined. According to this view, when the number of incisors falls short of three it is assumed that the absent ones are missing from the outer and posterior end of the series. Thus, when there is but one incisor present, it is ¢1; when two, they are 11 and i2. Further- more, when the premolars and the molars are below their typical number, the absent teeth are missing from the fore part of the premolar series, and from the back part of the molar series. If this were invariably so, the labours of those who describe teeth 1 By many writers the letters indicating the different kinds of teeth are printed in capitals, as 7, C, P, and M; while very frequently the symbol Pm is employed in place of p. 26 GENERAL ANATOMICAL CHARACTERS would be greatly simplified ; but there are so many exceptions that a close scrutiny into the situation, relations, and development of a tooth is required before its nature can be determined, and in some cases the evidence at our disposal is scarcely sufficient for the purpose. In other instances, however, as among the Polyprotodont Marsupials, we have decisive evidence to show that the missing premolar teeth are not those at the extremity of the series. The milk-dentition is expressed by a similar formula, d for deciduous or m for milk being commonly prefixed to the Fia. 3.—Milk and Permanent Dentition of Upper (I.) and Lower (II.) Jaw of the Dog (Canis Jamiliaris), with the symbols by which the different teeth are commonly designated. The third upper molar (m.3) is the only tooth wanting in this animal to complete the typical heterodont mammalian dentition. letter expressive of the nature of the tooth. Since the three molars, and almost invariably the first premolar of the permanent series, have no predecessors, the typical milk-dentition would be expressed as follows—di 8, de 1,dm 8,=4, total 28. In a few Ungulates, however, such as the Hyrax and Tapir, and in some instances the Rhinoceros and the extinct Paleotheriwm, the whole of the four premolars are preceded by milk-teeth ; when we have the fullest development of cheek-teeth in the whole of the Eutheria. The teeth which precede the premolars of the permanent series are all called molars in the milk-dentition, although as a general rule, in DENTAL SYSTEM 27 form and function they represent in a condensed form the whole premolar and molar series of the adult. When there is a marked difference between the premolars and molars of the permanent dentition, the first milk-molar resembles a premolar, while the last has the characters of the posterior true molar. The dentition of all the members of the orders Primates, Carnivora, Insectivora, Chiroptera, and Ungulata can clearly be derived from the above-described generalised type. The same may be said of the Rodents, and even the Proboscidea, though at least in the existing members of the order with greater modi- fication. It is also apparent in certain extinct Cetacea, as Zeuglodon and Squalodon, but it is difficult to find any traces of it in existing Cetacea, Sirenia, or any of the so-called Edentata. All the Marsupials, different as they are in their general structure and mode of life, and variously modified as is their dentition, present in this system of organs some deep-lying common characters which show their unity of origin. The generalised type to which their dentition can be reduced presents considerable resemblance to that of the placental mammals, yet differing in details. It is markedly heterodont, and susceptible of division into incisors, canines, premolars, and molars upon the same principles. The whole number is, however, not limited to forty-four. The incisors may be as numerous as five on each side above, and they are almost always different in number in the upper and the lower jaw. The premolars and molars are commonly seven, as in the placental mammals, but their arrangement is reversed, as there are four true molars and three premolars. The larger number of incisive and molar teeth among the Marsupials suggests that their additional teeth have disappeared in the Eutheria,! and Mr. O. Thomas has endeavoured to construct a generalised dental formula from which both the Marsupial and Eutherian modifications may have been derived by the suppression of particular teeth. Thus the hypothetical formula pe 2 ig eg, ee by the loss of the fifth lower incisor, OTs # iy, 3, a* "1, 3, 8.4, BP and of the second premolars (which we know to be those which disappear in the Marsupials) and the fifth molars, will give .1,2,3,4,5 .1 10,34 1, 2,8, 4. L25,40 “Spa e se Opossum (Didelphys), usually written i 2, ¢4,p3,m 4. Again, in the same formula the loss of the fourth and fifth incisors in ; : oth 2 8 G0: both jaws, and also of the fourth molars, gives us 2 12, 3,0,0 °7’ 1, 2, 3, 4 1, 2, . A : L234 oh, or the formula of a typical Eutherian, like the ! According to Mr. G. E. Dobson there are four upper incisors in some of the Soricide. or the formula of the 28 GENERAL ANATOMICAL CHARACTERS Pig, which we generally write as 14, ¢+, p4, m%. Such a generalised formula will admit of modification into that of all existing, and a large number of fossil Marsupials, but it is possible that some of the Mesozoic types may have had more than four premolars, although there is no absolutely decisive evidence that such was the case. The presence of seven or eight true molars in some Mesozoic forms merely entails the addition of two or three additional figures to the ideal generalised formula. The milk-dentition of all known Marsupials, existing or extinct, is (if not entirely absent) limited to a single tooth on either side of each jaw, this being the predecessor of the last permanent premolar. And if the view that the milk-dentition is an additional series grafted upon the original permanent series be correct, it is evident that we have in this single replacement the first stage of this additional development. In very few mammals are teeth entirely absent. Even in the Whalebone Whales their germs are formed in the same manner and at the same period of life as in other mammals, and even become partially calcified, but they never rise above the gums, and completely disappear before the birth of the animal. In some species of the order Edentata, the true Anteaters and the Pangolins, no traces of teeth have been found at any age. The adult Monotremata are likewise devoid of teeth of the same structure as those of ordinary mammals; but well-developed molars occur in the young Ornithorhynchus, although no traces of teeth have hitherto been detected in Echidna. Modifications of the Teeth in Relation to their Functions.—The principal functional modifications noticed in the dentition of mammalia may be roughly grouped as piscivorous, carnivorous, insectivorous, omnivorous, and herbivorous, each having, of course, numerous variations and transitional conditions. The essential characters of a piscivorous dentition are best exemplified in the Dolphins, and also (as modifications of the carnivorous type) in the Seals. This type consists of an elongated, rather narrow mouth, wide gape, with numerous subequal, conical, sharp-pointed, recurved teeth, adapted simply to rapidly seize, but not to divide or masticate, active, slippery, but not powerful prey. All animals which feed on fish as a rule swallow and digest them entire, a process which the structure of prey of this nature, especially the intimate interblending of delicate, sharp-pointed bones with the muscles, renders very advantageous, and for which the above- described type of dentition is best adapted. The carnivorous type of dentition is shown in its most specialised development among existing mammals in the Felide. The function being here to seize and kill struggling animals, often of large size and great muscular power, the canines are immensely developed, DENTAL SYSTEM 29 trenchant, and piercing, and are situated wide apart, so as to give the firmest hold when fixed in the victim’s body. The jaws are as short as is consistent with the free action of the canines, so that no power may be lost. The incisors are very small, so as not to interfere with the penetrating action of the canines, and the crowns of the molar series are reduced to scissor-like blades, with which to pare off the soft tissues from the large bones, or to divide into small pieces the less dense portions of the bones for the sake of nutriment afforded by the blood and marrow they contain. The gradual modification between this and the two following types will be noticed in their appropriate places. In the most typical insectivorous animals, as the Hedgehogs and Shrews, the central incisors are elongated, pointed, and project forwards, those of the upper and lower jaw meeting like the blades of a pair of forceps, so as readily to secure small active prey, quick to elude capture, but powerless to resist when once seized. The crowns of the molars are covered with numerous sharp edges and points, which, working against each other, rapidly cut up the hard- cased insects into little pieces fit for swallowing and digestion. The omnivorous type, especially that adapted for the con- sumption of soft vegetable substances, such as fruits of various kinds, may be exemplified in the dentition of Man, of most Monkeys, and of the less modified Pigs. The incisors are moderate, subequal, and cutting. If the canines are enlarged, it is usually for other purposes than those connected with food, and only in the male sex. The molars have their crowns broad, flattened, and elevated into rounded tubercles. The name Bunodont, or hillock- toothed, has been proposed for molars of this type, and will frequently be found convenient. In the most typically herbivorous forms of dentition, as seen in the Horse and Kangaroo, the incisors are well developed, trenchant, and - adapted for cutting off the herbage on which the animals feed ; the canines are rudimentary or suppressed ; the molars are large, with broad crowns, which in the simplest forms have strong transverse ridges, but may become variously complicated in the higher degrees of modification which this type of tooth assumes. Various forms of teeth of this type will be noticed among the Ungulates and Rodents. The natural groups of mammals, or those which in our present state of knowledge we have reason to believe are truly related to each other, may each contain examples of more than one of these modifications. Thus the Primates have both omnivorous and insectivorous forms. The Carnivora show piscivorous, carnivorous, insectivorous, and omnivorous modifications of their common type of dentition. The Ungulata and the Rodentia have among them the omnivorous and various modifications, both simple and complex, 30 GENERAL ANATOMICAL CHARACTERS of the herbivorous type. The Marsupialia exhibit examples of all forms, except the purely piscivorous. Other orders, more restricted in number or in habits, as the Proboscidea and Cetacea, naturally do not show so great a variety in the dental structure of their members. Taxonomy.—In considering the taxonomic value to be assigned to the modifications of teeth of mammals, two principles, often opposed to each other, which have been at work in producing these modifications, must be held in view:—(1) the type, or ancestral form, as we generally now call it, characteristic of each group, which in most mammals is itself derived from the still more generalised type described above; and (2) variations which have taken place from this type, generally in accordance with special functions which the teeth are called upon to fulfil in particular cases. These variations are sometimes so great as completely to mask the primitive type, and in this way the dentition of many animals of widely different origin has come to present a remarkable superficial resemblance, as in the case of the Wombat (a Marsupial), the Aye-Aye (a Lemur), and the Rodents, or as in the case of the Thylacine and the Dog. In all these examples indications may generally be found of the true nature of the case by examining the earlier conditions of dentition; for the characters of the milk- teeth or the presence of rudimentary or deciduous members of the permanent set will generally indicate the route by which the specialised dentition of the adult has been derived. It is perhaps owing to the importance of the dental armature to the well-being of the animal in procuring its sustenance, and preserving its life from the attacks of enemies, that great changes appear to have taken place so readily, and with such comparative rapidity, in the forms of these organs—changes often accompanied with but little modification in the general structure of the animal. Of this proposition the Aye-Aye (Chiromys) among Lemurs, the Walrus among Seals, and the Narwhal among Dolphins form striking examples ; since in all these forms the superficial characters of their dentition would entirely separate them from the animals with which all other evidence (even including the mode of development of their teeth) proves their close affinity. Trituberculism.—Recent researches, and more especially those of Professors Cope and Osborn, tend to show that almost all of the extremely different forms of tooth-structure found among Mammals may be traced to one common type, in which the crown of each tooth carried three cusps, and hence termed the fritubercular type ; these three cusps being arranged in a triangle, with the apex directed inwardly in the upper teeth (Fig. 4, 6), and outwardly in the lower ones (Fig. 4, 7). It is further probable that this tritubercular type was itself derived from a type of dentition in DENTAL SYSTEM 31 which the teeth were in the form of almost a quite simple cone ; such a presumably primitive type of dentition being apparently retained among some existing Edentates, like the Armadillos, while it is possible that we should regard the dentition of the existing Cetacea (Fig. 2) as a reversion to the same primitive type. None of the Mesozoic mammals at present known exhibit this simple conical type of teeth, although we have an approximation to it in the extremely generalised genus Dromatherium. Starting then Fia. 4.—Molar teeth of Mesozoic Mammals(enlarged). Triconodont type—1, Dromatherium ; 2, Microconodon ; 8, Amphilestes ; 4, Phascolotheriwm ; 5, Triconodon. Tritubercular type—6, 7, Spalacotherium ; 10, Asthenodon. Tubercular sectorial type—8, Amphitheriwm ; 9, Peramus ; 11- 13, Amblotherium; 14 (2) Amblotherium. pr, Protocone ; hy, hypocone; pa, paracone; me, metacone, in the upper teeth; and protoconid, hypoconid, paraconid, and metaconid in the lower, 6 and 15 are upper molars, and the rest lower molars. (After Osborn.) from this presumed simple cone it appears that the teeth of Dromu- therium (Fig. 4,1) present the first stage towards trituberculism, the crown of each tooth having one main cone, with minute lateral cusps, and the root being grooved. In the next or true Tricon- odont stage (Fig. 4, 3.5) the crown has become elongated antero- posteriorly, and consists of one central and two lateral cones or cusps, while the root is divided. From this the transition is easy to the tritubercular type, in which the three cusps, instead of being placed in a line, are arranged in a triangle; the upper teeth (Fig. 32 GENERAL ANATOMICAL CHARACTERS 4, 6) having one inner and two outer cusps, while the reverse condition obtains in those of the lower jaw (Fig. 4, 7). These three cusps of the simple tritubercular tooth are collectively desig- nated as the. primitive triangle ; in the upper tooth the inner cusp is termed the protocone, the antero-external one the paracone, and the postero-external the metacone ; the corresponding cusps of the lower tooth being named protoconid, paraconid, and metaconid— the protoconid being here on the outer side of the crown. It is thus apparent that in the first, or haplodont type, as well as in the triconodont type, the upper and lower molars are alike ; while in the simple tritubercular type they have a similar pattern, — but with the arrangement of the cusps reversed. This simple tritubercular type occurs in the Mesozoic genus Spalacotherium (Fig. 4, 6 and 7), and apparently in the existing Chrysochloris ; but in the majority of tritubercular forms, while this primitive triangle forms the main portion of the crown, other secondary cusps are added, the homologies of which in the upper and lower teeth are somewhat doubtful. At the same time that we have the addition of these secondary cusps we also find trituberculism differentiating into a secodont and a bunodont series, according as to whether the dentition becomes of a cutting or a crushing type. Thus in the lower molars (Fig. 4, s and 9) we very frequently find the three cusps of the primitive triangle elevated and connected by cross crests, while there is an additional low posterior heel or talon, which may be termed the hypoconid. This tubercular- sectorial sub-type, as it is termed, is found in the lower molars of many Polyprotodont Marsupials and Insectivores, and it also occurs in the lower carnassial teeth of the true Carnivora. The presence of two cusps (inner and outer) to the talon con- verts this modification into a quinquetubercular form ; while, by the sup- pression of one of the three primitive cusps, it develops into the quadri- ‘Fic. 5.—Diagram of two upper and two lower left tubercular type of the quadritubercular molars in mutual apposition. The cusps . and ridges of the upper molars in double lines, and those bunodont serles. of the lower in black lines. The lower molars are looked In the upper molars at from below, asif transparent. pr, Protocone; hy,hypo- the primitive triangle in f=) cone ; pa, paracone ; me, metacone; ml, protoconule ; pl, ch metaconule ; prd, protoconid ; hyd, hypoconid ; pad, para- the secodont Series may conid ; med, metaconid ; end, entoconid. (After Osborn.) Temain purely tricuspid ¥ but the addition of in- termediate cusps, both in the secodont and bunodont series, may give rise to a quinquetubercular type; these intermediate cusps being respectively designated as the protoconule and metaconule (Fig. 5, THE SKELETON 33 ml, pl). Finally, in the bunodont series, the addition of a postero- internal cusp (Fig. 5, hy), termed the hypocone, forms the sextuber- cular molar. The following table exhibits, in a collective form, the names and relations of all the above-mentioned cusps, and the letters by which they are indicated in the figures :— Upper Monars. Antero-internal cusp =protocone =pr. Postero ,, or 6th cusp =hypocone =hy. Antero-external cusp =paracone =pa. Postero,, s =metacone =me. Anterior intermediate cusp = protoconule = ml. Posterior 35 oF =metaconule = pl. Lower Motars. Antero-external cusp =protoconid =prd. Postero ,, 5 =hypoconid =hyd. Antero-internal or 5th cusp =paraconid =pad. Intermediate (or in quadritubercular molars antero-internal) cusp. =metaconid =med. Postero-internal cusp =entaconid =end. The common occurrence of trituberculism in the mammals of the earlier geological epochs is, as remarked by Osborn, very significant of the uniformity of molar origin. Thus, among the Mesozoic mammals (with the exception of the group known as Multituberculata, in which the molars are constructed on a different type), trituberculism occurs in the great majority of the genera; while out of 82 species, belonging to five different suborders from the Lowest or Puerco Eocene of the United States, all but four exhibit this feature ; and the same holds good for the mammals of the corresponding European horizon. At the present day trituber- culism persists in the Lemuroidea, Insectivora, Carnivora, and Mar- supialia. In the Carnivora there is a tendency to lose the meta- conid, while in the bunodont molars of the Ungulata it is the paraconid that disappears. Ill. THE SKELETON, Definition.—The skeleton is a system of hard parts, forming a framework which supports and protects the softer organs and tissues of the body. It consists of dense fibrous and cartilaginous tissues, portions of which remain through life in this state, but the greater part is transformed during the growth of the animal into bone or osseous tissue. This is characterised by a peculiar 3 34 GENERAL ANATOMICAL CHARACTERS histological structure and chemical composition, being formed mainly of a gelatinous basis, strongly impregnated with salts of calcium, chiefly phosphate, and disposed in a definite manner, con- taining numerous minute nucleated spaces or cavities called lacune, connected together by delicate channels or canaliculi, which radiate in all directions from the sides of the lacunz. Parts composed of bone are, next to the teeth, the most imperishable of all the organs of the body, often retaining their exact form and internal structure for ages after every trace of all other portions of the organisation has completely disappeared, and thus, in the case of extinct animals, affording the only means of attaining a knowledge of their characters and affinities.? In the Armadillos and their extinct allies alone is there an. ossified exoskeleton, or bony covering developed in the skin. In , all other mammals the skeleton is completely internal. It may be described as consisting of an axial portion belonging to the head and trunk, and an appendicular portion belonging to the limbs. There are also certain bones called splanchnic, being developed within the substance of some of the viscera. Such are the os cordis and os penis found in some mammals. It is characteristic of all the larger bones of the mammalia that their ossification takes its origin from several distinct centres. One near the middle of the bone, and spreading throughout its greater portion, constitutes the diaphysis, or “shaft,” in the case of the long bones. Others near the extremities, or in projecting parts, form the epiphyses, which remain distinct during growth, but ultimately coalesce with the rest of the bone. Axial skeleton.—The axial skeleton consists of the skull, the vertebral column (prolonged at the posterior extremity into the tail), the sternum, and the ribs. Skull.—In the skull of adult mammals, all the bones, except the lower jaw, the auditory ossicles, and the bones of the hyoid arch, are immovably articulated together, their edges being in close con- tact, and often interlocking by means of fine denticulations project- ing from one bone and fitting into corresponding depressions of the other ; they are also held together by the investing periosteum, or fibrous membrane, which passes directly from one to the other, and permits no motion, beyond perhaps a slight yielding to external pressure. In old animals there is a great tendency for the different bones to become actually united by the extension of ossification from one to the other, with consequent obliteration of the sutures. 1 See for the principal modifications of the skeleton of the class, the large and beautifully illustrated Ostéographie of De Blainville, 1835-54; the section devoted to the subject in Bronn’s Klassen wnd Ordnungen des Thier-Reichs, by Giebel, 1874-79; and An Introduction to the Osteology of the Mammalia, by W. H. Flower, 3d ed., 1885. THE SKELETON 35 The cranium, thus formed of numerous originally independent ossifications, which may retain throughout life more or less of their individuality, or be all fused together, according to the species, the age, or even individual peculiarity, consists of a brain-case, or bony capsule for enclosing and protecting the brain, and a face for the support of the organs of sight, smell, and taste, and of those concerned in seizing and masticating the food. The brain-case articulates directly with the anterior cervical vertebra, by means of a pair of oval eminences, called condyles, placed on each side of the large median foramen which transmits the spinal cord. It consists of a basal axis, continuous serially with the axes or centra of the yn ” Fic. 6.—Longitudinal and vertical section of the skull of a Dog (Canis familiaris), with mandible and hyoid arch. an, Anterior narial aperture; MT, maxillo-turbinal bone ; /T, ethmo- turbinal; Na, nasal; ME, ossified portion of the mesethmoid; CZ, cribriform plate of the ethmo-turbinal: Fr, frontal; Pa, parietal; IP, interparietal; SO, supraoccipital; Ex0, ex- occipital; BO, basioccipital; Per, periotic; BS, basisphenoid; Pt, pterygoid; AS, ali- sphenoid; OS, orbitosphenoid; PS, presphenoid; Pl, palatine; VO, vomer; M.x, maxilla; PMzx, premaxilla; sh, stylohyal; eh, epihyal; ch, ceratohyal; bh, basihyal; th, thyrohyal ; s, symphysis of mandible; cp, coronoid process ; cd, condyle; a, angle; id, inferior dental canal. The mandible is displaced downwards, to show its entire form; the * indicates the part of the cranium to which the condyle is articulated.1 vertebrae, and of an arch above, roofing over and enclosing the cavity which contains the cephalic portion of the central nervous system (see Fig. 6). The base with its arch is composed of three segments placed one before the other, each of which is comparable to a vertebra with a greatly expanded neural arch. The hinder or 1 This and many of the following figures in this chapter are taken from Flower’s Osteology of the Mammalia. 36 GENERAL ANATOMICAL CHARACTERS occipital segment consists of the basioccipital, exoccipital, and supraoccipital bones; the middle segment of the basisphenoid, ali- sphenoid, and parietal bones; and the anterior segment of the presphenoid, orbitosphenoid, and frontal bones. The axis is continued forwards into the mesethmoid, or septum of the nose, around which the bones of the face are arranged in a manner so extremely modified for their special purposes that anatomists who have attempted to trace their serial homologies with the more simple portions of the axial skeleton have arrived at very diverse interpretations. The characteristic form and structure of the face of mammals is mainly dependent upon the size and shape of (1) the orbits, a pair of cup-shaped cavities for containing the eyeball and its muscles, which may be directed forwards or laterally, placed near together or wide apart, and may be completely or only partially encircled by bone; (2) the nasal fosse, or cavities on each side of the median nasal septum, forming the passage for the air to pass between the external and the internal nares, and containing in their upper part the organ of smell; (3) the zygomatic arch, a bridge of bone for the purpose of muscular attachment, which extends from the side of the face to the skull, overarching the temporal fossa ; (4) the roof of the mouth, with its alveolar margin for the implanta- tion of the upper teeth. The face is completed by the mandible, or lower jaw, consisting of two lateral rami, articulated by a hinge joint with the squamosal (a cranial bone interposed between the posterior and penultimate segment of the brain-case, where also the bony capsule of the organ of hearing is placed), each being composed of a single solid piece of bone, and the two united together in the middle line in front, at the symphysis,—which union may be per- manently ligamentous or become completely ossified. Into the upper border of the mandibular rami the lower teeth are implanted. In addition to the bones already mentioned as entering into the formation of the cranium, there are many others, the most import- ant of which may be briefly noticed. The anterior extremity of the skull is formed by the premaxille (Figs. 6, 7, PJZc), which carry the incisors; behind them are the maxille, in which all the remaining upper teeth are implanted. Both the premaxille and maxille meet in a median suture on the palate, where they form a floor to the nasal passage ; this floor being continued backwards by the plate-like pala- tines, at the hinder extremity of which the posterior nares are usually situated. In a few instances, however, as in certain Edentates and Cetaceans, the small pair of bones forming the posterior continuation of the lateral borders of the palatines, and known as the pterygoids (Fig. 6, Pt), likewise meet in the middle line below the nasal passage, and thus cause the aperture of the posterior nares to be situated near the occiput. On the upper, or frontal aspect of the cranium the paired nasals roof over the nasal passage and fill the interval left THE SKELETON 37 between the premaxilla and maxilla of either side. Behind the nasals and maxille, the anterior part of the brain-case is formed by the large paired frontals (Figs. 6, 7, /r), behind which are the parietals, which may be of still larger size, and form the greater part of the brain-case. z475.1 Heart.—The heart of mammals consists of four distinct cavities, two auricles and two ventricles. Usually the ventricular portion is externally of conical form, with a simple apex, but in the Sirenia it is broad and flattened, and a deep notch separates the apical portion of each ventricle. A tendency to this form is seen in the Cetacea and the Seals. It is characteristic of mammals alone among verte- brates that the right auriculo-ventricular valve is tendinous like the left, consisting of flaps held in their place by fibrous ends (chorde tendinie) and arising from projections of the muscular walls of the ventricular cavity (musculi papillares). In the Monotremata a transition between this condition and the simple muscular flap of the Sauropsida is observed. In most of the larger Ungulates a dis- tinct but rather irregular ossification (0s cordis) is developed in the central tendinous portion of the base of the heart. Blood-vessels,—The orifices of the aorta and pulmonary artery are 1G. Gulliver, Proc. Zool. Soc., 1862, p. 91. 64 GENERAL ANATOIMHCAL CHARACTERS each guarded by three semilunar valves. ‘Tho aorta is single, and arches over tho left bronchial tube. After supplying tho tissues of the hoart itself with blood by means of tho coronary arterios, it gives off largo vessels (“carotid”) to the head and (“brachial”) to the anterior extremities. Tho mode in which these vessols arise from the aorta varies much in different mammals, and tho study of their disposition affords somo guide to classification, In nearly all cases the right brachial and carotid have a common origin (called the “innominate artery” in anthropotomy). The other two vessels may come off from this, as is the rule in Ungulates, the common trunk constituting tho “anterior aorta” of veterinary anatomy ; or they may be detached in various degrees, both arising separately from the aorta, as in Man, or tho left carotid from the innominate and the left brachial from the aorta, a vory common arrangement ; or tho last two from a common second or left innominate, as in some Bats and Insectivores. Tho aorta, after giving off the inter- costal arteries, passes through tho diaphragm into the abdomen, and, after supplying the viscora of that cavity by moans of tho gastric, hepatic, splenic, mesontoric, renal, and spermatic vessels, gives off in the lumbar region a large branch (iliac) to each of tho hindor extremities, which also supplies the polvic viscera, and is continued onwards in tho middlo line, greatly diminished in sizo, along the under surface of tho tail as the caudal artery. In certain mammals, arterial ploxusos, called retin mirahilia, formed by the breaking up of the vessel into an immense number of small trunks, which may run in a straight course parallel to one another (as in the limbs of Sloths and Slow Lemurs), or form a closely packed network, as in the intracranial plexuses of Ruminants, or a sponge-like mass of convoluted vessels, as in the intercostals of Cetaceans, are peculiarities of tho vascular system the meaning of which is not in all cases clearly understood. In the Cetacea they aro ob- viously receptacles for containing a largo quantity of oxygenated blood available during the prolonged immersion, with consequent absence of respiration, to which those animals are subject. The vessols returning the blood to the heart from the head and upper extremities usually unite, as in Man, to form tho single vena curd superior or procaval voin, but in somo Inscctivores, Chiroptora, and Rodents, in the Elophant, and all Marsupials and Monotromos the two suporior caval veins enter tho right auricle without uniting, as in birds. In Scals and somo other diving mammals there is a large venous sinus or dilatation of the inforior vena cava immediately below tho diaphragm, — In the Cotacoa the purposo of this is supplied by tho immenso abdominal venous ploxusos. As a rule tho voins of mammals aro furnished with valves, but those are said to he utogothor wanting in the Cetacoa, and in the suporior and inferior cava, subclavian and ilive veins, the veins of the liver (both portal ABSORBENT SYSTEM. 65 and hepatic), heart, lungs, kidneys, brain, and spinal cord of other mammals. Many of the veins within the cranium are included in spaces formed by the separation of the lamin of the dura mater, and do not admit of being dilated beyond a certain size; these are termed sinuses. The portal circulation in mammals is limited to the liver, the portal vein being formed by the superior and inferior mesenteric, the splenic, the gastro-epiploic, and the pancreatic veins. The kidney is supplied solely by arterial blood, and its veins empty their contents only into the inferior cava. Lymphatic Vessels.—The absorbent or lymphatic system of vessels is very fully developed in the Mammalia. Its ramifications extend through all the soft tissues of the body, and convey a colourless fluid called lymph, containing nucleated corpuscles, and also, during the process of digestion, the chyle, a milky fluid taken up by the lymphatics (here called lacteals) of the small intestine, and pour them into the general vascular system, where they mix with the venous blood. The lymphatic vessels of the hinder extremities, as well as those from the intestinal canal, unite in the abdomen to form the “thoracic duct,” the hinder end or commencement of which has a dilatation called the receptaculum chyli. This duct, which is of irregular size and sometimes double, often dividing and uniting again in its course, or even becoming plexiform, passes for- wards close to the bodies of the thoracic vertebre, and empties itself, by an orifice guarded by a valve, into the great left brachio-cephalic vein, having previously received the lymphatics from the thorax and the left side of the head and left anterior extremity. The lymph- atics from the right side of the head and right anterior limb usually enter by a small distinct trunk into the corresponding part of the right brachio-cephalic vein. The duct, and also the principal lymph- atic vessels, are provided with valves. Lymphatic glands, rarely met with in the Sauropsida, are usually present in mammals, both in the general and in the lacteal system ; the latter being called “mesenteric glands.” They are round or oval masses, situated upon the course of the vessels, which break up in them and assume a plexiform arrangement, and then reunite as they emerge. No structures corresponding to the pulsating “lymphatic hearts” of the lower vertebrates have been met with in mammals. Ductless Glands,—Associated with the vascular and lymphatic systems are certain bodies (the functions of which are not properly understood), usually, on account of their general appearance, grouped together under the name of “ductless glands.” The largest of these is the “spleen,” which is single, and always placed in mammals in relation to the fundus or left end of the stomach, to which it is attached by a fold of peritoneum. It is dark- coloured and spongy in substance, and has a depression or “hilus” 5 66 GENERAL ANATOMICAL CHARACTERS on one side, into which the splenic artery, a branch of the cceliac axis of the abdominal aorta, enters, and from which the vein joining the portal system emerges. The spleen varies much in size and form in different mammals, being relatively very small in the Cetacea. It is sometimes almost spherical, but more often flattened, oval, triangular, or elongated, and occasionally, as in Monotremes and most Marsupials, triradiate. The “suprarenal bodies” or “adrenals” are two in number, each situated either in contact with, or at a short distance in front of the anterior extremity of the kidney. They are abundantly supplied with nerves, and are much larger re- latively in early than in adult life. The “thyroid bodies,” of which there are generally two, though in Man and some other species they are connected by an isthmus passing across the middle line, are constant in mammals, though only met with in a rudimentary condition, if at all, in other vertebrates. They are situated in the neck, in contact with the sides of the anterior extremity of the trachea. The “thymus” lies in the anterior part of the thorax, between the sternum and the great vessels at the base of the heart, and differs from the thyroid in being median and single, and having a central cavity. It attains its greatest development during the period in which the animal is nourished by its mother’s milk, and then it diminishes, and generally disappears before full growth is attained. Nostrils— Mammals breathe occasionally through the mouth, but usually, and in many cases exclusively, through the nostrils or nares. These are apertures, always paired (except in the toothed Cetacea, where they unite to form a single external opening), and situated at the fore part of the face, generally at or beneath the end of the muzzle, a median prominence above the mouth. This is sometimes elongated to form a proboscis, to the extremity of which the nostrils are carried, and which attains its maximum of develop- ment in the Elephant. In the Cetacea the nostrils are situated at a considerable distance behind the anterior end of the face, upon the highest part of the head, and are called “blow-holes,” from the peculiar mode of respiration of those animals. The nostrils are kept open by means of cartilages surrounding their aperture, which many animals have the power of moving so as to cause partial dilatation or contraction. Jn diving animals, as Seals and Cetacea, they can be completely closed at will so as to prevent the entrance of water when beneath the surface. The passage to which the nostrils lead is in most mammals filled by a more or less complex sieve-like apparatus, formed of the convoluted turbinal bones and cartilages, over which a moist, vascular, ciliated mucous membrane is spread, which intercepts particles of dust, and also aids in warming the inspired air before it reaches the lungs. In the Proboscidea, in which these functions are performed by RESPIRATORY SYSTEM 67 the walls of the long tubular proboscis, this apparatus is entirely wanting. Lravhea.—The narial passages have the organ of smell situated in their upper part, and communicate posteriorly with the pharynx, and through the glottis with the “trachea” or windpipe. a tube by which the air is conveyed to and from the lungs. The permanent patency of the trachea during the varied movements of the neck is provided for by its walls being stiffened by a series of cartilaginous rings or hoops, which in most mammals are incomplete behind. Having entered the thorax, the trachea bifurcates into the two bronchi, one of which enters. and, dividing dichotomously. ramifies through each lung. In some of the Cetacea and Artiodactyla a third bronchus is given off from the lower part of the trachea, above its bifureation, and enters the right lung. Larynzy—The upper end of the trachea is modified into the organ of voice or “larynx,” the air passing through which to and from the lungs is made use of to set the edges of the “vocal cords,” or fibrous bands stretched one on each side of the tube, into vibra- tion. The larynx is composed of several cartilages, such as the “thyroid,” the “cricoid,” and the “arytenoid” which are moved upon one another by muscles, and suspended from the hyoidean arch. By alteration of the relative position of these cartilages the cords can be tightened or relaxed, approximated or divaricated, as required to modulate the tone and volume of the voice. A median tongue-shaped fibro-cartilage at the top of the larynx. the “epiglottis,” protects the “ glottis.” or aperture by which the larynx communi- cates with the pharynx, from the entry of particles of food during deglutition. The form of the larynx and development of the vocal cords present many variations in different members of the class. the greatest modification from the ordinary type being met with in the Cetacea, where the arytenoid cartilages and epiglottis are united in a tubular manner, so as to project into the nasal passage, and, being grasped by the muscular posterior margin of the palate, pro- vide a direct channel of communication from the lungs to the external surface. An approach to this condition is met with in the Hippopotamus and some other Ungulates: it is indeed so general as an abnormality, that Howes suggests that an internarial epi- glottis may have been a primitive feature common throughout the class. Nearly all mammals have a voice, although sometimes it is only exercised at seasons of sexual excitement. Some Marsupials and Edentates appear to be quite mute. In no mammal is there an inferior larynx, or “ syrinx,” as in birds. Diaphragm.—tThe thoracie cavity of mammals differs from that of the Sauropsida in being completely separated from the abdomen by a muscular partition, the “diaphragm,” attached to the vertebral 68 GENERAL ANATOMICAL CHARACTERS column, the ribs, and the sternum. This is much arched, with the convexity towards the thorax, so that when its fibres contract and it is flattened the cavity of the thorax is increased, and when they are relaxed the cavity is diminished. LIungs.—The lungs are suspended freely in the thorax, one on each side of the heart, being attached only by the root, which consists of the bronchus or air-tube and pulmonary arteries and veins by which the blood is passed backwards and forwards between the heart and the lungs. The remaining part of the surface of each lung is covered by serous membrane, the “pleura”; and what- ever the state of distension or contraction of the chest-wall, is accurately in contact with it. Inspiration is effected by the con- traction of the diaphragm and by the intercostal and other muscles elevating or bringing forward the ribs, and thus throwing the sternum farther away from the vertebral column. As the surface of the lung must follow the chest-wall, the organ itself is expanded, and air rushes in through the trachea to fill all the minute cells in which the ultimate ramifications of the bronchi terminate. In ordinary expiration very little muscular power is expended, the elasticity of the lungs and surrounding parts being sufficient to cause a state of contraction and thus drive out at least a portion of the air contained in the cells, when the muscular stimulus is with- drawn. The lungs are sometimes simple externally, as in the Sirenia (where they are greatly elongated) and the Cetacea, but are more often divided by deep fissures into one or more lobes. The right lung is usually larger and more subdivided than the left. It often has a small distinct lobe behind, wanting on the left side, and hence called lobulus azygos. Air-sacs. —Most mammals have inconnection with the air passages certain diverticuli or pouches containing air, the use of which is not always easy to divine. The numerous air sinuses situated between the outer and inner tables of the bones of the head, represented in Man by the antrum of Highmore and the frontal and sphenoidal sinuses, and attaining their maximum of development in the Indian Elephant, are obviously for the mechanical purpose of allowing expansion of the osseous surface without increase of weight. They are connected with the nasal passages. The Eusta- chian tubes pass from the back of the pharynx to the cavity of the tympanum, into which and the mastoid cells they allow air to pass. In the Hquide there are large post-pharyngeal air-sacs in connection with them. The Dolphins have an exceedingly complicated system of air-sacs in connection with the nasal passages just within the nostrils, and the Tapirs, Rhinoceroses, and Horses have blind sacs in the same situation. In the males of some Seals (Cystophora and Macerorhinus) large pouches, which the animal can inflate with air and which are not developed in the young animal or the female, URINARY SYSTEM _.. 69 arise from the upper part of the nasal passages, and lie immediately under the skin of the face. These appear analogous, although not in the same situation, to the gular pouch of the male Bustard. The larynx frequently has membranous pouches in connection with it, into which air passes. These may be lateral and opening just above the vocal cords, when they constitute the sacculi laryngis, found -in a rudimentary state in Man, and attaining an enormous development, so as to reach to the shoulders and axille, in some of the Anthropoid Apes; or they may be median, opening in front either above or below the thyroid and cricoid cartilages, as in the Howling and other Monkeys, and also in the Whalebone Whales and Great Anteater. Urinary Organs.—The kidneys of mammals are more compact and definite in form than in other vertebrates, being usually more or less oval, with an indent on the side turned towards the middle line, from and into which the vessels and ducts pass. They are distinctly divided into a cortical secretory portion, composed mainly of convoluted tubes, and containing the so-called Malpighian bodies ; and a medullary excreting portion, formed of straight tubes converging towards a papilla, embraced by the commencement of the ureter or duct of the organ. The kidneys of some mammals, as most Monkeys, Carnivores, Rodents, etc, are simple, with a single papilla into which all the renal tubuli enter. In others, as Man, there are many pyramids of the medullary portion, each with its papilla, opening into a division (calyx) of the upper end of the ureter. Such kidneys, either in the embryonic condition only, or throughout life, are lobulated on the surface. In some cases, as in Bears, Seals, and especially the Cetacea, the lobulation is carried further, the whole organ being composed of a mass of renules, loosely united by connective tissue, and with separate ducts, which soon join to form the common ureter. Bladder.—In all mammals except the Monotremes the ureters terminate by slit-like valvular openings in the urinary bladder. This receptacle when filled discharges its contents through the single median urethra, which in the male is almost invariably included in the penis, and in the females of some species of Rodents, Insectivores, and Lemurs has a similar relation to the clitoris. In the Monotremes, though the bladder is present, the ureters do not enter into it, but join the urino-genital canal some distance below ‘it, with the orifice of the genital duct intervening. VI. NERVOUS SYSTEM AND ORGANS OF SENSE. Brain.—The brain of mammals shows a higher condition of organisation than that of other vertebrates. The cerebral hemi- 7O GENERAL ANATOMICAL CHARACTERS spheres have a greater preponderance compared with other parts, especially to the so-called optic lobes, or corpora quadrigemina, which are completely concealed by them. The commissural system of the hemispheres is much more complex, both fornix and corpus callosum being present in some form; and when the latter is rudimentary, as in Marsupials and Monotremes, its deficiency is made up for by the great size of the anterior commissure, The lateral lobes of the cerebellum, wanting in lower vertebrates, are well developed and connected by a transverse commissure, the pons Varolii. The whole brain, owing especially to the size of the cerebral hemispheres, is considerably larger relatively to the bulk of the animal than in other classes, but it must be recollected that the size of its brain depends upon many circumstances besides the degree of intelligence which an animal possesses, although this is certainly one. Man’s brain is many times larger than that of all other known mammals of equal bulk, and even three times as large as that of the most nearly allied Ape. Equal bulk of body is here mentioned, because, in drawing any conclusions from the size of the brain compared with that of the entire animal, it is always necessary to take into consideration the fact that in every natural group of closely allied animals the larger species have much smaller brains relatively to their general size than the smaller species, so that, in making any effective comparison among animals belonging to different groups, species of the same'size must be selected. It may be true that the brain of a Mouse is, as compared with the size of its body, larger than that of a Man, but, if it were possible to reduce an animal having the general organisation of a Man to the size of a Mouse, its brain would doubtless be very many times larger ; and conversely, as shown by the rapid diminution of the relative size of the brain in all the large members of the Rodent order, a Mouse magnified to the size of a Man would, if the general rule were observed, have a brain exceedingly inferior in volume. Al- though the brain of the large species of Whales is, as commonly stated, the smallest in proportion to the bulk of the animal of any mammal, this does not invalidate the general proposition that the Cetacea have very large brains compared with terrestrial mammals, like the Ungulata, or even the aquatic Sirenia, as may be proved by placing the brain of a Dolphin by the side of that of a Sheep, a Pig, or a Manatee of equal general weight. It is only because the universally observed difference between the slower ratio of increase of the brain compared with that of the body becomes so enormous in these immense creatures that they are accredited with small brains. The presence or absence of “sulci” or fissures on the surface of the hemisphere, dividing it into “convolutions” or “ gyri,” and thus increasing the superficies of the cortical gray matter, as well NERVOUS SYSTEM 71 as allowing the pia mater with its nutrient blood-vessels to pene- trate into the cerebral substance, follow somewhat similar rules. The sulci are related partly to the high or low condition of organis- ation of the species, but also in a great degree to the size of the cerebral hemispheres. In very small species of all groups, even the Primates, they are absent, and in the largest species of groups so low in the scale as the Mar- supials and Edentates they are found. They reach their maximum of development in the Cetacea. The accompanying wood- cut (Fig. 23) shows the prin- cipal parts of a mammalian brain, as seen from the superior, lateral, and inner surfaces. The sylvian fissure (sf) is one of the most con- stant of the sulci found in the hemispheres. The researches of Pale- ontologists, founded upon studies of casts of the in- terior of the cranial cavity of extinct forms, have shown that, in many natural groups of mammals, if not in all, the brain has increased in size, and also in complexity of surface foldings, with the Fic. 23.—Brain of the Genet (Genetta tigrina). A, From above; B, from the right side; C, inner sur- face of right hemisphere; cc, corpus callosum ; e.m.s, calloso-marginal sulcus; ¢, notch represent- ing crucial sulcus of other forms; d, depression on superior lateral gyrus of hemisphere; hg, hippo- campal gyrus; i, inferior lateral gyrus of hemi- sphere; m, middle lateral gyrus of do. ; s, superior advance of time,—indicating in this, as in so many other respects, a gradual progress lateral gyrus of do.; 03, supraorbital sulcus of do. ; sf, sylvian fissure of do. ; ol, olfactory lobes. The deeply convoluted part behind the cerebral hemi- sphere is the cerebellum, below which lies the medulla oblongata, or commencement of the spinal f igh rom a lower to a hig er type cord. (Mivart, Proc. Zool. Soc. 1882, p. 516.) of development. Nerves.—The twelve pairs of cranial nerves generally recognised in vertebrates are usually all found in mammals, though the olfactory nerves are excessively rudimentary, if not altogether absent, in the Toothed Whales. The spinal cord, or continuation of the central nervous axis, lies in the canal formed by the neural arches of the vertebra, and gives off the compound double-rooted nerves of the trunk and the extremities, corresponding in number to the vertebra, through the interspaces between which they pass 72 GENERAL ANATOMICAL CHARACTERS out to their destination. The cord is somewhat enlarged at the two points where it gives off the great nerves to the anterior and the posterior extremities, which, from their interlacements soon after their origin, are called respectively the brachial and lumbar plexuses. The ganglionic or sympathetic portion of the nervous system is well developed, and presents few modifications. Sense of Touch.—The sense of touch is situated in the skin generally, but is most acute in certain regions more or less specialised for the purpose by the presence of tactile papille, such as portions of the face, especially the lips and end of the snout, and the extremities of the limbs when these are used for other purposes than mere progression, and the under surface of the end of the tail in some Monkeys. The “vibrisse” or long stiff bristles situated on the face of many mammals are rendered extremely sensitive to touch by the abundant supply of branches from the fifth nerve to their basal papilla. In Bats the extended wing membranes, and probably also the large ears and the folds and prominences of skin about the face of some species, are so sensitive as to receive impressions even from the different degrees of resistance of the air, and so enable the animals to avoid coming in contact with obstacles | to their nocturnal flight. Taste and Smell.—The organs of the other special senses are confined to the head. Taste is situated in the papille scattered on the dorsal surface of the tongue. The organ of smell is present in all mammals except the Toothed Whales. It consists of a ramifica- tion of the olfactory nerves over a plicated, moist, mucous membrane, supported by folded plates of bone, placed on each side of the septum nasi in the roof, or often in a partially distinct upper chamber, of the nasal passage, so arranged that, of the air passing into the lungs in inspiration, some comes in contact with it, causing the perception of any odorous particles with which it may be charged. Many mammals possess intense powers of smelling certain odours which others are quite unable to appreciate, and the influence which this sense exercises over the well-being of many species is very great, especially in indicating the proximity of others of the same kind, and giving warning of the approach of enemies. The development and modification of the sense of smell is probably associated with that of the odorous secretion of the cutaneous glands. Sight.—The organ of sight is quite rudimentary, and even concealed beneath the integument, in some burrowing Rodents and Insectivores, and is most imperfectly developed in the Platanista, or Freshwater Dolphin of the rivers of India. In all other mammals the eyeball has the structure characteristic of the organ in the higher Vertebrata, consisting of parts through which the rays of light are admitted, regulated, and concentrated upon the sensitive ORGANS OF SENSE 73 expansion of the optic nerve lining the posterior part of the ball. A portion of the fibro-vascular and highly pigmented layer, the choroid, which is interposed between the retina and the outer sclerotic coat, is in many mammals modified into a brilliantly- coloured light-reflecting surface, the tapetum lucidum. There is never a pecten or marsupium like that of the Sauropsida, nor is the sclerotic ever supported by a ring of flattened ossicles, as is so frequently the case in the lower vertebrated classes. The eyeball is moved in various directions by a series of muscles—the four straight, two oblique, and, except in the higher Primates, a pos- terior retractor muscle called choanoid. The superior oblique muscle passes through a tendinous pulley fastened to the roof of the orbit, which is a feature not found beyond the limits of the mammalian class. The eye is protected by the lids, generally distinctly separated into an upper and a lower movable flap, which, when closed, meet over the front of the eye in a more or less nearly horizontal line ; but sometimes, as in the Sirenia, the lids are not distinct, and the aperture is circular, closing to a point. In almost all mammals below the Primates, except the Cetacea, a “nictitating membrane” or third eyelid is placed at the inner corner of the eyeball, and works horizontally across the front of the ball within the true lids. Its action is instantaneous, being apparently for the purpose of cleaning the front of the transparent cornea ;—a function unneces- sary in animals whose eyes are habitually bathed in water, and which in Man and his nearest allies is performed by winking the true eyelids. Except in Cetacea the surface of the eye is kept moist by the secretion of the lachrymal gland, placed under the upper lid at its outer side, and the lids are lubricated by the Harderian and Meibomian glands, the former being situated at the inner side of the orbit, and especially related to the nictitating membrane, the latter in the lining membrane of the lids. Hearing.—The organ of hearing is inclosed in a bony capsule (periotic) situated in the side of the head, intercalated between the posterior (occipital) and the penultimate (parietal) segment of the skull. It has, in common with other vertebrates, three semicircular canals and a vestibule, but the cochlea is more fully developed than in the Sauropsida, and, except in the Monotremes, spirally con- voluted. The tympanic cavity is often dilated below, forming a smooth rounded prominence on the base of the skull, the auditory bulla (Fig. 8). The three principal ossicles, the “malleus,” “ incus,” and “stapes,” are always present, but variable in characters. In the Sirenia, Cetacea, and Seals they are massive in form, being in the first-named order of larger size than in any other mammals. In the Cetacea the malleus is ankylosed to the tympanic ; but in other mammals it is connected only with the membrana tympani. The stapes in the lower orders—LEdentates, Marsupials, and Monotremes 74 GENERAL ANATOMICAL CHARACTERS —has a great tendency to assume the columnar form of the corresponding bone in Sauropsida, its two rami entirely or partially coalescing.1 The tympanic membrane (drum of the ear) forms the outer wall of the cavity. In the fcetal state it is level with the external surface of the skull, and remains so permanently in a few mammals, as the American Monkeys ; but commonly, by the growth of the squamosal bone, it becomes deeply buried at the bottom of a bony tube (meatus auditorus eaternus), which is continued to the sur- face of the skin in a fibrous or fibro-cartilaginous form. In Whales, owing to the thickness of the subcutaneous adipose tissue, this meatus is of great length, and is also extremely narrow. In most aquatic and burrowing animals it opens upon the surface by a simple aperture, but in the large majority of the class there is a projecting fold of skin, strengthened by fibro-cartilages, called the pinna, auricle, or “external ear,” of very variable size and shape, generally movably articulated on the skull, and provided with muscles to vary its position ; this pinna helping to collect and direct the vibra- tions of sound into the meatus. VII. REPRODUCTIVE ORGANS. Testes.—In the male the testes retain nearly their primitive or internal position throughout life in the Monotremata, Sirenia, Cetacea, most Edentata, Hyracoidea, Proboscidea, and Seals, but in other groups they either periodically (as in Rodentia, Insectivora, and Chiroptera) or permanently pass out of the abdominal cavity through the inguinal canal, forming a projection beneath the skin of the perineum, or becoming suspended in a distinct pouch of integument called the scrotum. All the Marsupials have a pedunculated scrotum, the position of which differs from that of other mammals, being in front of, instead of behind, the preputial orifice. As regards the presence, absence, or comparative size and number of the accessory generative glands—prostate, vesi- cular, and Cowper’s glands, as they are called—there is much variation in different groups of mammals. Penis.—The penis is almost always completely developed, consisting of two corpora cavernosa attached to the ischial bones, and of a median corpus spongiosum enclosing the urethra, and forming the glans at the distal portion of the organ. In Marsupials, Monotremes, and the Sloths and Anteaters, the corpora cavernosa are not attached directly to the ischia, and in the last-named the penis is otherwise of a very rudimentary character, the corpus 1 The modifications of these bones are fully described by A, Doran, ‘‘ Morpho- logy of the Mammalian Ossicula auditus,” Trans. Linn. Soc. ser. 2, vol. i. pp. 371-497, pl. lviii.-lxiv. (1878). REPRODUCTIVE ORGANS 75 spongiosum not being present. In many Marsupials the glans penis is bifurcated. In most Primates, Carnivora, Rodentia, Insectivora, and Chiroptera, but in no other orders, an os penis is present. Ovaries and Oviduct.—In the female, the ovaries permanently retain their original abdominal position, or only descend a short distance into the pelvis. They are of comparatively smaller size than in other vertebrates, have a definite flattened oval form, and are enclosed in a more or less firm “tunica albigenia.” The oviduct has a trumpet-like, and usually fimbriated abdominal aperture, and is more or less differentiated into three portions :—(1) a contracted upper part, called in Man and the higher mammals the “ Fallopian tube”; (2) an expanded part with muscular walls, in which the ovum undergoes the changes by which it is developed into the foetus, called the “uterus”; (3) a canal, the “vagina,” separated from the last by a valvular aperture, and terminating in the urino- genital canal, or common urinal and genital passage, which in higher mammals is so short as scarcely to be distinct from the vagina. The complete distinction of the oviducts of the two sides through- out their whole length, found in all lower vertebrates, only occurs in this class in Monotremes ; a prevailing mammalian characteristic being their more or less perfect coalescence in the middle line to form a single median canal. In the Marsupials this union only includes the lower part of the vagina ; but in most Placentals it extends to the whole vagina and a certain portion of the uterus, which cavity is then described as “bicornuate.” In the higher mammals, as in Man, and also in some of the Edentates, the whole of the uterus is single, the contracted upper portion of the oviducts or Fallopian tubes, as they are then called, entering its upper lateral angles by small apertures. In certain lower forms the urino-genital canal opens with the termination of the rectum into a common cloaca, as in other vertebrates; but it is characteristic of the majority of the class that the two orifices are more or less distinct exter- nally. Mammary Glands. —Mammary glands secreting the milk by which the young are nourished during the first portion of their existence after birth, are present in both sexes in all mammals, though usually only functional in the female. In the Monotremes alone their orifices are mere scattered pores in the skin, but in all other forms they are situated upon the end of conical elevations, called mammille or teats, which, taken into the mouth of the young animal, facilitate the process of sucking. These are always placed in pairs upon some part of the ventral surface of the body, but vary greatly in number and position in different groups. In the Cetacea, where the prolonged action of sucking would be incom- patible with their subaqueous life, the ducts of the glands are dilated into large reservoirs from which the contents are injected 76 GENERAL ANATOMICAL CHARACTERS into the mouth of the young animal by the action of a compressor muscle. Secondary Seawal Characters—Secondary sexual characters, or modifications of structure peculiar to one sex, but not directly related to the reproductive function, are very general in mammals. They almost always consist of the acquisition or perfection of some character by the male as it attains maturity, which is not found in the female or the young in either sex. In a large number of cases these clearly relate to the combats in which the males of many species engage for the possession of the females during the breeding season; others are apparently ornamental, and of many it is still difficult to apprehend the meaning. Many suggestions on this subject will, however, be found in the chapters devoted to it in Darwin’s work on The Descent of Man and Selection in Relation to Ses, where most of the best-known instances are collected. Superiority of size and strength in the male of many species is a well- marked secondary sexual character related to the purpose indicated above, being probably perpetuated by the survivors or victors in combats transmitting to their descendants those qualities which gave them advantages over others of their kind. To the same category belong the great development of the canine teeth of the males of many species which do not use these organs in procuring their food, as the Apes, Swine, Musk and some other Deer, the tusk of the male Narwhal, the antlers of Deer, which are present in most cases only in the males, and the usual superiority in size and strength of the horns of the Bovidw. Other secondary sexual characters, the use of which is not so obvious, or which may only relate to ornament, are the presence of masses or tufts of long hair on different parts of the body, as the mane of the male Lion and Bison, the beards of some Ruminants and Bats (as Taphozous melano- pogon), Monkeys, and of Man, and all the variations of coloration in the sexes, in which, as a general rule, the adult male is darker and more vividly coloured than the female. Here may also be mentioned the presence or the greater development of odoriferous glands in the male, as in the Musk Deer, and the remarkable perforated spur with its glands and duct, so like the poison-tooth of the venomous serpents, found in the males of both Ornithorhynchus and Echidna, the use of which is at present unknown. Placenta.—The development of the mammalian ovum, and the changes which the various tissues and organs of the body undergo in the process of growth, are too intricate subjects to be explained without entering into details incompatible with the limits of this work, especially as they scarcely differ, excepting in their later stages, from those of other vertebrates, upon which, owing to the greater facilities these present for examination and study, the subject has been more fully worked out. There are, however, REPRODUCTIVE ORGANS 77 some points which require notice, as peculiar to the mammalian class, and as affording at least some hints upon the difficult subject of the affinities and classification of the members of the group. The nourishment of the fetus during intra-uterine life takes place through the medium of certain structures, partly belonging to the foetus itself and partly belonging to the inner parietes of the uterus of the parent. These in their complete form constitute the complex organ called the “placenta,” serving as the medium of communication between the mother and foetus, and in which the physiological processes that are concerned in the nutrition of the latter take place; but, as we shall see, though a placenta, in the usual acceptation of the term, is peculiar to the mammalian class, it is not in all of its members that one is developed. The structures to which we shall have especially to refer are the outer tunic of the ovum, to which, however formed, the term “chorion” is commonly applied, and two sac-like organs connected with the body-cavity of the embryo, both formed from the splanchnic mesoblast, lined by a layer of the hypoblast. These are the “umbilical vesicle” or “ yolk- sac” and the “allantois.” The umbilical vesicle is a thin membrane enclosing the yolk, which by the doubling in of the ventral walls of the embryo becomes gradually formed into a distinct sac external to the body, with a pedicle (the omphalo-enteric duct) by which for a time a communica- tion is maintained between its cavity and the intestinal canal. In the walls of this sac blood-vessels (omphalo-meseraic or vitelline) are developed in connection with the vascular system of the embryo, through which, either by their contact with the outer surface of the walls of the ovum, or by the absorption through them of the contents of the yolk-sac, the nutrition of the embryo in the lower vertebrates chiefly takes place. In mammals the umbilical ves- icle plays a comparatively subordinate part in the nourishment of the foetus, its function being generally superseded by the allantois. The last-named sac commences at a very early period as a diverticulum from the hinder end of the alimentary tract of the embryo. Its proximal portion afterwards becomes the urinary bladder, the contracted part between this and the cavity of the allantois proper constituting the urachus, which passes out of the body of the feetus at the umbilicus together with the vitelline duct. The mesoblastic tissue of the walls of the allantois soon becomes vascular ; its arteries are supplied with foetal blood by the two hypogastric branches of the iliacs, or main divisions of the abdominal aorta, and the blood is returned by venous trunks uniting to form the single umbilical vein which runs to the under surface of the liver, where, part of it joining the portal vein and part entering the vena cava directly, it is brought to the heart. These are 78 GENERAL ANATOMICAL CHARACTERS the vessels which, with their surrounding membranes, consti- tute the umbilical cord—the medium of communication between the foetus and the placenta, when that organ is fully de- veloped. The ege membranes of the Monotremes present many points of agreement with those of the ovum of the Marsupials,! and differ from those of the Placental types. Thus Monotremes and Marsu- pials agree in having a vitelline membrane, which appears between the young ovum and the follicular epithelium, persisting in the one case until the time of hatching, and in the other till a late uterine stage. There are also several other common features fully described in Mr. Caldwell’s memoir, but which cannot be detailed in this work. In the Marsupialia the observations made many years ago by Sir R. Owen upon the development of the Kangaroo have been confirmed by those of Dr. H. C. Chapman,? while Dr. Selenka,? and Professor H. F. Osborn have contributed important evidence as to the structure and relations of the foetal membranes of the Opossums and others. It thus appears that up to the period of the very premature birth of these animals the outer covering of the ovum, or false chorion, is free from persistent villi, and not adherent to the epithelium of the uterine walls; for, although fitting into the folds of the latter, it is perfectly and readily separable in its entire extent from them. The umbilical vesicle or yolk-sac is large, vascular, and adherent to a considerable portion of the false chorion or subzonal membrane, while the allantois is relatively small, and although the usual blood-vessels can be traced into it, it does not appear to contract any connection with the false chorion, and, there- fore, much less with the walls of the uterus, of such a nature as to constitute a placenta. In some forms, however, such as the Opossums, the umbilical vesicle or yolk-sac develops temporary villi, which unite with the subzonal membrane, or false chorion, to form a disc-like area closely attached to the cells covering the utricular glands of the uterine epithelium, and thus forming a so-called yolk-sac placenta. The function of this organ is considered to be the transmission of the secretions of the utricular glands to the embryo by means of the umbilical vesicle ; the function of the allantois being [either respiratory or the absorption of the fluid secreted in the uterine cavity by the utricular glands. While in the uterus the nourishment of the fcetus seems, there- fore, to be derived from the umbilical vesicle, as in reptiles and 1 See B. H. Caldwell—‘‘ The Embryology of Monotremata and Marsupialia,” Phil. Trans. for 1887, p. 463. 2 Proc. Acad. Nat. Sct. Philadelphia, 1881, p. 468. ° © Studien uéber Entwickelungeschichte der Thierc,” pt. 4, Wiesbaden, 1886. 4 Journal of Morphology, vol. i. p. 873 (1887). REPRODUCTIVE ORGANS 79 birds, rather than from the uterine walls by means of the allantoic vessels, as in the higher mammals. The latter vessels, in fact, play even a much less important part in the development of these animals, not only than in the placental mammals, but even than in the Sauropsida, for they can scarcely have the respiratory function assigned to them in that group: pulmonary respiration and the lacteal secretion of the mother very early superseding all other methods of providing the due supply both of oxygen and of food required for the development and growth of the young animal. In this sense the Marsupials may be looked upon as the most typically “‘mammalian” of the whole class. In no other group do the milk-secreting glands play such an important part in pro- viding for the continuity of the race. In the third primary division of the Mammalia, the so-called Placentalia, the umbilical vesicle generally does not quite unite with the chorion, and disappears as development proceeds, so that no trace of it can be seen in the membranes of an advanced embryo; but it may persist until the end of the intra-uterine life as a distinct sac in the umbilical cord, or lying between the allantois and amnion. The disappearance or persistence of the umbilical vesicle does not, according to our present knowledge, appear to be correlated with a higher or lower general grade of de- velopment, as might be presupposed. It is stated to have been found in Man even up to the end of intra-uterine life, and also in the Carnivora, while in the Ungulata and Cetacea it disappears at an earlier age. In many, if not all, of the Rodentia, Insectivora, and Chiroptera, it plays a more important part, becoming adherent to a considerable part of the inner surface of the chorion, to which it conveys blood-vessels, although villi do not appear to be developed from the surface of this part, as they are on the portion of the chorion supplied by the allantoic vessels. These orders thus present to a certain extent a transitional condition from the Mar- supials, although essentially different, in possessing the structures next to be described. The special characteristic of the whole of the placental mammals constituting the majority of the class, is that the allantois and its vessels become intimately blended with a smaller or greater part of the parietes of the ovum, forming a structure on the outer surface of which villi are developed, and which, penetrating into corresponding cavities of the “decidua,” or soft, vascular, hypertrophied. lining membrane of the uterus, constitutes the placenta. This organ may be regarded, as Sir William Turner says, both in its function and in the relative arrangement of its constituent textures, as a specially modified secreting gland, the ducts of which are represented by the extremities of the blood-vessels of the fetal system. The passage of material from the maternal to the foetal system of vessels is not 80 GENERAL ANATOMICAL CHARACTERS a simple percolation or diffusion through their walls, but is oc- casioned by the action of a layer of cells derived from the maternal or uterine structures, and interposed between the blood-vessels of the maternal part of the placenta and those of the villi covering the chorion, in which the embryonic vessels ramify. The numerous modifications in the details of the structure of this organ relate to augmenting the absorbing capacity of the vessels of the chorion, and are brought about either by increasing the com- plexity of the foetal villi and maternal crypts over a limited area, or by increasing the area of the part of the chorion covered by the placental villi, or by various combinations of the two methods. The first class of variations has given rise to a distinction into two principal kinds of placenta: (1) simple or non-deciduate, and (2) deciduate. In the former the fcetal villi are received into corre- sponding depressions of the maternal surface, from which at the period of parturition they are simply withdrawn. In the second, or more complex form, the relation is more intimate, a layer of greater or less thickness of the lining membrane of the uterus, called “decidua,” becoming so intimately blended with the chorion as to form part of the placenta proper, or that structure which is cast off as a solid body at parturition. In other words, in the one case the line of separation between the placenta and uterus at birth takes place at the junction of the foetal and maternal structures, in the other through the latter, so that a portion of them, often of con- siderable thickness, and containing highly organised structures, is cast off with the former. It was once thought that the distinction between these two forms of placentation is so important as to con- stitute a sufficiently valid basis for a primary division of the pla- cental mammals into two groups. It has, however, been shown that the distinction is one rather of degree than of kind, as inter- mediate conditions may exist, and it is probable that in different primary groups the simpler, non-deciduate form may have become developed independently into one or other of the more complex kinds. Apart from its intimate structure, the placenta may be met with of very varied general form. It may consist of villi scattered more or less regularly over the greater part of the surface of the chorion, the two extremities or poles being usually more or less bare. This form is called the “diffused placenta.” It is probably a primitive condition, from which most of the others are derived, although its existence must presuppose the absence of the umbilical vesicle as a constituent of the chorionic wall. It is found at present in the Manis among Edentates, the Cetacea, the Perissodactyle Ungulates, and the Camels, Pigs, and Chevrotains among the Artiodactyles. Such placente are always non-deciduate. Recent observations by Sir W. Turner on the placentation of the Dugong show that the REPRODUCTIVE ORGANS 81 Sirenia present the peculiarity of having a zonary placenta, which is either entirely or in great part non-deciduate, and is, therefore, transitional between the diffused and the true zonary type. In the true Ruminants or Pecora, among the Artiodactyle Ungulates, the villi are aggregated in masses called cotyledons, with bare spaces between. Such a placentation is called “ poly- cotyledonary.” In another modification the villi are collected in a more or less broad band encircling the chorion, leaving a very large portion of the two poles bare, constituting the “ zonary placenta,” characteristic of the Carnivora, and also occurring in the Elephant, Hyrax, and Orycteropus. The fact of the form of the placenta of these three last-named animals agreeing together, and with that of the Carnivora, does not, however, necessitate the ascription of zoological affinities, as the same ultimate form may have been attained by different processes of development. In another form one pole only of the chorion is non-vascular, the placenta assuming a dome or bell shape, as in the Lemurs and the Sloths. The transition from this, by the gradual restriction of the vascular area, is easy to the oval or discoidal form of placenta of the Anteaters, Armadillos, and higher Primates. The discoidal placenta of the Rodents, Insectivores, and Chiroptera, though show- ing so much superficial resemblance to that of the last-named order as to have led to the inclusion of all these forms in one primary group, is now known to be developed in another manner, not by the concentration of villi from a diffused to a limited area, but by retaining the area to which it was originally restricted in con- sequence of the large surface of the chorion occupied, as before mentioned, by the umbilical vesicle. To compensate for the small- ness of area, the complex or deciduate structure has been developed. Among some Rodents there is evidence to show that the discoidal placenta has been derived from a zonary one, of which distinct vestiges have been detected in the Mouse. We may conclude that, although the characters and arrangement of the fetal structures may not have that extreme importance which has been attributed to them by some zoologists, they will form, especially when more completely understood, valuable aids in the study of the natural affinities and evolution of the Mammalia.? 1 For a full exposition of the present state of knowledge on this subject, see the various memoirs of Sir William Turner, also F. M. Balfour’s Treatise on Comparative Embryology, vol. ii. (1881), and J. A. Ryder in American Naturalist, vol. xxi. p. 780 (1887). CHAPTER III ORIGIN AND CLASSIFICATION OF THE MAMMALIA Origin.—Although, as stated in the first chapter, the mammalian class, as at present known either by existing or extinct forms, is completely isolated from all other groups of the animal kingdom, yet it is impossible to refrain from speculating as to its origin and nearest affinities. In arranging the classes of vertebrates in a linear series it is customary to place them in the following order—Pisces, Amphibia, Reptilia, Aves, Mammalia,—an order which probably indicates the relative degree of elevation to which the mos highly developed members of each class has attained. Such an arrangement appears to express the true relationship of the first four classes to one another, but it is quite clear that the Mammalia have no sort of affinity with the Aves. Writing in 1879, Professor Huxley + came to the conclusion that, in looking among vertebrates for the progenitors of the Mammalia, we must pass over all known forms of birds and reptiles, and go straight down to the Amphibia. In addition to the characters derived from the conformation of the pelvis upon which the argument was primarily based, the following reasons were given for this conclusion: “The Amphibia are the only air-breathing Vertebrata which, like mammals, have a dicon- dylian skull. It is only in them that the articular element of the mandibular arch remains cartilaginous, while the quadrate ossifica- tion is small, and the squamosal extends down over it to the osseous elements of the mandible, thus affording an easy transition to the mammalian condition of those parts. The pectoral arch [girdle] of the Monotremes is as much amphibian as it is sauropsidian; the carpus and the tarsus of all Sauropsida, except the Chelonia, are modified away from the Urodele type, while those of the mammal are directly reducible to it. Finally, the fact that in all'Sauropsida it is a right aortic arch which is the main conduit of arterial blood leaving the heart, while in mammals it is a left aortic arch which 1 Proceedings of the Royal Soctety of London, vol. xxviii. p. 395 (1879). ORIGIN . 83 performs this office, is a great stumbling-block in the way of the derivation of the Mammalia from any of the Sauropsida. But, if we suppose the earliest forms of both the Mammalia and the Saur- opsida to have had a common Amphibian origin, there is no difficulty in the supposition that, from the first, it was a left aortic arch in the one series, and the corresponding right aortic arch in the other, which became the predominant feeder of the arterial system.” Subsequently Professor E. D. Cope! in a suggestive paper called attention to the remarkable resemblances to the Monotremes pre- sented by the skeleton of that group of early secondary reptiles which he then designated the Theromorpha, but which may be included in the Anomodontia of Sir R. Owen, and came to the conclusion that in that group we have the true ancestors of the Mammalia. This conclusion was, however, disputed by Dr. Baur,? who considered that the Anomodontia were too specialised to have been the actual progenitors of the Mammalia, and that they should rather be regarded as a divergent branch of the stem which had given origin to the Mammalia. Since that date observations made on the structure of the South African Anomodonts have shown such an intimate connection between that group and the Labyrinthodont Amphibians, that there can be no hesitation in regarding the one as the direct descendant of the other; and we may probably regard the Mammalia as having originated from the same ancestral stock at the time the Amphibian type was passing into the Reptilian. From this point of view, some of the mammalian features found in the more specialised Anomodonts may probably be regarded as having been acquired during a parallel line of development. Both the Anomodontia and the Mammalia differ from the Amphibians in the loss of. the splint-like parasphenoid which underlies the basisphenoid axis of the skull, and by the ossification of that axis; but while the former have become monocondylic by the participation of the basioccipital in the support of the cranium, the latter retain the Amphibian dicondylic plan. The skull of the Anomodonts presents mammalian resemblances not found in any other Reptiles, this being especially noticeable in the region of the squamosal; and it is only in this group and mammals that the temporal or zygomatic arch is a squamoso-maxillary one (see p. 37). The resemblance between the pectoral and pelvic girdles of the Anomodonts and those of the Monotreme Mammals is noticed under the head of the latter, where reference is also made to the similarity in the structure of the humerus in the two groups. 1 “The Relations between the Theromorphous Reptiles and the Monotreme Mammalia,” Proceedings of the American Association for the Advancement of Science, vol. xxxiii. p. 471 (1885). 2 “On the Phylogenetic Arrangement of the Sauropsida,” Journal of Morphology, vol. i. pp. 98-104 (1887). 84 ORIGIN AND CLASSIFICATION The pes of the Amphibia and Anomodontia agree in having a distinct intermedium, tibiale, fibulare, and centrale, whereas in other Reptiles these bones are not generally distinct ; in Mammals the intermedium, fibulare, and centrale are distinct, and according to Cope’s interpretation there may be a distinct tibiale. Classification.—In the present condition of the world, mammals have become so broken up into distinct groups by the extinction of intermediate forms, that a systematic classification is perfectly practicable. Most of the associations of species, which we call “orders,” and even the “suborders” and “families,” are natural groups. In isolating, defining, and naming them, we are really dealing with facts of nature of a totally different order from the artificial and fanciful divisions formed in the infancy of zoological science. When, however, we pass to the extinct world, all is changed. In many cases the boundaries of our groups become enlarged until they touch those of others. New forms are discovered which cannot be placed within any of the existing divisions. As the horizon of our vision is thus expanded, the principles upon which a scheme of classification is constructed must be altogether changed. Our present divisions and terminology are no longer sufficient for the purpose; and some other method will have to be invented to show the complex relationships existing between different animal forms when viewed as a whole. The present time, pre-eminently distinguished by the rapidly changing and advancing knowledge of extinct forms, is scarcely one in which this can be done with any satisfactory result; so that all attempts to form a classification embracing even the already known extinct species must be only of a provisional and temporary nature. In systematic descriptions in books, in lists, and catalogues, and in arranging collections, the objects dealt with must be placed in a single linear series. But by no means whatever can such a series be made to coincide with natural affinities. The artificial character of such an arrangement, the constant violation of all true relation- ships, are the more painfully evident the greater the knowledge of the real structure and affinities. But the necessity is obvious; and all that can be done is to make such an arrangement as little as possible discordant with facts. The following table contains a list of the orders, suborders, and families of existing mammals as recognised by the authors, and placed in the order in which they will be treated of in this work. The more important of the groups containing only extinct forms are added in a different type, being interpolated, as near as may be, among those that appear to be their existing relatives. A few explanatory remarks upon the mutual relations of some of the principal groups mentioned in the table may be useful here, CLASSIFICATION 85 but the subject will be more fully developed in treating separately of each division. One of the most certain and fundamental points in the classifica- tion of the Mammalia is, that all the animals now composing the class can be grouped primarily into three natural divisions, which, presenting very marked differential characters, and having no exist- ing, or yet certainly demonstrated extinct, intermediate, or trans- itional forms, may be considered as subclasses of equal value, tax- onomically speaking, though very different in the numbers and importance of the animals at present composing them. These three groups are often called by the names originally proposed for them by Blainville—(1) Ornithodelphia, (2) Didelphia, (3) Monodelphia— the first being equivalent to the order Monotremata, the second to the Marsupialia, and the third including all the remaining members of the class. Although actual paleontological proof is wanting, there is much reason to believe that each of these, as now existing, are survivors of distinct branches to which the earliest forms of mammals have successively given rise, and for which hypothetical branches Professor Huxley has proposed the names of Prototheria, Metatheria, and Eutheria, names which, being far less open to objection than those of Blainville, are here used as equiva- lents of the latter. The only known existing PRoTOTHERIA, although agreeing in many important characters, evidently represent two very divergent stocks, perhaps as far removed as are the members of some of the accepted orders of the Kutheria. It would, however, be merely encumbering zoological science with new names to give them any other than the ordinarily known family designations of Ornitho- rhynchide and Echidnide. Similarly with regard to the METaTHERIA, although the great diversity in external form, in anatomical characters, and in mode of life of the various animals of this section might lead to their division into groups equivalent to the orders of the Eutheria, we do not think it advisable to depart from the usual custom of treating them all as forming one order, called Marsupialia, the limits of which are equivalent to those of the subclass. The characters of the six families which compose the group are extremely well marked and easily defined ; and since they form a regular gradation between two extreme types, they can be satisfactorily arranged in a serial order. A marked distinction in the dentition enables us to divide them into primary groups or suborders. The remaining mammals are included in the EUTHERIA, PLACEN- TALIA, or MoNoDELPHIA. Their affinities with one another are so complex that it is impossible to arrange them serially with any regard to natural affinities. Indeed each order is now so isolated that it is almost impossible to say what its affinities are; and none 86 ORIGIN AND CLASSIFICATION of the hitherto proposed associations of the orders into larger groups stand the test of critical investigation. All serial arrangements of the orders are therefore perfectly arbitrary ; and although it would be of very great convenience for reference in books and museums if some general sequence, such as that here proposed, were generally adopted, such a result can scarcely be expected, since equally good reasons might be given for almost any other combination of the various elements of which the series is composed. In fact, we have already seen reason to depart in some respects from that used in the “ Encyclopedia.” The Edentata, Sirenia, and Cetacea stand apart from all the rest in the fact that their dentition does not conform to the general heterodont, diphyodont type to which that of all other Eutheria can be reduced, and which is such a close bond of union between them. In all three orders, however, some indications may be traced of relationship, however distant, with the general type. With regard to the Edentata, reasons will be given for believing that both the Sloths and Anteaters are nearly related, and that the Armadillos, though much modified, belong to the same stock, but that the Pangolins and the Aard-varks represent very isolated forms. There is no difficulty about the limits of the order Sirenia, com- prising aquatic, vegetable-eating animals, with complete absence of hind limbs, and low cerebral organisation, represented in our present state of knowledge only by two existing genera, Halicore and Mana- tus, and a few extinct forms, which, though approaching a more generalised mammalian type, show no special characters allying them to any of the other orders. The few facts as yet collected relating to the former history of the Sirenia leave us as much in the dark as to the origin and affinities of this peculiar group of animals as we were when we only knew the living members. They lend no countenance to their association with the Cetacea - and, on the other hand, their supposed affinity with the Ungulata receives no very material support from them. Another equally well-marked and equally isolated, though far more numerously represented and diversified order, is that of the Cetacea, placed simply for convenience next to the Sirenia; with which, except in their fish-like adaptation to aquatic life, they have little in common. ‘The old association of these orders in one group can only be maintained either in ignorance of their structure or in an avowedly artificial system. Among the existing members of the order, there are two very distinct types, the toothed Whales or Odontoceti, and the Baleen Whales or Mystacoceti, which present as many marked distinguishing structural characters as are found between many other divisions of the Mammalia usually reckoned as orders. Since the extinct Zeuglodonts, so far as their characters CLASSIFICATION 87 are known, do not fall into either of these groups, but are in some respects annectant forms, we have placed them provisionally, at least, in a third group by themselves, named Archeoceti. There is nothing known at present to connect the Cetacea with any other order of Mammals; but it is quite as likely that they are offsets of a primitive Ungulate as of a Carnivorous type, or perhaps of a still more generalised mammalian stock. The remaining Eutherian mammals are clearly united by the characters of their teeth, being all heterodont and diphyodont, with their dental system reducible to a common formula. Although older views of, the relationship of Ungulate mammals expressed by the terms Pachydermata, Runvinantia, and so forth, still linger in some corners of zoological literature, no single point in zoological classification can be considered so firmly established as the distinction between the Perissodactyle and Artiodactyle Ungulates ; both being in the existing fauna of the world perfectly natural and distinctly circumscribed groups. The breaking-up of the latter into four equivalent sections, the Pecora, Tylopoda, Tragulina, and Suina, is equally in accordance with all known facts. Less certain, however, is the association of the Proboscidea and the Hyracoidea with the true Ungulates. By many zoologists they are each, although containing so very few existing species, made into distinct orders; and much is to be said in favour of this view. The discovery, however, of a vast number of extinct species of Ungu- lates which cannot be brought under the definition of either Perisso- dactyla or Artiodactyla, and yet are evidently allied to both, and to a certain extent bridge over the interval between them and the isolated groups just mentioned, make it necessary either to intro- duce a number of new and ill-defined ordinal divisions, or so to widen the scope of the original order as to embrace them all, considering the Elephants and the Hyraces as representing sub- orders equivalent to the great Perissodactyle and Artiodactyle groups. It is the latter alternative that we have adopted. The Rodentia, although generally presenting a low grade of development, are a very specialised and distinct group. The position here assigned to them would accord with apparent relation- ships with the Ungulates, through the Elephant on the one hand and the extinct Typotherium on the other. In the present state of the fauna of the earth, the Carnivora form a very distinct order, though naturally subdivided into two groups, the members of the one being more typical, while those of the other (the Pinnipedia) are aberrant, having the whole of their organisation specially modified for living habitually in the water. The Insectivora comprise various lowly organised and generalised forms, exhibiting considerable divergence of character, and ap- parently connected through transitional extinct species with the 88 ORIGIN AND CLASSIFICATION Carnivora. As no other order can claim the family Galeopithecide, it is placed here, but rather for convenience than for any other consideration, since it has but little if any relationship with any of the other members. Its isolated position is indicated by assigning it a distinct subordinal rank. The Chiroptera have always been placed near the Insectivora ; but they are really a highly specialised group, as much isolated from all other mammals by the modification of their anterior limbs in adaptation to aerial locomotion, as the Cetacea and the Sirenia, by the absence of hind limbs, are specially adapted for an aquatic life. Lastly, the Primates, which in any natural system must be placed at the head of the series, are divisible into two very distinct groups—one containing the various forms of Lemurs (Lemuroidea), and the other the Monkeys and Man (Anthropoidea). Whether the Lemuroidea should form part of the Primates (according to the traditional view), or a distinct order altogether removed from it, is as yet an undetermined question, for both sides of which there is much to be said. There can, however, be no doubt that the Anthropoidea form a perfectly natural group, presenting a series of tolerably regular gradations from the Marmosets (Hapale) to Man. Certain breaks in the series, however, enable us to divide it into five distinct families:—Hapalide or Marmosets ; Cebide or American Monkeys, with three premolar teeth on each side of each jaw; Cercopithecide, containing the majority of Old-world Monkeys ; Simiide, consisting of the genera Hylobates, Simia, Gorilla, and Anthropopithecus, the true Man-like Apes; and, lastly, Hominude, containing the genus Homo alone. Subclass I. PRoTOTHERIA. Order i. MonoTREMATA—Monotremes. Fam. 1. Ornithorhynchide—Duck-bill. 2, Echidnide—Spiny Anteater. Group. MULTITUBERCULATA.! Fam. 1. Plagiaulacide—Plagiaulax. 2. Polymastodontidee—Polymastodon. 3. Tritylodontide—tTritylodon. Subclass IT. METATHERIA. Order ii. Marsuprati14—Marsupials. Suborder 1. Potyproropont1a—Polyprotodonts. 1 The names of the groups containing only extinct forms are printed in heavier type than those which contain species still existing. CLASSIFICATION 89 Fam. 1. Dromatheriide—Dromatherium. 2. Amphitheriide—Amphitherium, etc. 3. Spalacotheriidz—Spalacotherium. 4. Tritylodontide—Tritylodon. 5. Didelphyide—Opossums. 6. Dasyuride—Thylacine and Dasyures. 7. Peramelide—Bandicoots. Suborder 2. DrpRoropontra—Diprotodonts. Fam. 8. Phascolomyide—W ombats. 9. Phalangeride—Phalangers. 10. Diprotodontidze—Diprotodon. 11. Nototheriidee—Notothere. 12. Macropodide—Kangaroos. Subclass IIT. Euruerta. Order iii. EDENTATA—Edentates. Fam, 1. Bradypodide—Sloths. 2. Megatheriidee—Ground Sloths. . Myrmecophagide—Anteaters. . Dasypodide—Armadillos. . Glyptodontide—Glyptodonts. . Manide—Pangolins. . Orycteropodide—Aard-varks. TO op Order iv. SrRENIA—Sirenians. Fam. 1. Manatide—Manatees. 2. Rhytinide—Rhytina. 3. Halicoride—Dugongs. 4. Halitheriide—Halithere. Order v. CreTacEA—Cetaceans. Suborder 1. Mystacoceti—Baleen Whales. Fam. 1. Balenide—Greenland Whale, etc. Suborder 2. ARCHAOCETI. Fam. 2. Zeuglodontide—Zeuglodonts. Suborder 3. Opontoceti—Toothed Whales. Fam. 3. Physeteride—Sperm Whale. 4. Platanistide—Freshwater Dolphins. 5. Delphinide—Dolphins, Porpoises, ete. Order vi. UNGuLATA—Hoofed Mammals. Suborder 1. ArtiopactyLa—Artiodactyles. Section A. Suina—Pig-like Artiodactyles. Fam. 1. Hippopotamide—Hippopotamus. 2. Suwida—Pigs and Peccaries. go ORIGIN AND CLASSIFICATION . Cheropotamide—Cheeropotamus. . Anthracotheriide—Anthracothere. . Merycopotamide—Merycopotamus. types. eae eee ea, OCIA AB w Annectant Section B. 9. Section C. 10. 11. Section D. 12. 13. 14. 15. Suborder 2. Fam. 16. 17. 18. 19. 20. 21. 22. 23. 24, Suborder 3. Fam. 25. 26. Suborder 4. Fam. 27. 28. 29. Suborder 5. Fam. 30. Suborder 6. Fam. 31. : 32. 33. Suborder 7. Fam. 34. 35. . Cotylopidze—Oreodonts. . Anoplotheriidz—Anoplothere. . Dichodontidee—Dichodon. TRAGULINA—Chevrotains. Tragulide—Chevrotains. TyLopopa—Camels. Camelide—Camels and Llamas. Poebrotheriidee—Poébrotherium. Precora—True Ruminants. Cervide—Deer. Giraffide—Giraffe, Antilocapride—Prong-buck. Bovide—Sheep, Cattle, etc. PERISSODACTYLA—Perissodactyles. Tapiride—Tapirs. Lophiodontide—Lophiodonts. Paleotheriidee—Palzotheres. Lquide—Horses. Rhinocerotide—Rhinoceroses. Lambdotheriide— Palzosyops. Chalicotheriidae—Chalicothere. Titanotheriidee—Titanothere. Macraucheniide— Macrauchenia. TOXODONTIA—Toxodonts. Toxodontide—Toxodon. Typotheriide—Typothere. CONDYLARTHRA. Periptychide—Periptychus. Phenacodontidz—Phenacodus. Meniscotheriidee—Meniscothere. Hyracoipra—Hyraces. Hyracide—Hyrax. AMBLYPODA. Pantolambdidea—Pantolambda, Coryphodontidz—Coryphodon. UVintatheriidze—Uintathere. Prosposcip—Ea—Proboscideans, Dinotheriide— Dinothere. Elephantide—Elephants. CLASSIFICATION gI Group. TILLODONTIA—Tillodonts, Fam. Anchippodontidze—Anchippodus. Calamodontidz—Calamodon. Order vii. RopENTIA—Rodents. Suborder 1. SIMPLICIDENTATA. Fam. 1. Anomaluride—Anomalurus. 2. Sciuride—Squirrels and Marmots. 3. Haplodontide—Haplodon. 4, Ischyromyide—Ischyromys. 5. Castortdee—Beavers. 6. Myoxide—Dormice. 7. Lophiomyide—Lophiomys. 8. Muride—Rats, Mice, and Voles. 9. Spalactde—Mole-rats. 10. Geomyide—Pouched Rats. 11. Dipodide—Jerboas. 12. Theridomyide—Theridomys. 13. Octodontide—Spiny Mice. 14. Castoroididze—Castoroides, 15. Hystricide—Porcupines. 16. Chinchillide—Chinchillas. 17. Dinomyide—Dinomys. 18. Cavitde—Cavies. 19. Dasyproctide—Agouties. Suborder 2. DuPLICIDENTATA. Fam. 20. Lagomyide—Picas. 21. Leporide—Hares and Rabbits. Order viii. CARNIVORA—Carnivores. Suborder 1. Carnivora VERA—Fissipedes. Fam. 1. Felide-—Cats. . Hyenide—Hyeenas. . Proteleide—Earth-wolf. . Viverride—Civets and Ichneumons. . Canide—Wolves and Foxes. . Urside—Bears. . Mustelide—Weasels and Otters. . Procyonide—Racoons and Cat-bear. Suborder 2. Prnnipep1a—Pinnipedes. Fam. 9. Otartide—Eared Seals. 10. Trichechide—Walrus. 11. Phocide—Seals. Suborder 3. CREODONTA—Creodonts. Fam. 12. Hyenodontide—Hyznodon. 13. Proviverride—Proviverra. 14, Arctocyonidze— Arctocyon. 15. Mesonychidze—Mesonyx. ODIDoapP wD 92 ORIGIN AND CLASSIFICATION Order ix. INsEcTIVoRA—Insectivores. Suborder 1. Fam. 1. Corn ow w po Suborder 2. Fam. 10. INSECTIVORA VERA. Tupaiide—tTupaias. . Macroscelidide—Elephant-Shrews. . Hrinaceide—Hedgehogs. . Soricide—Shrews. . Talpide—Moles. . Potamogalide—Potamogale. . Solenodontide—Solenodon. . Centetide—Centetes. . Chrysochloridee—Golden Moles. DERMOPTERA. Galeopithecide—Galeopithecus. Order x. CHIROPTERA—Bats. Suborder 1. Fam. 1. Suborder 2. Fam. 2. 2. 4 5. 6 Mrcacurroprera—Frugivorous Bats. Pteropodide—Flying Foxes. MicrocHiRoprERA—Insectivorous Bats. Vespertiliontde—Common Bats. Nycteride—Nycteris. . Rhinolophide—Leaf-nosed Bats. Emballonuride—Emballonura. . Phyllostomatide—Vampyres. Order xi. PRIMATES. Suborder 1. Fam. 1. 2, Dy 4, Suborder 2. Fam. 5. oO aorta Lemuromipgea—Lemuroids. Hyopsodontidea—Hyopsodus. Chiromyide—Aye-Aye. Tarstide—Tarsier. Lemuride—Lemurs. ANTHROPOIDEA—Anthropoids. Hapalide—Marmosets. . Cebide—American Monkeys. . Cercopithecide—Old World Monkeys. . Simide—Gibbons and Man-like Apes. . Hominide,—Man. The distinctive character of these subclasses and orders, with an account of their subdivisions and the principal forms contained in each, will be given in subsequent chapters. CHAPTER IV GEOGRAPHICAL AND GEOLOGICAL DISTRIBUTION I, GEOGRAPHICAL DISTRIBUTION.! In considering the present distribution of mammals over the globe, we may, in the first place, direct our attention to terrestrial or land types, reserving the consideration of aerial types, like the Bats, and aquatic forms, as exemplified by the Cetaceans, Sirenians, and Seals, to separate sections. Among terrestrial forms each species has a certain definite area of distribution in space, which may be of very wide extent, or may be confined to a restricted region. This distributional area is, however, always connected, or continuous; that is to say, that although we may have a single species inhabiting two continents, like the Lion in Asia and Africa, or dwelling both on a continent and adjacent continental islands, like the Javan Rhinoceros of India, Java, and Borneo, yet we shall always find that such areas, if not still connected, show evident signs of having been so connected in comparatively late geological epochs; and we never find instances of the same species inhabiting totally disconnected areas, such as India and South America. As examples of mammals with a wide distribution we may mention the Lion and the Leopard, which are now found throughout Africa, and also occur in India, as well as in the intervening areas of Arabia and Persia. In the case of the former species, paleontology further teaches us that its distribution in the last geological epoch was even more extensive, since we have good evidence to show that it formerly ranged over the greater part of Europe, including the British Isles. The Jackal affords another well-known instance of a species common 1 On this subject see A. Murray, Geographical Distribution of Mammals, 1866 ; and especially A. R. Wallace, The Geographical Distribution of Animals, 2 vols., 1876, and Island Life, 1881; also A. Heilprin, The Geographical and Geological Distribution of Animals, 1887. 94 GEOGRAPHICAL DISTRIBUTION to India and Africa. The American Puma, again, may be cited as an example of a mammal having a very wide range in latitude, since it is found from Patagonia in the south to Canada in the north. As instances of wide range in the opposite direction we have only to mention the Reindeer and the Elk or Moose, found in the northern regions of both the Old and New Worlds, which are only separated from one another by the narrow channel of Behring Strait. Of mammals with extremely restricted distributional areas, we may mention many of the Insectivora, such as the Desman of the Pyrenees, and some of the Madagascar types of this order, the Lemurs from the same island, some of the species of Marmots, the remarkable bear-like Aluropus of Eastern Tibet, one species of Zebra, and other Ungulates from Africa. The distribution of a genus (except of course when the genus is represented only by a single form) is very generally more exten- sive than that of a species; and this may be markedly the case when there are only some two or three species in a genus. In genera, moreover, we meet with what is known as discontinuous distribution, that is, where the distributional area of one or more’ species is totally separated from that of others. The best instance of this occurs in the case of the Tapirs, where we find one species inhabiting the Malayan Peninsula, and no others anywhere in the world, with the exception of South America. The explanation of such an apparently anomalous feature in distribution is to be found in the past history of the globe, which shows us that Tapirs once existed in China, Europe, and North America, and, therefore, indicates that the existing isolated species are the sole survivors of a group once spread over a large portion of the earth’s surface. In regard to generic distribution it must, however, be mentioned that this depends to a great extent on the limits which we are disposed to assign to genera themselves. As the distributional area of a genus generally exceeds that of a species, so that of a family, or group of genera, is larger than that of a single genus; and similarly the distribution of an order, or assemblage of families, usually occupies a larger area than that of a single family. Thus, for instance, the genus Thylacinus, re- presented only by the so-called Tasmanian Wolf or Thylacine, is now entirely restricted to Tasmania ; but the family Dasyuride, to which that genus belongs, ranges all over Australia, while the order Marsupialia, which includes the Dasyuride, is found both in Aus- tralia and America, and in past epochs was probably spread over the entire globe. ; A remarkable feature in connection with the distribution of the terrestrial Mammalia is the circumstance that, with the exception of certain species introduced by human agency, and small forms which TERRESTRIAL DISTRIBUTION 95 can easily have been transported on floating timber or other similar means, they are totally absent from what are known as oceanic islands—that is islands arising from great depths in the ocean, mainly composed of coral or volcanic rocks, and showing no signs of having ever been connected with the existing continents, or the larger and so-called continental islands. The obvious explanation of this feature is, that from their total isolation these islands have never been able to receive a mammalian fauna from the great continental areas on which mammalian life was probably first developed. As an intermediate step between these islands which are practically void of mammalian life and the continents which teem with such a variety of forms, are certain larger islands and portions of continents containing a mammalian fauna more or less markedly distinct from that of the whole of the other regions of the globe. The best instance of this is Australia, which, with the exception of one dog—the Dingo—and certain Muride and Bats, has no mammals except Monotremes and Marsupials. The latter are, moreover, per- fectly distinct from those of America, which, if we exclude the islands in the neighbourhood of Australia, is the only other region which now possesses any Marsupials at all. Here also we have a ready and full explanation which accords with all the facts; since it is evident that Australia has been isolated from the Asiatic continent from some very remote geological epoch, at which period it is probable that Monotremes and Marsupials were the dominant if not the sole representatives of the Mammalia then existing. Consequently Australia has never been able to receive an influx of the Eutherian orders, which have probably swept away all the Marsupials except the small American Opossums from the rest of the globe. Again, the large island of Madagascar, which has a fauna of an African type, but still very markedly different from that of the mainland, may be considered to have been connected with the latter at a time when the Eutheria had become the dominant forms, but has been separated for a sufficiently long period to have enabled a large number of its species and genera to have become distinct from those of the adjacent continent. Similarly, there is evidence to show that South America was probably cut off for a considerable period _from the northern half of the American continent, in consequence of which its lowly organised fauna of Edentates were enabled to attain such a remarkable development in the later geological periods. In contrast to the mammalian fauna of islands of the preceding type is, or rather was, that of the British Islands, which in the early historic and prehistoric periods was identical with that of the Continent. This leads to the inference that at a comparatively late epoch there was a direct land communication between Britain 96 GEOGRAPHICAL DISTRIBUTION and the Continent, which is shown by geological evidence to have actually been the case. The above instances are sufficient to show what an important influence the date of separation of islands from the adjacent continents has had upon their existing mammalian fauna, and how largely the present distribution of mammalian life is bound up with the past history of our globe. We must, however, not omit to mention another very important agency of past times which has likewise had great influence on the present distribution of the various faunas of the northern hemisphere. This is the so-called glacial epoch, which took place immediately before the establish- ment of the present condition of things, and appears to have been the cause of the extinction of many of the larger mammalian types which formerly inhabited Europe, and whose retreat to the warmer regions of the south was apparently cut off by the Mediterranean. Zoological Regions.—Zoologists are now generally agreed in dividing the land surfaces of the globe into a number of zoological regions or provinces, characterised by a more or less distinctly marked general Jacies of their fauna asa whole. Some of these regions are much more distinctly defined than the others; and in the majority of cases there is a kind of neutral ground or No-man’s-land at the junction between any two of these regions. It must also be remembered that in the Old World proper as we go back in time we find a gradual assimilation in the mammalian faunas of the different regions, indicating that originally there was one large fauna of a generally similar type occupying the greater portion of this area. Thus we find that Hippopotami, Giraffes, Kudus, Elands, and other types of Antelopes now restricted to Africa, formerly extended to Europe and India, while there is also evidence to show that the group of large anthropoid Apes, now found only in Africa and the Bornean region, were likewise spread over a large part of the south-western half of the Old World. Moreover, while at the present day there is a marked connection between the mammals of the northern regions of both the Old and New Worlds, in the Tertiary period it appears that the fauna of the whole of North America was much more nearly allied to that of the central regions of the Old World than is now the case. Thus in the Tertiary rocks of America we meet with remains of what we are accustomed to regard as such essentially Old World genera as Horses and Rhinoceroses. On the other hand there are no traces in America of the existence at any period of Apes, Giraffes, Hippopotami, or Hyznas, while that continent has yielded evidence of groups of Ungulates totally unrepresented in the eastern hemisphere. The chief zoological regions of the globe, proposed by Mr. Sclater in 1857, and now recognised by the majority of authorities, are six in number, and are named as follows. Firstly, the Palearctic ZOOLOGICAL REGIONS—PALAZARCTIC 97 region, embracing the whole of Europe, Persia, Northern Arabia, and all of Asia northward of the line of the Himalaya proper, Japan, that part of Africa lying northward of the Sahara Desert, and the oceanic islands of the North Atlantic. Secondly, the Ethiopian region, which comprises all Africa lying to the south of the Sahara, the southern part of Arabia, Madagascar, and the Mascarene Islands. Thirdly, the Oriental or Indian region, which is taken to include India south of the Himalaya, and to the north-west as far as Beluchistan, the Malay peninsula, southern China, Sumatra, Java, Borneo, and the Philippines. Fourthly, the Australasian region, which -is usually defined as being bounded to the north-west by the deep sea channel lying between Borneo and Celebes known as Wallace’s line, and is taken to include Celebes, Lumbok, New Guinea, Australia, Tasmania, New Zealand, and the host of oceanic islands in the South Pacific. Several writers, how- ever, prefer to regard Celebes and some of the adjacent islands as representing a transitional Austro-Malayan region. Fifthly, the Nearctic region, comprising Greenland and North America as far south as the north of Mexico. And, sixthly, the Neotropical region, which embraces the remaining portion of the American continent and the West Indies. Various minor modifications of this scheme have been proposed. Thus some writers are disposed to raise India to the rank of a distinct primary region, while others propose the same for New Zealand. The Palearctic and Nearctic regions have a large number of common types, more especially among the mammals, and Dr. A. Heilprin! has expressed his opinion that they should be regarded as a single primary region under the name of the Holarctic. The same writer would also separate the South Pacific Islands as con- stituting a Polynesian region. Minor divisions or sub-regions have also been marked out, but it will be unnecessary to indicate their limits in the present work. We may, however, mention the Mediterranean sub-region of the Palearctic, which includes the peninsular portion of southern Europe, North Africa, Asia Minor, Persia, Afghanistan, Beluchistan, and Northern Arabia, as a good instance of the transition from one region to another, since its fauna has a mingling of Palearctic, Ethiopian, and Oriental types, the former being, however, the predominant ones. Of the chief mammalian types characteristic of these various regions only a ‘brief sketch can be given in this work. Palearctic Region.—The Palearctic region is of enormous extent, and includes countries varying greatly in their flora, climate, and elevation. Thus it embraces the Arctic plains of Siberia, the warm regions of Italy, Southern France, and Northern Africa, the forest- 1 Distribution of Animals, 7 98 GEOGRAPHICAL DISTRIBUTION clad slopes of the outer Himalaya, and the lofty arid plains of Turk- estan and Tibet, scorched by a burning sun in summer and chilled by a still more terrible cold in winter. Its extreme limits in the west are marked by the Canaries and Azores, and in the east by distant Japan ; and yet throughout this vast expanse we find a great uni- formity of life, as exemplified by the large number of British genera which occur also in Japan. The mammals which are on the whole the most characteristic of this region are the Sheep and Goats, forming a section of the great family of Bovide, nearly all the species of which are Palearctic, although we meet with one Goat (Capra) in the Nilgherries of Southern India, and a Sheep (Ovis) in the Nearctic region. The Musk Ox (Ovibos) is characteristic of the Palearctic and Nearctic regions. At least one species of Camel is characteristic of this region, and it is not improbable that the second may also have originated in it. There are a few characteristic types of Antelopes, such as the Alpine Chamois (fupicapra), the Saiga of Tartary, and the Chiru (Pantholops) of Tibet, each of which is represented by only a single species; and we miss the host of Antelopes so characteristic of the Ethiopian region. Deer (Cervus) are abundant, although by no means confined to this region; and the Musk Deer (JJoschus), the sole representative of the subfamily Moschine, is exclusively Palearctic. Monkeys, as a rule, are absent, although we meet with one species of Macacus in Northern Africa and at Gibraltar, and some other types on the southern border of Tibet. The Moles (Zalpa) are mainly Palearctic, although one species enters Northern India, while the Desmans (Myogale) of the Pyrenees and Southern Russia are unknown beyond the limits of this region. The Water-shrew (Nectogale) is likewise a peculiar eastern Palearctic type. Among the Rodents, the Picas or Tailless Hares (Lagomys) and the Dormice (J/yorus) are essentially Palearctic forms, only one species of each being found beyond the limits of the region, and the one extra-Palzarctic species of Lagomys occurring in the cognate Nearctic region. The Mice and Rats are represented by the typical genus J/us and other types, and Hares (Lepus) and one species of Squirrel (Scivrus) are common. The Carnivora include two species of Bears (Ursus), Wolves and Foxes (Canis), a Lynx and a few species of Cats (Felis), as well as numerous weasels (Mustela), and some other types. Ethiopian Region—The Ethiopian region is of great interest to the student of mammals, since it is inhabited by a number of forms remarkable for their large size. A considerable portion of the area consists of desert, especially in the north; but there is also a wide extent of grassy plains (veltd), as well as vast tracts of equatorial forests of great density. Perhaps the most striking feature in the Ethiopian fauna is the number of Ungulates, both of the Artio- dactyle and Perissodactyle sections. In the former section we have ETHIOPIAN REGION 99 the Giraffes (Giraffa) represented by one species, which is the type of a family, and is unknown elsewhere. Equally characteristic are the Hippopotami, which likewise form the type of a family, while the Pigs are represented by the Wart-hogs (Phacocherus) and the River-hogs, forming an aberrant group of the genus Sus. The Oxen (Bos) are represented by Buffaloes, but there are no species of true Oxen or Bison. The Antelopes attain an extraordinary develop- ment, the number of species being estimated at from eighty to ninety, which are referred to a large number of genera, although several of these are more or less ill defined. Most of these genera are peculiar to this region, but the Gazelles (Gazella) are also found in the desert regions of other parts of the Old World, and Orya ranges into Arabia and Persia. In contrast to this abundance of Antelopes is the total absence of the Deer family, or Cervidw, which are so characteristic of the Palearctic and Oriental regions. The Chevrotains or Tragulide are, however, represented by Dorcatherium.1 In the Perissodactyle section we may notice the presence of two species of Rhinoceros, both furnished with two horns, and distinguished from those of the Oriental region by the absence of incisor and canine teeth. The Horse family (Zquide) is also represented by several species, and includes the peculiar group of Zebras, characterised by their beautifully striped skins. Of other Ungulates the Ele- phants, which, like the Rhinoceroses, are now peculiar to the Ethiopian and Oriental regions, have one species, which is widely different from its Indian congener. The Hyraces are mainly characteristic of this region, although one species occurs in Syria and Palestine. The Carnivora include some forms like the Lion, Leopard, and Jackal, common to the Oriental region, but likewise include certain peculiar types like the Earth-wolf (Proteles), which may be regarded as the type of a distinct family, and two species of Hyznas, which are referred by some authorities to a distinct genus (Crocuta), There is also the Hunting-dog (Lycaon), and the peculiar group of Foxes known as the Fennecs, together with Ofocyon. Bears, Wolves, and true Foxes are absent; but Civets, etc., are abundant, although not characteristic of the region. The Primates yield several very characteristic types, such as the Gorilla and the Chimpanzee (Anthropopithecus) among the Simiide, which, with the exception of the Orangs of Borneo, are the only existing large man-like Apes, and the group of Dog-faced Baboons (Cynocephalus) in the Cercopithe- cide. The genus Colobus is also a group of the latter family, absolutely characteristic of the region. Lemurs, again, occur on the continent of Africa, but the great development of this group is in the adjacent island of Madagascar, where several peculiar genera occur, and where the larger Carnivora and Ungulata are 1 Generally known as Hyomoschus, but first described as an extinct form under the above name. 100 GEOGRAPHICAL DISTRIBUTION absent. These peculiarities of the fauna of Madagascar apparently point, as previously mentioned, to its separation from the mainland before the latter was overrun by the larger types, and at a time when its chief mammals were Lemurs and Insectivores. There are two genera of Edentates, the Pangolins (Munis), and the Aard- vark (Orycteropus), the latter being peculiar. Although the foregoing groups of mammals are now so characteristic of the Ethiopian region, it cannot be too strongly insisted that their restriction to this region is, so to speak, merely a feature of the present day, and that at a late geological epoch nearly all the peculiar genera were represented in India, and many of them also in Europe. Oriental Region—The third or Oriental region is likewise of very considerable extent, and is the only one, in addition to the Ethiopian, which is the home of huge Ungulates, like Elephants and Rhinoceroses, and the large man-like Apes. A large proportion of this extensive area is occupied by tropical and subtropical forests and swamps; these being especially abundant in Burma, Southern China, Siam, and the southern ridges of the Himalaya, collectively constituting the Indo-Chinese sub-region, and also in the Indo- Malayan sub-region of the Malay peninsula and adjacent islands. In the third or Indian sub-region, comprising peninsular India, with the exception of the Carnatic, there are large tracts of open country, including some of the hottest regions in the world, parts of which form plains more or less covered with vegetation during the cooler and rainy seasons, while others are barren rocky table-lands, as in the Deccan, or arid deserts like those of parts of the Punjab and Sind. Finally, in the fourth or Cingalese sub-region, represented by the Carnatic and the island of Ceylon, we find vast areas of luxuriant forest and jungle. In the north-western desert area of the Indian sub-region the fauna includes a mixture of Palearctic and Ethiopian forms, with those characteristic of the Oriental region. Among the chief features of the mammalian fauna of this region we may notice the absence of Hippopotami and Giraffes, the greatly diminished number of Antelopes, as compared with those of Africa, and the abundance of Deer and true Pigs. The Antelopes comprise the two peculiar genera Boselaphus (Nilghai) and the typical Antilope (Black-buck), each of which is represented by only a single species, while the Deer belong to the so-called Rusine group, which is markedly different from that to which the Paleearctic Red Deer belongs. True Chevrotains (Tragulus) are peculiar to this region. The Oxen include the true Buffalo, differing in many respects from the African species of the same group, and also certain species of true Oxen, such as the Gaour and Banting, belonging to the Bibovine group, which is confined to this region. In the Perissodactyla Horses (Equus) are represented ORIENTAL REGION Iol only by a single species in the desert area of the Indian sub-region, while the two species of Rhinocerss differ from those of Africa in being furnished with canines and incisors. The Malayan Tapir is the only Old World species of its genus. The Indian Elephant differs, moreover, so markedly from its African ally that some writers regard the two as types of distinct genera, The Carnivora include the Lion, Leopard, Jackal, and Hunting-Leopard, which are common to Africa; but the Tiger is very characteristic of this region, although extending northwards into the Palearctic. Civets are abundant, comprising some peculiar genera, of which it will suffice to mention the well known Paradorurus. Wolves closely allied to the Palearctic species occur in Northern India, and there are also Foxes related to the typical species. The Dog-hke animals which hunt in packs, and are separated by some writers from Canis under the name of Cyen, occur in the present and the Palearctic region. The striped Hyena is the Indian representative of its genus. Ratels are common to this and the Ethiopian region, and constitute the genus Vellivora, The most striking feature in the Carnivorous fauna of this region, as distinguished from the Ethiopian, is, however. the presence of Bears, some of which belong to the typical genus Ursus, while one species is usually generically separated under the name of Velursus. Among the Rodents we may especially notice the abundance of the Vuride and Sziurida. In the former family we have numbers of true Mice (us), and also the peculiar genus Nesocia (Bandicoot-Rat), while in the latter both the true Squirrels (Seiurus) and the Flying-Squirrels (Pieremys) attain great develop- ment. The genus (Pizromys) is, indeed, mainly characteristic of this region, although in Kashmir and Japan it enters the Palearctic. The Bats are very numerous, being represented by all the families, with the exception of the Phallesiomatid, or Vampyres, of South America. Among the Insectivora the genera Tupaia and Gales pithecus (Flying Lemur) are peculiar to this region, although not found in Indis. Finally, in the Primates we have the genera Macacus and Sennopithzeus very abundantly represented, although both also enter the Palearctic region; but the Anthropoid types are confined to the south-eastern half of the region, and include the Orangs (Sitnia) of Borneo. and the smaller long-armed Gibbons (Hylobaies), which are abundant in the Malay peninsula, both genera not being found beyond this region. The Lemurs are much less abundant than in the Ethiopian region, but they include the peculiar Tarsier of Sumatra, Borneo, and Celebes (Austro-Malayan region), which differs so markedly in dentition and structure of the feet from all other forms that it has been made the type of a separate family. The Edentates, so poorly represented in the Old World, include only Pangolins (Afanis\, which, ss we have already seen, also occur in the Ethiopian region. 102 GEOGRAPHICAL DISTRIBUTION Australasian Region —With the fourth or Australasian region we come to a mammalian fauna so peculiar that we have no difficulty whatever in defining it from all the other regions of the globe, although it should be observed that in the Austro-Malayan islands we have a partial mingling of the Australasian and Malayan faunas. If we exclude Celebes from this region we find that, with the exception of a Pig in New Guinea, of the Dingo in Australia, of numerous Mice and Rats (Muride), and Bats, there are no Eutherian mammals throughout the area. The mammals of this region are restricted to the Australian mainland, the island of Tasmania, New Guinea, and the Aru islands, the whole area of New Zealand having been totally devoid of mammalian life until introduced by man. The whole of the Monotremata, constituting the subclass Prototheria, and all the Marsupials, exclusive of the few outlying forms ranging into the transitional Austro-Malayan area, and with the exception of the American family of the Opossums (Didelphyide), are absolutely confined to this region. Celebes.—The mammals of Celebes—the typical representative of the Austro-Malayan transitional region or sub-region—include the peculiar Ape known as Cynopithecus, Tarsius (also Oriental), the Anoa, and the single species of Babirusa. Several other types of placental mammals are found in this transitional area, while the Marsupials are represented by Phalanger and Petaurus. Nearctic Region.—The two remaining regions we have to consider are comprised in the New World. The first of these is the Nearctic, which, as already mentioned, has a fauna showing such a strongly. marked relationship to that of the Palearctic region, that it has been proposed to unite the two regions. Among types common to these two regions we may mention closely allied species of true Deer (Cervus) as exemplified by the Red Deer and the Wapiti; the allied Bisons of the two regions; the Reindeer and Elk common to both; as well as nearly related, and in some cases identical, species of Cats, Lynxes, Bears, Wolves, Foxes, Beavers, Squirrels, Marmots, and Hares. The Glutton or Wolverene, and the Musk Ox is also common to the Arctic portions of the two regions, The Ungulates are very poorly represented, but we have, in addition to the forms already mentioned, one species of the Palearctic genus Ovis, namely the Big-horn, and the Prong-buck (Antilocapra), which is quite peculiar. There are, however, no Perissodactyla. The Racoons and Coatis (Procyonide) constitute a family represented out of the New World only by the aberrant Cat-Bear (#lurus) of Nipal. The characteristic American feline known as the Puma extends over this region; but there are no Edentates, and the Marsupials are represented only by a single species of Opossum. Rodents are ex- tremely numerous, and comprise several characteristic types, which alone would tell us what part of the globe we were visiting. The NEOTROPICAL REGION 103 most distinctive are the Pouched Rats (472000) /.2r), and the Beaver-like rodents known as the Hapluvatide. True Rats and Mice (Vus, which are represented throughout the Old World, are totally wanting in the New, where they are replaced by the Vesper-mice. which may be included in the European genus Crie/‘s, although often separated as Hesveromys, This feature alone would seem to justify the dis- tinction of the Nearetic from the Palearctic region. The Musquash (fiver) is a genus of Nearetie rodents unknown in the Old World. Among other characteristic genera We may Mention, in the Carnivora, the Skunk (Mephitis) and che American Badger (Tazidea). Primates are absent from the entire region. cpinl) Fexiem—The last of the six main regions is the Neotropical, including Mexico, South America, and the West Indies. Avery large extent ‘of this area is occupied by forests. which are described as being denser and more luxuriant than those of any other part of the globe. Alternating with these forest areas are the vast grassy plains known in different rezions as Hanos, savannas, and pampas. The back-bone of the region is formed by the greaz chain of the Andes. Next to the Australasian. this region is perhaps better characterised by its mammalian fauna than any of the others. Commencing with the Un sulates, we find a total absence of Antelopes, Sheep. and Oxen, and also of all Perissodac- trles except Tapirs. Deer are, however, represented. although br peculiar forms (C2sitevs) unknown beyond the New World. The Pecearies (IMevfulzs), which are citen made the type of a discinet family, take the place of the Old World Pizs, while the Llamas and Alpacas (4ucienia) are the substitutes Tor the Palearctic Camels. The Carnivora inelude several Cats (F27is\, among which the Puma and the Jaguar are the mest noticeable; and there are also Raccoons, Coatis, Foxes, and one species of Bear. Insectivora are tozally wanting: but the Bats are characterised by the presence of the Vampyres (Piullastomatid:), which are almost restricted to this region. The Rodents likewise inelude three families unknown elsewhere, namely the Chinchillas and Viscacha aoe pies the Agouties (1). fide’, and the Cavies (Carudz); while a large number of the Cctoduntide are Neotropical. all the other forms being Ethiopian. In the Frimaces, again. we have all the forms quite peculiar to this rezion. and constituting two families. viz the Celidz or Prehensile-zatied Monkeys. and the Hevalide, or Mar- mosets, both of which differ decidedly in their dentizion, as well as in other features. from the Old World Monkeys. Lemuroids are unknown. Perhaps, however, the mammals which may be considered ag most characteristic of the Nearetic rezion are the numerous Edentates, which form three families. mostly confined to it. These the Brotypelide or Sloths. which solely inhabit the forest region; the Myrmceophamiix or Anceaters ; and 104 GEOGRAPHICAL DISTRIBUTION the Dasypodide or Armadillos, of which one species has crept northward as far as Texas. Almost equally characteristic are the numerous Opossums, the majority of which belong to the genus Didelphys. Finally, it should be observed that the West Indies are distinguished from the rest of the region by the absence of Primates, Carnivora, and Edentates. Aquatic Mammals.—Many mammals grouped for the present . purpose as terrestrial pass a great portion of their lives in brooks, lakes, or rivers, and, being dependent upon such waters for ob- taining their subsistence, are necessarily confined to their vicinity ; | but the truly aquatic mammals, or those living constantly in the water, and unable to move their quarters from place to place by land, are the orders Cetacea and Sirenia, with which may also be grouped the Seals, forming the Pinniped division of the order Carnivora. For the marine Cetacea, animals mostly of large size and endowed with powers of rapid locomotion, there are obviously no barriers to universal distribution over the surface of the earth covered by sea, except such as are interposed by uncongenial temperature or absence of suitable food. Nevertheless it was thought some years ago that the fact of a Whale or a Dolphin occurring in a sea distant from that in which it had usually been found was sufficient justification for considering it as a distinct species and imposing a new name upon it. There are now, however, so many cases known in which Cetaceans from the northern and southern seas, from the Atlantic and Pacific Oceans, present absolutely no distinguishing external or anatomical charac- ters upon which specific determination can be based that the opposite view is gaining ground; and, since some species are un- doubtedly very widely distributed, being in fact almost cosmopolitan, there seems little reason why many others should not be included in the same category. The evidence is satisfactory enough in those instances in which the intermediate regions are inhabited by the same forms ;—the cases of “ continuous areas ” of distribution. In those in which the areas of distribution are apparently discontinuous, there may be more room for doubt; but it must not be forgotten that the negative evidence is here of much less value than in the case of land animals, since the existence of Cetaceans in any particular part of the ocean may be easily overlooked. The great Sperm Whale (Physeter macrocephalus) is known to be almost cosmopolitan, in- habiting or passing through all the tropical and temperate seas, although not found near either pole. At least three of the well- known species of Rorqual (Balenoptera) of the British coasts are represented in the North Pacific, on the South American shores, and near New Zealand, by species so closely allied that it is difficult to point out any valid distinctive characters, though it may perhaps AQUATIC MAMMALS 105 be desirable to wait for a more exhaustive examination of a large series of individuals before absolutely pronouncing them to be specifically identical. There is nothing yet known by which we can separate the “Humpback Whales” (Megaptera) of Greenland, the Cape of Good Hope, and Japan. The same may be said of the common Dolphin of the European seas (Delphinus delphis) and the so-called D. bairdi of the North Pacific and D. forstert of the Australian seas. The Pilot Whale (Globicephalus melas) and the Pseudorca of the North Atlantic and of New Zealand are also, so far as present knowledge enables us to judge, respectively alike. Many other similar cases might be given. Captain Maury collected much valuable evidence about the distribution of the larger Cetacea, and, finding Right Whales (Balena) common in both northern and southern temperate seas, and absent in the intermediate region, laid down the axiom that ‘the torrid zone is to the Right Whale as a sea of fire, through which he cannot pass.” Hence all cetologists have assumed that the Right Whale of the North Atlantic (B. biscayensis), that of the South Seas (B. australis), and that of the North Pacific (B. japonica), are necessarily distinct species. The anatomical structure and external appearance of all are, however, so far as yet known, ‘marvellously alike, and, unless some distinguishing characters can be pointed out, it seems scarcely justifiable to separate them from geographical position alone; as, though the tropical seas may be usually avoided by them, it does not seem impossible, or even improbable, that some individuals of animals of such size and rapid powers of swimming may have at some time traversed so small a space of ocean as that which divides the present habitual localities of these supposed distinct species. If identity or diversity of structural characters is not to be allowed as a test of species in these cases, as it is usually admitted to be in others, the study of their geographical distribution becomes an impossibility. Although many species are thus apparently of such wide dis- tribution, others are certainly restricted; thus the Arctic Right Whale (Balena mysticetus) has been conclusively shown to be limited in its range to the region of the northern circumpolar ice, and no corresponding species has been met with in the southern hemisphere. In this case, not only temperature, but also the peculiarity of its mode of feeding, may be the cause. The Narwhal and the Beluga have a very similar distribution, though the latter occasionally ranges farther south. The common Hyperoddon is restricted to the North Atlantic, never entering, so far as is yet known, the tropical seas. Other species are exclusively tropical or austral in their range. One of the true Whalebone Whales (Neobulena marginata) has only been met with hitherto in the seas round Australia and New Zealand; and a large Ziphioid (Berardius arnouxi) only near the last-named islands. 106 GEOGRAPHICAL DISTRIBUTION The Cetacea are not limited to the ocean, or even to salt water, some entering large rivers for considerable distances, and others being exclusively fluviatile. One species of Platanista is extensively distributed throughout nearly the whole of the river systems of the Ganges, Brahmaputra, and Indus, ascending as high as there is water enough to swim in, but apparently never passing out to sea. The individuals inhabiting the Indus and the Ganges must therefore have been for long ages isolated without developing any definite distinguishing anatomical characters; for those by which the sup- posed P. indi was formerly separated from 2. ygangetica have been shown by Anderson to be of no constant value. Oreella fluminalis appears to be limited to the Irawaddy river, and at least two distinct species of Dolphin belonging to different genera are found in the waters of the upper Amazon. A Neomeris has been found in the great Chinese river, the Yang-tsi-Kiang, nearly a thousand miles from the sea. It is remarkable, however, that none of the great lakes or inland seas of the world are, according to our present knowledge, inhabited by Cetaceans. A regular seasonal migration has been observed in many of the oceanic Cetacea, especially those inhabiting the North Atlantic, but further observations upon this subject are still much needed. The great difference in the manner of life of the Sirenia, as compared with that of the Cetacea, causes a corresponding difference in their geographical distribution. Slow in their movements, and feeding exclusively upon vegetable substances, water-grasses, or fuci, the Sirenia are confined to rivers, estuaries, or coasts where these grow, and are not denizens of the open sea, although of course there is a possibility of accidental transport by the assistance of oceanic currents across considerable distances. Of the three genera exist- ing within historic times, one (J/anatus) is exclusively confined to the shores of the tropical Atlantic and the rivers entering into it, individuals scarcely specifically distinguishable being found both on the American and the African side of the ocean. The Dugong (Haticore) is distributed in different colonies, at present isolated, throughout the Indian Ocean from Arabia to North Australia. The Rhytina or Northern Sea-Cow was, for some time before its extinction, limited to a single island in the extreme north of the Pacific Ocean. The Pinnipeds, although capable of traversing long reaches of ocean, are less truly aquatic than the last two groups, always resorting to the land or to extensive ice-floes for the purpose ‘of breeding. The geographical range of the various species is generally more or less restricted, usually according to climate, as they are mostly inhabitants either of the Arctic or Antarctic seas and adjacent temperate regions, very few being found within the tropics. For this reason the northern and the southern species are for the most part GEOLOGICAL DISTRIBUTION 107 quite distinct. In fact, the only known exception is the case of a colony of the Sea-Elephant (Macrorhinus leoninus), the general range of which is in the southern hemisphere, inhabiting the coast of California. Even in this case a different specific name has been given to the northern form; but the characters by which it is distinguished are not of great importance, and probably, except for the abnormal geographical distribution, would never have been noticed. The most remarkable circumstance connected with the distribution of the Pinnipeds is the presence of members of the suborder in the three isolated great lakes or inland seas of Central Asia—the Caspian, Aral, and Baikal; these forms, notwithstanding their long isolation, having varied but slightly from species now inhabiting the Polar Seas. II. GEOLOGICAL DISTRIBUTION, Geological Sequence. —In order to understand the geological distribution, or in other words the distribution in time of mammals, it is necessary to be acquainted with the chief divisions, or time- periods, of the strata constituting the crust of the globe. These are shown in the following table, which commences with the uppermost or most recent beds and ends with the lowest and oldest. I. Cartnozoic on TERTIARY— 1. Pleistocene—River alluvia, ete. 2. Pliocene—Suffolk Crag, 3. Miocene—Hempstead Beds of Hampshire. 4, Eocene—Paris Gypsum and London Clay. II. Mesozoic or SEconDaRy— 1. Cretaceous—Chalk, Greensands, etc. 2. Jurassic—Oolites and Lias. 3. Triassic—Red Marls, Dolomites, ete. III. Patzozorc on Primary— . Permian—Beds overlying the Coal. . Carboniferous—Coal-measures, ete. . Devonian—Old Red Sandstone. . Silurian— Wenlock Limestone, ete. . Cambrian—Llanberis Slate, ete. . Archean—Gneiss and other schists. Qa Pk whe The names in the first column indicate the primary divisions or life-periods, while those in the second column are the great systems, each of which is again divided into minor groups, the popular names of a few of these minor groups being given in the third column. There are at present no means of arriving at any satis- factory conclusion as to the absolute length of time indicated by 108 GEOLOGICAL DISTRIBUTION either the primary or secondary divisions ; but there is little doubt that the whole of the Tertiary period is only equal to a fraction of the Mesozoic as regards its duration, while it is probable that the duration of the Mesozoic epoch was largely exceeded by that of the Paleozoic. Mesozoic Mammals.—The earliest date at which mammals are at present known is in the upper part of the Triassic period, which forms the base of the great Mesozoic epoch ; and from this date they are represented more or less abundantly in various horizons of the Jurassic and Cretaceous. The very rapid advances in our knowledge of these forms which have been made in the last few years, especially in consequence of the explorations of rich fossiliferous beds in North America, have not only completely changed the present aspect of the science, but give such promise for the future, that any sketch which we may now attempt of this branch of the subject can only be regarded as representing a transient phase of knowledge. It will be well, however, to gather together in this place the leading facts now ascertained with regard to the most ancient forms, as, owing to the uncertainty of their relationship with any of the existing orders, they will be most conveniently treated of separately, while the ascertained facts relating to the geological history of the forms more nearly allied to those now living will be more appropriately described under the account of the different groups into which the class may now be divided. The remains of mammals which existed anterior to the Tertiary period hitherto discovered nearly all belong to creatures of very small size, many of the largest scarcely exceeding the common Pole- cat or Squirrel. Some are known only by a few isolated teeth, others by nearly complete sets of these organs, and the majority by more or less nearly perfect specimens of the rami of the lower jaw. It is a very curious circumstance that this part of the skeleton alone has been preserved in such a large number of instances. Only very rarely has a nearly complete cranium been found; and there is no satisfactory evidence of the structure of the vertebral column of any single individual, and only one known case of a com- plete limb.!' The species already described from European strata are numerous, although the number of genera and species has lately been reduced. Of these by far the greater number have been found at a single spot near Swanage in Dorsetshire, in a bed of calcareous mud only forty feet long, ten feet wide, and averaging five inches in depth. The marvellous results obtained by the exploration by Mr. S. H. Beckles of this small fragment of the earth’s surface show by what accidents, as it were, our knowledge of the past history of life 1 The fore limb from S. Africa described as Theriodesmus, which appears to be mammalian, and may belong to T'ritylodon. MESOZOIC MAMMALS 109 has been gained, and what may still remain in store where little thought of at present. A bed, apparently equally rich, has been discovered in the Jurassic of Wyoming, North America, the contents of which have been made known by Professor Marsh, while another fertile source of these remains occurs in the Laramie beds of the Upper Cretaceous of the United States.! The whole of the Mesozoic mammals at present known may be divided into two great groups, the one characterised by a type of dentition more or less clearly resembling that found among the existing Polyprotodont Marsupials, while the other presents an altogether peculiar modification, recalling in some respects that of the Diprotodont Marsupials, although differing so decidedly as to Fic, 24.—Frontal and oral aspects of the cranium of Yritylodon longcevus; from the Karoo system of Basuto-land, South Africa, 4% natural size. (After Owen.) show that the owners of this form of dentition cannot be included in that group. Multituberculata.—The name Multituberculata has been proposed for the group exhibiting the type of dentition last mentioned, and is generally adopted, although the term Allotheria has been also suggested. The essential characteristic of the dentition of this group is the presence of a single scalpriform incisor on each side of the 1 The subjects referred to under this heading are mostly described and figured in detail in Owen’s ‘‘ Monograph of the Fossil Mammalia of the Mesozoic Forma- tions,” Paleontographical Society's Publications, 1871 ; and in various papers by Marsh, in the American Journal of Science and Arts, 1878-89. Important con- tributions to our knowledge of these forms have also been made hy Professors Cope and Osborn, and the reader should especially consult the memoir by the latter writer on the ‘‘Structure and Affinities of the Mesozoic Mammals,” published in the Journal of the Philadelphia Academy (1888), vol. ix. 110 GEOLOGICAL DISTRIBUTION lower jaw (Fig. 25) and of one larger incisor, and in some instances of one or two smaller ones in each premaxilla (Fig. 2+). These incisors are separated by an interval or diastema from the first of the premolars. The true molars, and in some instances the pre- molars (ig. 24), are characterised by having longitudinal rows of tubercles separated by one or More grooves : there being either two or three of these rows in the upper molars of -—The right ramus of the mandible of Plagicaulae those forms tty which deklesi; trom the Purbeck of Swanage, Twiee natural size. these teeth are known, i, Incisor; ne, molar ; b, coronvid process ; c, condyle. (Atte while there are, at least aii usually, only two in those of the lower jaw. In other cases the premolars ave of a secant type, with a highly convex cutting-edge, and usually cither serrated or obliquely grooved (Figs. 25, 26). 9 From a certain resemblance between these secant premolars and those of some of the smaller Jacropodid it was at one time considered that we had in these mammals representatives of Diprotodont Marsupials. The great difference in the strueture of the molar teeth of these forms, Fic. 26,—The imperfect right ramus of the Fra. * Sterecgnathus odlithicus, Frag: mandible of Migiaulae minor; from Swanage. ment of jaw with three teeth (a, d, e), it Four times natural size. p, Premolars ym, matrix; from the Stonestield Slate. Natue molars. (After Lyall.) ral size. (After Owen.) coupled with the circumstance that when the number of upper incisors is reduced helow three it is the second in place of the tirst which becomes enlarged and opposed to the incisor of the lower jaw, seems to prevent the acceptation of this view. Moreover, in their peculiar structure the molars seem, on the whole, to make a nearer approximation to the teeth of Ornithorhynchus than to any other known mammal; and it has accordingly been sueeested that the Multituberculata: may veally represent an order of Prototheria. Some support is afforded to this sugeestion by certain fragmentary bones from the Cretaceous of the United States, which are rewarded MESOZOIC MAMMALS III by Marsh as parts of a coracoid and interclavicle. The peculiar character of the whole dentition of these forms indicates that if they are really Prototherians they cannot be regarded as primitive and ancestral types. It would be beyond the scope of the present work to describe in detail, or even to mention the names of all the members of this group, and it will therefore suffice to refer to a few of the principal types. Of the forms with tubercular premolars the best known is the genus Tritylodon (Fig. 24), which occurs typically in beds of Lower Mesozoic in South Africa, but is also known from the Trias of Stuttgart. In the Stonesfield Slate, near Oxford, which belongs to the lower part of the Jurassic system, and is separated from the Trias by the intervening Lias, a fragmentary jaw with three teeth (Fig. 27) appears to indicate an allied type, the teeth having three longitudinal ridges separated by grooves. In the Purbeck beds of Dorsetshire, forming the top of the Jurassic system, we find another member of this group, which has been described as Bolodon, closely allied to which is ¢ ( 7 3 Prmy Incisors sharp and cutting, those of the lower jaw frequently having a scissor-like action against one another ; upper canine, if present, small. Penultimate premolar shed with the fourth milk-molar, which is molariform and long persistent. Molars wide, and either transversely ridged or bluntly tuberculate. Premolars and molars moving forwards in the skull as the age of the animal increases, this being most marked in the larger species. Masseteric fossa of mandible hollowed out below into a deep cavity walled in externally by a plate of bone and communicating with the inferior dental canal by a large foramen. Hind limbs usually larger than the anterior ones, antl progression generally saltatorial. Fore feet with five digits; hind feet syndactylous, the fourth digit being very large and strongly clawed ; hallux usually absent. Tail generally long and hairy, MACROPODID.E 159 oceasionally prehensile ; stomach sacculated. Pouch large and opening forwards. The JMucropodide or Kangaroos, taken as a whole, form a very well-marked family, easily distinguished from the other members of the suborder by their general conformation, and by peculiarities in the structure of their limbs, teeth, and other organs. They vary in size from that of a sheep down to a small rabbit. The head, especially in the larger species, is small, compared with the rest of the body, and tapers forward to the muzzle. The shoulders and fore limbs are feebly developed, and the hind limbs usually of disproportionate strength and magnitude, which gives them a peculiarly awkward appearance when moving about on all fours, as they occasion- ally do when feeding. Rapid progression is, how- ever, performed only by the powerful hind limbs, the animal covering the ground by a series of immense bounds, during which the fore part of the body is inclined forwards, and balanced by the long, strong, and tapering tail, which is carried horizontally backwards. When not moving they often assume a perfectly upright position, the tail aiding the two hind legs to form a sort of support- ing tripod, and the front limbs dangling hy the side of the chest. This position gives full scope for the senses of sight, hearing, and smell to warn of the approach of enemies, from which these animals save themselves by their bounding flight. The fore paws have five distinct digits, each armed with a strong curved claw. The hind foot (Fig. 52), as being a typical example of the syndactylous modification, may be — Fre. 52,—Skeleton noticed in some detail. It is extremely long and $f rsht hind foot of narrow, and (with only one exception) without any ~~" hallux or great toe. It consists mainly of one very large and strong toe, corresponding to the fourth of the human or other typically developed foot, ending in a strong, curved, and pointed claw. Close to the outer side of this lies a smaller fifth digit, and to the inner side two excessively slender toes (the second and third), bound together almost to the extremity in a common integument. The two little claws of these toes, projecting together from the skin, may be of use in scratching and cleaning the fur of the animal, but the toes themselves must have quite lost all connexion with the functions of support or progression. The dentition of the Kangaroos, functionally considered, 160 WARSUPIALIA consists of sharp-edged incisors. most fully developed near the median line of the mouth, for the purpose of cropping the various kinds of herbage on which they feed. and ridged and tuberculated molars for crushing it, there beinz no tusks or canines for offensive or defensive purposes. The number of vertebre is—in the cervical region 7, dorsal 13. lumbar 6, sacral 2. caudal varying according to the length of the tail, but generally from 21 to 25. In the fore limb the clavicle and the radius and ulna are well developed. allowing of considerable freedom cf motion of the hand. The pelvis has large epipubic or “marsupial” hones. The femur is short. and the tibia and fibula are of great length. as is the foot, the whole of which is applied to the wen Sipsciesys af The moles. da owhich the w de sand AT ayoredimahan ta the WV ant: Hags : ee samc Teamre being eee ees) by s gheeneh ar odes at the Asgorian PRocene. 8. ftom BIN daptateus ot The Panes Tabi poagepher wih Se acecuns and Semantics at the covesponsiig Paroposn ceposus. are veri Pay SASS Sa oss aS : Thesr ss ee ah oe Sar ae a tlh a oe Tay ny. No fyse HS ae RE oe GE WSs Se SN A 1a, ae OF that at Barope. are Ta alhed Sones NT Soa! See PAIAIA LO SN Cairo nen wid etch they vee Dao molar simichirs, SN oammeticusn of the Upper Phoeene Red ayers Ta he alhed to S etecanusy: whe In the ENO So ehicns> ot the PRoeene of Noi Vi ~. Incha we prohahly have the Qyoct sneesion at S xa/ranins. eS Pre VON) Bland af Rphiooen (Aon gaan ates) tilly, Sh: 3 DEES OF 3 m 2s total 34. The total wmambher af weth os therefore considerably redmeed. the earer mope: haagor and The Pwo antestar premolars of beth jis hemg abso The sesics esmedally the fosh are sander and scee or than WS yar rhe STOAD POST IATIN at this SPs: aS the ox V develop Mont ot aie eaivues of the mala Thes: : ever etowing. Jong, Sider. and carved, and entre foal @amel oes aft the wpper Jaw are Dreaded is wacds from ihe feiss Bo: thal ther pis a ae Come the month. ‘hai a the Se oes oat he t veble hovos rather than tecth. s. we has wards. and fmally offen foowicus age actias; ar qrite Weta the skin af the forchaad, Vortehee OT, DIRT UA Se Shes as at ame sneues 8 aboeus Famd only In the : Snes amd Rera. lis cvternal s.toee is ales ‘ alae Uhre Wr er nay tr R57 (e si Qo Rahirasn.” 288 UNGULATA entirely devoid of hair. With regard to the curiously modified dentition, Wallace (Malay Archipelago, vol. i. p. 435) makes the following observations: —‘“It is difficult to understand what can be the use of these horn-like teeth. Some of the old writers supposed that they served as hooks by which the creature could rest its head on a branch. But the way in which they usually diverge just over and in front of the eye has suggested the more probable idea, that they serve to guard these organs from thorns and spines while hunting for fallen fruits among the tangled thickets of rattans and other spiny plants. Even this, however, is not satisfactory, for the female, who must seek her food in the same way, does not possess them. I should be inclined to believe rather that these tusks were once useful, and were then worn down as fast as they grew, but that changed conditions of life have rendered them unnecessary, and they now develop into a monstrous form, just as the incisors of the Beaver and Rabbit will go on growing if the opposite teeth do not wear them away. In old animals they reach an enormous size, and are generally broken off as if by fighting.” Phacocherus1—The Wart-Hogs, so called from the large cutaneous lobes projecting from each side of the face, have the teeth still more remarkably modified than in Babirusa. The milk-dentition, and even the early condition of the per- manent dentition, is formed on the same general type as that of Sus, except that certain of the typical teeth are absent, the formula being 7 4, ¢ +, p 3, m %, total 34; but as age advances all the teeth have a tendency to disappear, except the canines and the posterior molars, which in some cases are the only teeth left in the jaws, and attain an extraordinary development. The upper canines especially are of great size, and curve outwards, forwards, and upwards. Their enamel covering is confined to the apex, and soon wears away. The lower canines are much more slender, but follow the same curve ; except on the posterior surface, their crowns are covered with enamel. Unlike those of the Babirusa, the canines of the Wart-Hog are large in both sexes. The third molar tooth of both jaws is of great size, and presents a structure at first sight unlike that of any other mammal, being composed of numerous (22-25) parallel cylinders or columns, each with pulp-cavity, dentine, and enamel covering, and packed together with cement. Careful examination will, however, show that a similar modification to that which has transformed the comparatively simple molar tooth of the Mastodon into the extremely complex grinder of the Indian Elephant has served to change the tooth of the common Pig into that of Phacochwrus ; and, as already mentioned, some of the fossil Indian and African species of Sus indicate the mode in which this ? Cuvier, Regne-Animal, vol. i. p, 236 (1817). ° DICOTYLID.E 289 transition came about. The tubercles which cluster over the surface of the crown of the molars of the common Pig are elongated and drawn out into columns in the Wart-Hog, as the low transverse ridges of the Mastodon’s tooth become the leaf-like plates of the Elephant’s. Two species of this genus are commonly but rather doubtfully distinguished :—P. africanus, Ailian’s Wart-Hog, widely distributed over the continent; and P. cethiopicus, Pallas’s Wart-Hog, confined to South-Eastern Africa. In specimens attributed to the latter species the dentition reaches its most complete reduction, as in adult animals the upper incisors are absent and the lower ones worn down to the roots. Family DICOTYLIDA. Snout as in Swide. Dentition: i 2, ¢ 4, p 3, m 4; total 38. Incisors rooted; upper canines directed downwards, with sharp cutting hinder edges. Toes, four on the fore feet and three on the hind feet (the fifth wanting). Stomach complex. A cecum. Confined to the New World. Dicotyles..—The teeth of the Peccaries (Dicotyles) differ from those of the true Pigs (Sus) numerically in wanting the upper outer incisor and the anterior premolar on either side of each jaw, and also in the circumstance that the last premolar is nearly as complex as the molars. The upper canines have their points directed down- wards, not outwards or upwards as in the Boars, and are very sharp, with cutting hinder edges, and completely covered with enamel until worn. The lower canines are large, directed up- wards and outwards, and slightly curved backwards. The pre- molar and molar teeth form a continuous series, gradually increasing in size from the first to the last. The true molars have square quadricuspidate crowns. The stomach is much more complex than in the true Pigs, almost approaching that of the ruminants. In the feet the two middle (third and fourth) metapodial bones, which are completely separate in the Pigs, are united at their upper ends, as in the ruminants. On the fore foot the two (second and fifth) outer toes are equally developed as in Pigs, but on the hind foot, although the inner (or second) is present, the. outer (or fifth) toe is entirely wanting, giving an unsymmetrical appearance of the member, very unusual in Artiodactyles. Vertebre: C 7, D14, L 5, 8 4, C7. As in the Pigs, the snout is truncated, and the nostrils are situated in its flat, expanded, disc-like termination. The ears are rather small, ovate, and erect; and there is no external appearance of a tail, The surface of the body is well covered with thick bristly hair, and rather behind the middle of the back is a large and 1 Cuvier, Regne Animal, vol. i. p. 237 (1817). 19 290 UNGULATA peculiar gland, which secretes an oleaginous substance with a power- ful musky odour. This was mistaken by the old travellers for a second navel, a popular error. which suggested to Cuvier the name of Dicotyles. When the animal is killed for food, it is necessary speedily to remove this gland, otherwise it will taint the whole flesh so as to render it uneatable. There are two species,! so nearly allied that they will breed together freely in captivity. Unlike the true Pigs, they never appear to produce more than two young ones at a birth. The Collared Peccary (D. tajacu, Linn., torquatus, Cuvier), Fig. 109, ranges from the Red River of Arkansas through the forest districts of a5 ae gel] Fic. 109.—The Collared Peccary (Dicotyles tujacu). Central and South America as far as the Rio N egro of Patagonia. Generally it is found singly or in pairs, or at most in small herds of from eight to ten, and is a comparatively harmless creature, not being inclined to attack other animals or human beings. Its colour is dark gray, with a white or whitish band passing across the chest from shoulder to shoulder. The length of the head and body is about 36 inches. The White-lipped Peccary or Warree (D. labiatus, Cuvier) is rather larger, being about 40 inches in length, of a blackish colour, with the lips and lower jaw white. Its range is less ex- tensive, since it is not found farther north than British Honduras or south of Paraguay. It is generally met with in large herds of from fifty to a hundred or more individuals, and is of a more pugnacious disposition than the former species, and capable of ' Professor Cope considers that there is a third species, for which he has pro- posed the name D. angularis, DICOTYLIDE 291 indiisting severe wounds with it: sharp tu:k:. A hunter who en- counters a herd cof them in a forest has often to climb a tree as his only chance of safety. Both species are omnivorous. living on roots, fallen fruits, worms, and carrion; and when they approach the neighbourhood «oi villages and cultivated lands ther ofter inflict great devastation upon the crops of the inhabitants. Remains of the two existing species of Peceary, a: well as of one much larger extinct form, are found in the cavern-deposiis of Brazil : while large Pecearies also occur in the Pleistocene of the United States. which, although ther have been referred to a distinct genus. Platygonus, on account of their relatively smaller incisors and some- what simpler premolars, may well be included in Dicstules. Allied Extinet Genert.—In the Tertiary deposits of both the Old and New World oceur remains of Pig-like animal: which, =9 far a: we can judge. appear to connect the Peccaries so closely with the true Pigs as to render the Distylida really inseparable Irom the Suma. Of these the American genus Chenohyus has the lower canine with a triangular cross ction and received into a notch in the upper jaw, a: in the Pec- caries, but the iourth upper premolar is simpler than the molars, as in the under-mentioned genus Hyctlerium. The trpical forms have ouly three premolars. but in others, which it haz been proposed ts separate generically as Bothriolalis, there are four of these teeth. Aystherium, ot the Pliocene and Miocene of the Old World, is a generalised form allied both to Sus and Dieotyles az wellas tw certain extinct genera. The upper molars (Fig. 119) are char- acterized by their square crowns, the last having no Fis. 110.—The three left upper molars of Hystherium distinet third lobe, and com- perimense, fro the Pliocene of India ing into use before the first is much worn, while the last premolar is simpler than the true molars. The canines, which have an oval section and are scarcely larger than the incisors, are not received into a notch in the upper jaw. In the Pliocene of India there occurs an apparently allied genus known as Hippohyus, in which the crowns of the molars are much taller, and have lateral inivldinys of the enamel, producing a very complex pattern on the worn crowns. The European Miocene genus Listriodon, with the dental formula i 4,¢4, p 3, m4, differs from all the preceding in having the anterior and posterior pairs of tubercles of the molar: united into ridges running across their crowns, 20 that these teeth resemble the lower molars of the Tapir. The genus is also found in the Lower Pliocene of India. 292 UNGULATA EXTINCT TRANSITIONAL ARTIODACTYLES. In this place it will be convenient to notice briefly a few of the extinct types of Tertiary Artiodactyles which connect the existing bunodont Suina with the more specialised selenodont groups mentioned below so closely as to show that in a strictly paleontological classification such groups cannot be maintained. It should be mentioned that while some of these extinct forms were in all probability actual ancestral links between the bun- odonts and selenodonts, others, like the Anoplotheres, died out entirely without giving rise to any more specialised descendants. Cheropotamide.—In this family the molars are intermediate in structure between those of the Suidw and the next family. The upper ones have very broad crowns, with the five columns arranged as in Anthracotherium , while the premolars are not secant, and may be very large. The best known forms are the small Cebochwrus of the Phosphorites of Central France ; Cheropotamus of the Upper Eocene, the type species of which was of the size of a large Pig, with the dental formula 2 3, ¢ 4,p 4, m %, and no distinctly selenodont structure in the molars; the much larger Elotherium, from the Upper Eocene and Lower Miocene of both the Old and New Worlds, which presents the very rare feature of the absence of a third lobe to the last. lower molar; and the equally large Tetraconodon of the Pliocene of India, in which this third lobe was present and the premolars were of enormous size. The remarkable North American Eocene genus Achenodon should perhaps also be placed here. Anthracothertide.—The genera Anthracotherium and Hyopotamus, of the upper Eocene and Miocene, have the typical Eutherian dental for- mula; the upper molars (Fig. 111) , carrying three columns on the anterior 7 and two on the posterior half of the crown, all of which are of a more or less decidedly selenodont structure. The mandible has a descending flange at the angle. The figured tooth (in which the antero-internal and antero- median columns are imperfect) may be compared with the diagram given in Fig. 5, p. 32, when the homology of Fic. 111.—The imperfect third lett the columns or tubercles will be at otamus git e pram Homan genee, once epparent, the broken antaro- logia Indica.) presenting the proto- conule. Some of the species are of large size, while others are comparatively small. ENXTINCY FAMILIES 205 Merucopotamus.—The genus Vervespotemus of the lower Phocene of India may be regarded as an Anthracotheroid which has lost the antero-median column to the upper molars (Fig. 112), so that these teeth are consequently quadritubereulate : and may thus be regarded typical examples of the brachy-selenodont modifica- tion of molar structure. Cofalspide.—The Miocene genus Cetileps (Ore- eden}) is the type of a large American family in which the upper molars are selenodont and usually | Fis: WA msht E E - Upper molar of Mery Ue haye four columns, while the lower canine is APprOXt- estamus mated to the incisors and its form and function Fliocene, India, assumed by the first premolar. The last upper ainaas ante: premolar is simpler than the molars. There is no flange to the angle of the mandible: and the feet have four digits. The aftinities of this peeuliar family are probably widely spread. but they may have been derived from the fnthrecctieriide, The type genus has the full Eutherian dentition, but in some of the more “specialised forms (C yelopidius) the upper incisors may be wanting, and large vaeuities occur in the lachrymal region. The g eneralised genus Preferesden, of the Upper or Uinta Eocene. has tive cusps on the upper molars, arranged as in the Luthrocetheriida. The pollex is retained in the manus of the type genus. The family may be divided into subfamilies as follows -—— I. Upper molars with four columns. 1. Orbits open, no lachrymal fossa, a diastema, the last upper premolar with two euter columns, outer wall of upper molars coneaye and inelined inwards. — Linn. Syst. Nat. 12th ed. vol. i. p. 90 (1766). ® There is much confusion as to the proper use of the names Camel and Dromedary. It is now generally accepted that the former is tho common term for all the members of the genus, and that Dromedary should be confined to the lighter and swifter breeds of the one-humped species. One of the oldest pictures of CAMELIDE 297 is quite unknown in a wild state, but it is reported that wild Bactrian Camels occur in the more remote parts of Turkestan. The latter species is found in a domesticated state throughout a large portion of Turkestan and the neighbouring region, extending as far as the Crimea in the west and to Lake Baikal and Pekin in the east. It is a heavier and more clumsy animal than the Arabian Camel, with thicker hair, shorter legs, and the feet more callous and better adapted to a hard ground. The hair is most developed upon the top of the head, neck, humps, arm, and wrist. Bactrian Camels are occasionally brought over the stupendous mountain Fic. 114.—The Bactrian Camel (Camelus_bactrianus). passes south of Yarkand to within a few days’ journey of Leh, in Kashmir territory. The Arabian Camel is commonly employed as a beast of burden in Africa and India, and has of late years been introduced into Australia for the same purpose ; it is especially valuable in crossing long stretches of arid desert from its power of existing for a con- siderable period of time without water. The female goes fully eleven months with young, and produces but a single calf at a birth, which is suckled for a whole year. In disposition the Camel is surly and subject to furious outbursts of temper, especially during the rutting season. Atsuch periods the male utters a peculiar and highly disagreeable bubbling noise in its throat, well known to all who have travelled in India with Camels as their transport. It has been said that the Camel is docile, but Palgrave observes :— the two-humped Camel extant, painted on the wall of the Chapter House of Westminster Abbey, has, however, ‘‘ Dromedary” inscribed under it. 298 ‘UNGULATA “ Tf docile means stupid, well and good; in such a case the Camel is the very model of docility. But if the epithet is intended to designate an animal that takes an interest in its rider so far as a beast can, that in some way understands his intentions, or shares them in a sub- ordinate fashion, that obeys from a sort of submissive or half-fellow- feeling with his master, like the horse or elephant, then I say that the camel is by no means docile—very much the contrary. He takes no heed of his rider, pays no attention whether he be on his back or not, walks straight on when once set agoing, merely because he is too stupid to turn aside, and then. should some tempting thorn or green branch allure him out of the path, continues to walk on in the new direction simply because he is too dull to turn back into the right road. Ina word, he is from first to last an undomesticated and savage animal, rendered serviceable by stupidity alone, without much skill on his master’s part, or any co-operation on his own save that of an extreme passiveness. Neither attach- ment nor even habit impress him; never tame, though not wide- awake enough to be exactly wild.” The two species breed together freely, and among the Yourouks of Asia Minor, hybrids, or mules, the produce generally of a male Bactrian and a female Arabian camel are preferred to either of the pure breeds. Fossil remains of Camels are found in the Pliocene of the Siwalik Hills in Northern India. These differ from the existing representatives of the genus in having a vertical ridge at the antero-external angle of the lower molars, whereby they resemble Auchenia ; their cervical vertebre are also intermediate in structure between those of the latter and the existing Camels. A fossil Camel is also found in the Pleistocene of Algeria. Auchenia.\—Dentition of adults normally: i 4, ¢ 4, p 2, m 8; total 32—one of the lower premolars may, however, be wanting. In the upper jaw there is a compressed, sharp, pointed laniariform incisor near the hinder edge of the premaxilla, followed, in the male at least, by a moderate-sized, pointed, curved true canine in the anterior part of the maxilla. The isolated canine-like premolar which follows in the Camels is not present. The teeth of the molar series, which are in contact with each other, consist of two very small premolars (the first almost rudimentary) and three broad molars, constructed gener- ally like those of Camelus. In the lower jaw the three incisors are long, spatulate, and procumbent ; the outer ones being the smallest. Next to these is a curved, suberect canine, followed after an interval by an isolated, minute, and often deciduous simple conical premolar ; then a contiguous series of one premolar and three molars, which differ from those of existing species of Camelus in having a small accessory column at the anterior outer edge. The skull generally resembles that of Camelus, the relatively larger brain-cavity and 1 Illiger, Prodromus Syst. Mamm. p. 103 (1811). CAMELIDA 299 orbits and less developed cranial ridges being due to its smaller size. The nasal bones are shorter and broader, and are joined by the premaxille. Vertebre: C 7, D 12, L 7, 8 4, C 15-20. Ears rather long and pointed. No dorsal hump. Feet narrow, the toes being more separated than in the camels, each hav- ing a distinct plantar pad. Tail short. Hairy covering long and woolly. Size (in existing forms) smaller, and general form lighter than in the Camels. At present and within historic times the Fic, 115.—Llama (Auchenia glama), from an animal living in the Gardens of the Zoological Society of London. genus is entirely confined to the western side and southernmost parts of South America, but fossil remains have been found in the caves of Brazil, in the pampas of the Argentine republic, and in Central and North America. The word Llama, sometimes spelt Lama, is the name by which the Peruvians designated one of a small group of closely allied animals, which, before the Spanish conquest of America, were the only domesticated hoofed mammals of the country, being kept, not only for their value as beasts of burden, but also for their flesh, hides, and wool,—in fact, supplying in the domestic economy of the people the place of the horse, the ox, the goat, and the sheep of the Old World. The word is now sometimes restricted to one 300 UNGULATA particular species or variety of the group, and sometimes used in a generic sense to cover the whole. Although they were often com- pared by early writers to sheep, and spoken of as such, their affinity to the camel was very soon perceived, and they were included in the genus Camelus in the Systema Nature of Linneus. They were, however, separated by Cuvier in 1800 under the name of Lama, changed by Illiger in 1811 to Auchenia (in allusion to the great length of neck, adxjv), a term afterwards adopted by Cuvier, and almost universally accepted by systematic zoologists, although there has been of late a disposition to revive the earlier name. In essential structural characters, as well as in general appear- ance and habits, all the animals of this genus very closely resemble each other, so that the question as to whether they should be considered as belonging to one, two, or more species has been one which has led to a large amount of controversy among naturalists. The question has been much complicated by the circumstances of the great majority of individuals which have come under observa- tion being either in a completely or partially domesticated state, and descended from ancestors which from time immemorial have been in like condition, one which always tends to produce a certain amount of variation from the original type. It has, however, lost much of its importance since the doctrine of the distinct origin of species has been generally abandoned. The four forms commonly distinguished by the inhabitants of South America are recog- nised by some naturalists as distinct species, and have had specific designations attached to them, though usually with expressions of doubt, and with great diffi- culties in defining their dis- tinctive characteristics. These are (1) the Llama, Auchenia glama (Linn.), or Lima peruana (Tiedemann) ; (2) the Alpaca, 4. pueos Linn.) ; (3) the Guanaco or Huanaco, 1. huenacus (Mo- Fic. 116.—Head of Vicugna, from an animal living ru oe (4) the V aeenss in the Gardens of the Zoological Society of London. d. TLcUgna (Molina), or -d. vieunna, (Cuv.) The first and second are only known in the domestic state, and are variable in size and colour, being often white, black, or piebald. The third, and fourth are wild, and of a nearly uniform light-brown colour, passing into white below. They certainly differ from each other, CAMELIDA 301 the Vicugna being smaller, more slender in its proportions, and having a shorter head (Fig. 116) than the Guanaco (Fig. 117). It may therefore, according to the usual view of species, N A\ ‘ be considered distinct. It lives in herds on the bleak and elevated parts of the mountain range bordering the region of perpetual snow, amidst rocks and precipices, occurring in various suitable localities throughout Peru, in the southern part of Ecuador, and as far south as the middle of Bolivia. Its manners very much re- semble those of the Chamois of the European Alps; and ; Vales wl a ee timid. The wool is ex- tremely delicate and soft, and highly valued for the purposes of weaving, but the quantity which each animal produces is not great. The Guanaco has an extensive geographical range, from the highlands of the Andean region of Ecuador and Peru to the open plains of Patagonia, and even the wooded islands of Tierra del Fuego. It constitutes the principal food of the Patagonian Indians, and its skin is invaluable to them, as furnishing the material out of which their long robes are constructed. It is about the size of a European Red Deer, and is an elegant animal, being possessed of a long, slender, gracefully curved neck and fine legs. Dr. Cunningham,! speaking from observation on wild animals, says :— “Tt is not easy to describe its general appearance, which combines some of the characters of a camel, a deer, and a goat. The body, deep at the breast but very small at the loins, is covered with long, soft, very fine hair, which on the upper parts is of a kind of fawn- colour, and beneath varies from a very pale yellow to the most beautiful snow-white. The head is provided with large ears, in general carried well back, and is covered with short grayish hair, which is darkest on the forehead. Occasionally the face is nearly black. Asa rule it lives in flocks of from half a dozen to several hundreds, but solitary individuals are now and then to be met with. They are very difficult to approach sufficiently near to admit of an easy shot, as they are extremely wary, but, on being disturbed, canter off at a pace which soon puts a safe distance between them 1 Natural History of the Strait of Magellan, 1871. 302 UNGULATA and the sportsman, even though he should be mounted. Despite their timidity, however, they are possessed of great curiosity, and will sometimes advance within a comparatively short distance of an unknown object, at which they will gaze fixedly till they take alarm, when they effect a speedy retreat. Their cry is very peculiar, being something between the belling of a deer and the neigh of a horse. It would be difficult to overestimate their numbers upon the Patagonian plains; for in whatever direction we walked we always came upon numbers of portions of their skeletons and detached bones.” Darwin, who has given an interesting account of the habits of the Guanaco in his Natwralist’s Voyage, says that they readily take to the water, and were seen several times at Port Valdes swimming from island to island. The Llama is only known as a domestic animal, and is chiefly met with in the southern part of Peru. Burmeister, a very com- petent writer on the subject, says that he is perfectly satisfied that it is the descendant of the wild Guanaco, an opinion opposed to that of Tschudi. It generally attains a larger size than the Guanaco, and is usually white or spotted with brown or black, and sometimes altogether black. The earliest and often-quoted account of this animal by Agustin de Zarate, treasurer-general of Peru in 1544, will bear repeating as an excellent summary of the general character and uses to which it was put by the Peruvians at the time of the Spanish conquest. He speaks of the Llama as a sheep, observing, however, that it is camel-like in shape though destitute of a hump :— “Tn places where there is no snow the natives want water, and to supply this they fill the skins of sheep with water and make other living sheep carry them; for, it must be remarked, these sheep of Peru are large enough to serve as beasts of burden. They can carry about one hundred pounds or more, and the Spaniards used to ride them, and they would go four or five leagues a day. When they are weary they lie down upon the ground; and as there are no means of making them get up, either by beating or assisting them, the load must of necessity be taken off. When there is a man on one of them, if the beast is tired and urged to go on, he turns his head round and discharges his saliva, which has an un- pleasant odour, into the rider’s face. These animals are of great use and profit to their masters, for their wool is very good and fine, particularly that of the species called Pacas, which have very long fleeces ; and the expense of their food is trifling, as a handful of maize suffices them, and they can go four or five days without water. Their flesh is as good as that of the fat sheep of Castile. There are now public shambles for the sale of their flesh in all parts of Peru, which was not the case when the Spaniards came first 3 for CAMELIDA 303 when one Indian had killed a sheep his neighbours came and took what they wanted, and then another Indian killed a sheep in his turn.” The disagreeable habit here noticed of spitting in the face of persons whose presence is obnoxious is common to all the group, as may be daily witnessed in specimens in confinement in the menageries of Europe. One of the principal labours to which the Llamas were subjected at the time of the Spanish conquest was that of bringing down ore from the mines in the mountains. Gregory de Bolivar estimated that in his day as many as three hundred thousand were employed in the transport of the produce of the mines of Potosi alone; but since the introduction of horses, mules, and donkeys the importance of the Llama as a beast of burden has greatly diminished. The Alpaca, though believed by many naturalists to be a variety of the Vicugna, is more probably, like the Llama, derived from the Guanaco, having the naked callosities on the hind limbs, and the relatively large skull of the latter, It is usually found in a domesticated or semi-domesticated state, being kept in large flocks which graze on the level heights of the Andes of southern Peru and northern Bolivia at an elevation of from 14,000 to 16,000 feet above the sea-level, throughout the year. It is smaller than the Llama, and, unlike that animal, is not used as a beast of burden, but is valued only for its wool, of which the Indian blankets and ponchas are made. Its colour is usually dark brown or black. Mention has already been made of the occurrence of fossil Llamas in America, but some diversity of view obtains as to the generic position of some of these forms, owing to variations in their dental formula. Remains apparently referable to the existing species occur in the cavern-deposits of Brazil. In the Pleistocene of Mexico we meet with 4. (Palauchenia) magna, which attained the size of a Camel, and had always two, and occasionally three, lower premolars ; while in one South American Pleistocene species, which has been generically separated as Hemiauchenia, there were invariably three premolars in each jaw. In A. (Holomeniscus) hesterna, from the Pleistocene of North America, which was equal in size to A. magnu, the premolars were reduced to one in each jaw; and the same condition obtains in 4. (Eschatius) vitakeriana, where, however, the upper one is of simpler structure. Extinct Cameloids.— Until within the last few years the existence of two genera having so very much in common as the Camels and the Llamas, and yet so completely isolated geographically, had not received any satisfactory explanation ; for the old idea that they in some way “represented” each other in the two hemispheres of the world was a mere fancy without philosophical basis. The dis- coveries made mostly within the past twenty years of a vast and 304 UNGULATA previously unsuspected extinct fauna in the American continent of the Tertiary period, as interpreted by Leidy, Cope, Marsh, and others, has thrown a flood of light upon the early history of this family, and upon its relations to other mammals. There have been found in these regions many Camel-like animals exhibiting different generic modifications; and, what is more interesting, a gradual series of changes, coinciding with the antiquity of the deposits in which they are found, have been traced from the thoroughly differentiated species of the modern epoch down through the Pliocene to the early Miocene beds, where, their characters having become by degrees more generalised, they have lost all that specially distinguishes them as Camelidw, and are merged into forms common to the ancestral type of all the other sections of the Artiodactyles. Hitherto none of these annectant forms have been found in any of the fossiliferous strata of the Old World; and it may therefore be fairly surmised (according to the evidence at present before us) that America was the original home of the Tylopoda, and that the true Camels have passed over into the Old World, probably by way of the north of Asia, where we have every reason to believe there was formerly a free communica- tion between the continents, and then, gradually driven southward, perhaps by changes of climate, having become isolated, have under- gone some further special modifications; while those members of the family that remained in their original birthplace have become, through causes not clearly understood, restricted solely to the southern or most distant part of the continent. The occurrence in the dentition of the fossil Siwalik Camels of a feature now found only in Auchenia is especially interesting from this point of view. Briefly referring to some of these fossil types, we may note that Pliauchenia, of the Loup Fork beds (Lower Pliocene) of the United States, has three lower premolars, while in Procamelus there were four of these teeth. In Protolabis of the Miocene we have a more generalised form, in which the dental formula isi 2, c4,p 4, m3; and from this type a transition may be traced to Poébrotherium, which, while having the same dental formula, was no larger than a Fox, and had the third and fourth - metacarpals separate, with rudiments of the fourth and fifth. The earliest undoubted representative of the group is Leptotragulus, of the Uinta Eocene, which appears to have been closely allied to Poebrotherium. It is, however, probable that the first lower pre- molar was wanting; while the other premolars of the mandible were much shorter antero-posteriorly than in the last-named genus. The manus, moreover, appears to have been less reduced, the second metacarpal retaining its connection with the magnum. It is suggested that Leptotragulus may have been derived from the TRAGULIDE 305 Bunodont genus Homacodon of the Bridger Eocene, mentioned among the Cenotherude. TRAGULINA. Family TRAGULIDA. No teeth in premaxille. Upper canines well developed, especi- ally in the males; narrow and pointed. Lower canines incisiform. No caniniform premolars in either jaw, all the premolars except the last in the upper jaw being secant. Molariform teeth in a con- tinuous series, consisting of p #, m 3. Odontoid process of axis vertebra conical. Fibula complete. Four complete toes on each foot. The middle metapodials generally confluent, the outer ones (second and fifth) very slender but complete, i.e. extending from the carpus or tarsus to the digit. Navicular, cuboid, and ectocunei- form bones of tarsus united. Tympanic bull of skull filled with cancellar tissue. No frontal appendages. Ruminating, but the stomach with only three distinct compartments, the maniplies or third cavity of the stomach of the Pecora being rudimentary. Placenta diffused. This section is represented only by the single family Tragulide, containing a few animals of small size, commonly known as Chevrotains, intermediate in their structure between the Deer, the Camels, and the Pigs. The large size of the canines of the male and the absence of horns caused them to be associated formerly with foschus, one of the Cervide ; hence they are often spoken of as “Pigmy Musk-Deer,” although they have no musk-secreting gland, or, except in the above-named trivial external characters, no special affinities with the true Musk-Deer. There has scarcely been a more troublesome and obdurate error in zoology than in this association of animals so really distinct. It has been troublesome, not only in preventing a just conception of the relations of existing Artiodac- tyles, but also in causing great confusion and hindrance in paleonto- logical researches among allied forms; and most obdurate, inasmuch as all that has been recently done in advancing our knowledge of both groups has not succeeded in eradicating it, not only from nearly every one of our zoological text-books, whether British or Continental, but even from works of the highest scientific pre- tensions. The family is now generally divided into two genera. Tragulus,) containing the smallest of the existing Ungulates, animals having more of the general aspects and habits of some Rodents, as the Agoutis, than of the rest of their own order. The best-known species are 7. javanicus, T. napu, T. stanleyanus, and 1 Pallas, Spicilegia Zoologica, vol. xiii. p. 27 ita) 20 306 UNGULATA LT. memmina. The first three are from the Malay Peninsula, or the islands of the Indo-Malayan Archipelago, the last from Ceylon and India. A fossil species occurs in the Pliocene of the latter country. Dorcatheriwm+ is distinguished chiefly by the feet being stouter and shorter, the outer toes better developed, and the two middle metacarpals not ankylosed together. Its dental formula (as that of Tragulus) is usually 7 9, ¢4, p 3, m#% =34. Vertebre: C 7, D138, L 6,85, C 12-13. The only existing species, D. aquaticwm (Fig. 118), from the west coast of Africa, is rather larger than any Fic. 118.—The African Water-Chevrotain (Dorcatherium aquaticum). of the Asiatic Chevrotains, which it otherwise much resembles, but it is said to frequent the banks of streams, and have much the habits of Pigs. It is of a rich brown colour, with back and sides spotted and striped with white. It is evidently the survivor of a very ancient form, as remains of the type species (D. nani), only differing in size, occur in the lower Pliocene and Miocene of Europe ; fossil species are also found in the Indian Pliocene. In D. nawi there are, at least frequently, four lower premolars while the existing species has but three of these teeth. Extinct Traguloids.—A number of small selenodont Artiodactyles » Kaup, Ossemens Fossiles de Darmstadt, pt. 5, p. 92 (1836). This name which was proposed for a fossil species, antedates Ayomoschus, Gray, applied to the living form. PECORA 307 from various Miocene and Pliocene deposits appear to connect the modern Tragulina so closely with Gelocus (p. 294), and thus with the ancestral Cervidw, that their classification is almost an impossi- bility. Thus Leptomeryx, from the Miocene of the United States, is regarded as a Traguloid, having four premolars in each jaw and with the metatarsals fused into a cannon-bone. Prodremotherium, of the Upper Eocene Phosphorites of France, differs in that the metacarpals also form a cannon-bone; while in the American Hypertragulus, both metacarpals and metatarsals remain separate. Bachitherium, of the French Phosphorites, apparently presents affinity with Gelocus, Prodremotherium, and Dorcatherium. In this genus the first of the four lower premolars assumes the character and function of a canine, the true canine being incisor-like, and there are traces of minute upper incisors. PECORA, OR COTYLOPHORA. No premaxillary teeth or caniniform premolars. Upper canines generally absent, though sometimes largely developed. Inferior incisors, three on each side with an incisiform canine in contact with them. Molariform teeth consisting of p 3, m %, in con- tinuous series. Auditory bulle simple and hollow within. Odon- toid process in the form of a crescent, hollow above. Distal extremity of the fibula represented by a distinct malleolar bone of peculiar shape, articulating with the outer surface of the lower end of the tibia. Third and fourth metacarpals and metatarsals con- fluent. Outer or lateral toes small and rudimentary, or in some cases entirely suppressed ; their metapodial bones never complete in existing forms. Navicular and cuboid bones of tarsus united. Horns or antlers usually present, at least in the male sex. Left brachial artery arismg from a common innominate trunk, instead of coming off separately from the aortic arch as in the preced- ing sections. Stomach with four complete cavities. Placenta cotyledonous.1 The Pecora or true Ruminants form at the present time an extremely homogeneous group, one of the best-defined and most closely united of any of the Mammalia. But, though the original or common type has never been departed from in essentials, varia- tion has been very active among them within certain limits; and the great difficulty which all zoologists have felt in subdivid- ing them into natural minor groups arises from the fact that the changes in different organs (feet, skull, frontal appendages, teeth, cutaneous glands, ete.) have proceeded with such apparent irregularity and absence of correlation that the different modifica- 1 For the anatomy of this group see A. H. Garrod, Proc. Zool. Soc. 1$77, p. 2. 308 UNGULATA tions of these parts are most variously combined in different members of the group. It appears, however, extremely probable that they soon branched into two main types, represented in the present day by the Cervide and the Bovide,—otherwise the antlered and horned Ruminants. Intermediate smaller branches produced the existing Musk-Deer and Giraffe, as well as the extinct Helladotherium inclining to the first-named group, and the extinct Sivatherium, Brahmatherium, Hydaspitherium, and others more allied to the latter, although upon the true relationship of these forms there is a difference of opinion. The earliest forms of true Pecora, as Paleomeryx, generally had no frontal appendages, and some few forms continue to the present day in a similar case. In the very large majority, however, either in both sexes or in the male only, a pair or occasionally two pairs (Tetraceros and the extinct Sivatherium) of processes are de- veloped from the frontal bones as weapons of offence and de- fence, these being almost always formed on one or other of two types. 1, “ Antlers” are outgrowths of true bone, covered during their growth with vascular, sensitive integument coated with short hair. When the growth of the antler is complete, the supply of blood to it ceases, the \w, skin dies and peels off, leaving sy the bone bare and _insensible, and after a time, by a process of absorption near the base, it becomes detached from the skull Fic. 119.—A shed right antler of the Red Deer and is “ shed” (Fig. 119). A (Cervus elaphus), found in an Trish lake. a, Brow more or less elongated portion tine; b, bez tine; ¢, tres tine; @d, crown or royal a3 . ” . ue (Adler Omen or “pedicle” always remains on the skull, from the summit of which a new antler is developed. In the greater number of exist- ing species of Deer this process is repeated with great regularity at the same period of each year. The antler may be simple, straight, subcylindrical, tapering and pointed, but more often it sends off one or more branches called “tines” or “snags” (Fig. 119). In this case the main stem is termed the “beam.” Commonly all the branches of the antler are cylindrical and gradually tapering. Sometimes they are more or less expanded and flattened, the PECORA 309 antler being then said to be “palmated.” In young animals the antlers are always small and simple, and in those species in which they are variously branched or palmated, this condition is only gradually ac- quired in several successive annual growths. An interesting paral- \ lel has been ob- served here, as & in so many other cases, between the development of the race and that of the in- dividual. Thus the earliest known forms of Deer, those of the Lower Mio- cene, generally have no antlers, as in the young of the existing species. The Deer of the Middle Miocene have simple ant- lers, with not more than two branches, as in existing Deer of the second year ; but it is not until the Pliocene and Pleistocene times that Deer occur with antlers developed with that luxuriance of growth and beauty of form characteristic of some of the existing species in a perfectly adult state. Among recent Cervidw, antlers are wanting in the genera Moschus and Hydropotes ; they are present in both sexes in Tarandus (the Reindeer), and in the male sex only in all others. In those forms with the most complex antlers (Figs. 119, 120) the tine immediately over the forehead is termed the brow tine, the next one the bez tine, and the third one the fres tine; the mass of points at the summit of the antler being termed either the royal and surroyal tines, or collectively the crown. The nodulated bony ring at the base of the antler, just above the point at which it separates from the pedicle when it is shed, is termed the burr. Fic. 120.—Head of Deer (Cervus schomburgki), showing antlers. From Sclater, Proc. Zool. Soc. 1877, p. 682. 310 UNGULATA 2. The horns of the Bovide consist of permanent, conical, usually curved bony processes, into which air-cells continued from the frontal sinuses often extend, called “horn- cores,” ensheathed in a case of true horn, an epider- mic development of fibrous struc- ture, which grows continuously, though slowly, from the base, and wears away at the apex, but is very rarely shed entire. The only existing species in which the latter process occurs regularly and periodically is the American Prong-Buck (dntilocapra), in which the horns also differ from those of all others in being bifure- ated. Horns are not present at birth, but begin to grow very soon afterwards. The Fic, 121.—Head of Antelope (Gazella granti), showing horns. From 7 Sir V. Brooke, Proc. Zool. Soc. 1878, p. 724, males of all pages ing Bovide possess them, and they are also present (though usually not so fully developed) in the females of all except the genera Boselaphus, Strepsiceros, Tragelaphus, Antilope, AEpyceros, Saiga, Cobus, Cervicapra, Pelea, Nanotragus, Neotragus, Cephalophus, and Tetraceros y as well as in some species of Gazella, such as G. picticaudata and G. walleri. Another character by which different members of the Pecora can be distinguished among themselves is derived from the nature of the molar teeth. Although there is nothing in the general mode and arrange- ment of the enamel-folds, or in the accessory columns, absolutely distinctive between the two principal families, existing species may PECORA 311 generally be distinguished, inasmuch as the true molars of the Cerridi are more or less brachydont, and those of the Dovid generally hypsodont, ie. the teeth of the former have comparatively short crowns (Fig. 122), which, as in most mammals, take their place at once with the neck (or point where the crown and root join) on a level with or a little above the alveolar border, and remain in this position throughout the animal’s life; whereas in the other forms (Fig. 123), the crown beinglengthened bs and the root small, the neck docs not come up — Fis. 122.—Crown sur. to the alveolar level until a considerable part a eee of the surface has worn away, and the crown of ensis,to shew brachydont the tooth thus appears for the greater part of type. (From the Patwonto- the animal's life partially buried in the socket. cuits In this form of tooth (which is almost always most developed in the posterior molars of the permanent series) the constituent columns of the crown are neeessarily nearly parallel, whereas Pia. 128.—Inner and outer aspeets of an almost unworn left upper molar of the Nilghai (Roselaphus tragoviamelus), to show hypsodont type. (From the Paleontologia Indica.) in the first-deseribed they diverge from the neck towards the free or grinding surface of the tooth. In the completely hypsodont form the interstices of the lengthened columnar folds of enamel and dentine ave filled up with eement, which gives stability to the whole organ, and is entirely or nearly wanting in the short- crowned teeth. The same modifieation from low to high crowns without essential alteration of pattern is seen in an even still more marked manner in some of the Perissodactyle Ungulates, the tooth of the Horse bearing to that of -nchitheriun the same relation as that of an Ox does to the early selenodont Artiodactyles. 312 UNGULATA A parallel modification has also taken place in the molar teeth of the Proboscidea. As the hypsodont tooth is essentially a modification of, and, as it were, an improvement upon, the brachydont, it is but natural to expect that all intermediate forms may be met with. Even among the Deer themselves, as pointed out by Lartet, the most ancient have very short molars, and the depressions on the grinding surface are so shallow that the bottom is always visible ; while in the Cerzidu of the more recent Tertiary periods, and especially the Pleistocene and living species, these same cavities are so deep that whatever be the state of the dentition the bottom cannot be seen. Some existing Deer, as the Axis, are far more hypsodont than the majority of the family ; and, on the other hand, many of the Antelepes (as Tragelaphus) retain much of the brachydont character, which is, however, completely lost in the more modern and highly specialised Sheep and Oxen. Fic. 124.—Stomach of Ruminant opened to show internal structure. a, Gsophagus ; D, rumen or paunch ; ¢, reticulum or honey-comb bag; d, psalterium or many plies ; e, abomasum or reed; f, duodenun. The complicated stomach of the Pecora (Fig. 124), which is necessary for the performance of the peculiar function known as “chewing the cud”—a function common also to the Tragulina and Tylopoda—is divided into four well-defined compartments, known as (1) the Rumen or Paunch, (2) the Reticulum or Honey- comb Bag, (3) the Psalterium or Manyplies, (4) the Abomasum or Reed. The paunch is a very capacious receptacle, shaped like a blunted cone bent partly upon itself. Into its broader base opens the oesophagus or gullet at a spot not far removed from its wide orifice of communication with the second stomach or honey- comb bag. Its inner walls are nearly uniformly covered with a pale mucous membrane, which is beset with innumerable close-set short, and slender villi, resembling very much the “pile” on velvet The honey-comb bag is very much smaller than the paunch. CERVIDE 313 It is nearly globose in shape, and receives its name on account of the peculiar arrangement of its mucous membrane which forms shallow hexagonal cells all over its inner surface. Running along its upper wall there is a deep groove, coursing from the first to the third stomach. This groove plays an important part in the act of rumination. Its walls are muscular, like those of the viscus with which it is associated, which allows its calibre to be altered. Some- times it completely closes round so as to become converted into a tube by the opposition of its edges. At others it forms an open canal. The manyplies is globular in form, and its lining membrane is raised into longitudinal folds or lamine arranged very much like the leaves of a book, and very close together. Their surfaces are roughened by the presence of small projections or papille. The reed is the proper digestive stomach, corresponding with the same organ in man. Its shape is somewhat pyriform, and its walls are formed of a smooth mucous membrane, which secretes the gastric juice. When the food is first swallowed it is conveyed into the paunch, and after undergoing a softening process there it is regurgitated into the mouth, and undergoes a further trituration by the molar teeth and mixture with the secretion of the salivary and buccal glands. It is then swallowed again, but now passes directly through the before-mentioned groove into the manyplies, and, after filtering through the numerous folds of the lining membrane of this cavity, finally reaches the fourth or digestive stomach. The placenta of the Pecora is characterised by the foetal villi being collected into groups or cotyledons, which may present either a convex or a concave surface to the uterus. These cotyledons are received into permanent elevations in the mucous membrane of the uterus, the surfaces of which present a curvature which is the reverse of the cotyledons. Family CERVIDA. Frontal appendages, when present, in the form of antlers. First molar, at least, in both jaws brachydont. Two orifices to the lachrymal duct, situated on or inside the rim of the orbit. An antorbital or lachrymal vacuity of such dimensions as to exclude the lachrymal bone from articulation with the nasal. Upper canines usually present in both sexes, and sometimes attaining a very great size in the male (see Fig. 134). Lateral digits of both fore and hind feet almost always present, and frequently the distal ends of the metapodials. Placenta with few cotyledons. Grall-bladder absent (except in Moschus). This family contains numerous species, having a wide geographical distribution, ranging in the New World from the Arctic 3I4 ' UNGULATA Circle as far south as Chili, and in the Old World throughout the whole of Europe and Asia, though absent in the Ethiopian and Australian regions. It may be divided into two subfamilies. Subfamily Mosechinze.—This subfamily is represented solely by the Musk-Deer, which differs so remarkably from the true Deer that it is considered by several writers as the representative of a separate family. The late Professor Garrod even suggested that it should be regarded as an extremely aberrant member of the Bovide. Moschus.1—The Musk-Deer (Fig. 125) in many respects stands by SS h Fic. 125.—The Musk-Deer (Moschus moschiferus). itself as an isolated zoological form, retaining characters belonging to the older and more generalised types of ruminants before they were distinctly separated into the horned and the antlered sections now dominant upon the earth. One of these characters is that both sexes are entirely devoid of any sort of frontal appendage. In this, however, it agrees with one existing genus of true Deer (Hydropotes) ; and, as in that animal, the upper canine teeth of the males are remarkably developed, long, slender, sharp pointed, and gently curved, projecting downwards out of the mouth with the ends turned somewhat backwards. Vertebre: C7, D 14,L5,8 5, C6. Among the anatomical peculiarities in which it differs from all true Deer and agrees with the Bovide is the presence of a gall- * Linn. Syst. Nat. 12th ed. vol. i. p. 91 (1766). CERVID.E 315 bladder. The hemispheres of the brain are but slightly convoluted, and the cotyledons of the placenta are arranged in a peculiar linear manner.? Although, owing to variations of colour presented hy different individuals in different localities and seasons, several nominal species have been described, zoologists are now generally agreed that there is but one, the J/osehus moschiferus of Linneus. In size it is rather less than the European Roe Deer, being about 20 inches high at the shoulder. Its limbs, especially the hinder ones, are long. “The feet are remarkable for the great development of the lateral pair of hoofs, and for the freedom of motion they all present, so that they appear to have the power of grasping projecting rocky points, —a power which must be of great assistance to the animal in steadying it in its agile bounds among the crags of its native haunts. The ears are large, and the tail quite rudimentary. The hair covering the body is long, coarse, and of a peculiarly brittle and pith-like character, breaking with the application of an extremely slight force; it is generally of a grayish-brown colour, sometimes inclined to yellowish- red, and often variegated with lighter patches. The Musk-Deer has a wide distribution over the highlands of central and eastern Asia, including the greater part of southern Siberia, and extends to Kashmir on the south-west and Cochin-China on the south-east, always, however, at considerable elevations,—being rarely found in summer below 7000 feet above the sea-level, and ranging as high as the limits of the thickets of birch or pines, among which it mostly conceals itself in the day-time. It is a hardy, solitary, and retiring animal, chiefly nocturnal in its habits, and almost always found alone, rarely in pairs, and never in herds. It is exceedingly active and sure-footed, having few equals in traversing rocky and precipitous ground; and it feeds on moss, grass, and leaves of the plants which grow on the mountains among which it makes its home. Most of the animals of the group to which the Musk-Deer helongs, in fact the large majority of mammals, have some portion of the cutaneous surface peculiarly modified and provided with glands seeveting some odorous and oleaginous substance specially characteristic of the species. This, correlated with the extraordin- ary development of the olfactory organs, appears to offer the princi- pal means by which animals in a state of nature become aware of the presence of other individuals of their own species, or of those inimical to them, even at very great distances, and hence it is of extreme importance both to the well- being of the individual and to the continuance of the race. The situation of this specially modified portion of skin is extremely various, sometimes between the toes, as in Sheep, sometimes on the face in front of the eyes, as in many 1 For the anatomy of Aoschus sce Flower, Prov. Zool. Sve. 1875, p. 159 ; and Garrod, ibid, 1877, p. 287. 316 UNGULATA Deer and Antelopes. Sometimes it is in the form of a simple depres- sion or shallow recess, often very deeply involuted, and in its fullest state of development it forms a distinct pouch or sac with a narrow tubular orifice. In this sac a considerable quantity of the secretion can accumulate until discharged by the action of a compressor muscle which surrounds it. This is the form taken by the special gland of the Musk-Deer, which has made the animal so well known, and has proved the cause of an unremitting persecution to its possessor. It is found in the male only, and is a sac about the size of a very small orange, situated beneath the skin of the abdomen, the orifice being immediately in front of the preputial aperture. The secretion with which the sac is filled is of dark-brown or chocolate colour, and when fresh described as being of the consist- ence of “moist gingerbread,” but becoming dry and granular after keeping. It has a peculiar and very powerful scent, which when properly diluted and treated forms the basis of many of our most admired perfumes. When the animal is killed the whole gland or “pod” is cut out and dried, and in this form reaches the market of the Western World, chiefly through China. Subfamily Cervinse.—This subfamily includes all the true Deer. According to the arrangement proposed by Sir V. Brooke! the. existing Cervine may be divided into the sections Plesiometacarpalia and Telemetacarpalia. Plesiometacarpalia.—In this section, which is mainly character- istic of the Old World, the proximal portions of the lateral (second and fifth) metacarpals persist, and the vomer is never so ossified as to divide the posterior osseous nares into two distinct passages. The premaxille nearly always articulate with the nasals. Cervulus.2—Antlers half the length of the head, placed on pedicles nearly equal to them in length. Brow tine short, inclined inwards and upwards; terminal extremity of beam unbranched, and curved downwards and inwards. Lachrymal fossa of skull very large, and extending into facial part of jugal; lach- rymal (antorbital) vacuity moderate. Ascending portion of pre- maxille at least as long as nasals. A permanent ridge extending from each pedicle over the orbit, lachrymal fossa and vacuity. Auditory bulla much inflated. Upper canines of males very large. Ectocuneiform united with naviculo-cuboid of tarsus. No traces of the phalanges of the lateral digits. The native name Muntjac has been generally adopted in European languages for a small group of Deer indigenous to the southern and eastern parts of Asia and the adjacent islands, which are separated by very marked characters from all their allies. They are also called “Kijang” or “Kidjang,” and constitute the genus 1 Proc. Zool. Soe. 1878, p. 889. 2 De Blainville, Bull. Soc. Philom. 1816, p. 74. CERVID.E 317 Cerrulus of Blainville and most zoologists :—Stulocervs of Hamilton- Smith, and Prox of Ogilby. They are all of small size compared with the majority of Deer, and have long bodies and rather short limbs and neck. The antlers, which as in most Deer are present in the male only, are small and simple, and the main stem or beam, after giving off a very short brow tine, inclines backwards and up- wards, is unbranched and pointed, and when fully developed curves inwards and somewhat downwards at the tip. These small antlers are supported upon pedicles or permanent processes of the frontal bones, longer than in any other Deer, and the front edges of which are continued downwards as strong ridges passing along the sides of the face above the orbits, and serving to protect the large supra- orbital glands lying on their inner sides. The lachrymal fossa of the skull, in which is lodged the large suborbital gland or crumen, is of great depth and extent. The upper canine teeth of the males are strongly developed and sharp, curving downwards, backwards, and outwards, projecting visibly outside the mouth as tusks, and loosely implanted in their sockets. In the females they are very much smaller. The limbs exhibit several structural peculiarities not found in other Deer. The lateral digits of both fore and hind feet are very little developed, the hoofs alone being present and their bony supports (found in all other Deer) wanting. There is a tufted gland on the outer side of the metatarsus. The Muntjacs are solitary animals, very rarely even two being seen together. They are fond of hilly ground covered with forests, in the dense thickets of which they pass most of their time, only coming to the skirts of the woods at morning and evening to uvaze. They carry the head and neck low and the hind-quarters high. their action in running being peculiar and not very elegant, somewhat resembling the pace of a sheep. Though with no power of sustained speed or extensive leap, they are remarkable for flexibility of body and facility of creeping through tangled underwood. They are often called by Indian sportsmen “ Barking Deer,” a name given on account of ‘their alarm cry, a kind of short shrill bark, like that of a fox but louder, which may often be heard in the jungles they frequent both by day and by night. When attacked by dogs the males use their sharp canine teeth with great vigour, inflicting upon their opponents deep and even dangerous wounds. There is some difference of opinion among zoologists as to the number of species of the genus Cerrulus. Sir Victor Brooke, who investigated this question in 1878 (see Preecedings of the Zcological Society of Londen for that year, p. 898). came to the conclusion that there are certainly three which are quite well marked, vizi— C. muntja: (Fig. 126), found in British India, Burma, the Malay Peninsula, Sumatra, Java, Hainan, Banca, and Borneo. The general 318 UNGULATA colour is a bright yellowish-red, darker in the upper parts of the back ; the fore legs from the shoulder downwards and the lower part of the hind legs, dark bluish-brown ; anterior parts of the face from the muzzle to between the eyes, brown—a blackish line running up the inside of each frontal ridge; chin, throat, inside of hind legs, and under surface of tail white. The female has a black bristly tuft of hair on the spot from which the pedicles of the antlers of the male grow. The average length of the male, according to Jerdon, is 34 feet, tail 7 inches, height 26 to 28 inches. The female is a little smaller. The specimens from Java, Sumatra, and Borneo are Fic. 126.—The Muntjac (Cervulus muntjac). of larger size than those from the mainland, and may possibly be of distinct species or race. C. lacrymans of Milne-Edwards, or Sclater’s Muntjac of Swin- hoe, from Moupin, and near Hangchow, China. C. reevesi, a very small species from southern China. Subsequently the name C. crinifrons has been applied to a Munt- jac from Ningpo, China, readily distinguished from all other species by its bushy forehead and long tail. Another species from Tenas- serim has been described as C. fea. ; Small Deer from the European Pliocene have been provisionally referred to Cervulus, but the so-called Prox furcatus, of the Miocene is now included in Paleomeryz. : Elaphodus..—Antlers very small, unbranched, supported on long, 1 Milne-Edwards, Nouv. Arch. du Muséum, vol. vii. Bull. p. 93 (1872). CERVIDAS 319 slender, converging pedicles. Ascending rami of premaxille shorter than nasals. No supraorbital ridges or frontal glands. Upper canines of male long, but not everted. A distinct frontal tuft of hair. Other characters as in Cervulus. This genus (which has also received the name of Lophotragus) is represented by a small Deer (Fig. 127) from China of about the same size as the Indian Muntjac. The male has minute simple antlers and very large canine teeth. There are no supraorbital SS i Fig. 127.—Male of Elaphodus michianus. From Selater Proc. Zool. Soc. 1876, p. 278. glands, nor is there a tufted gland on the metatarsus. The limbs have the same peculiarities as in Cervudus, but the mesocuneiform may also ankylose with the ectocuneiform, and traces of the meta- carpals may remain. The hair is coarse and somewhat quill-like. Cervus.1—The great majority of the Deer of the Old World may be included in this large genus, which is one not easy of definition. The antlers of the male are, however, large, and two or three times the length of the head, and may be either rounded or palmate ; the canines are never large; the ectocuneiform of the tarsus remains distinct from the naviculo-cuboid ; the lateral digits are represented by their phalanges ; and the skull does not carry prominent frontal ridges. Vertebre: C7, D13,L6,S 4, C11-14. The size of the 1 Linn. Syst. Nad. 12th ed. vol. i. p. 92 (1766). 320 UNGULATA lachrymal fossa and vacuity, and the degree of inflation of the audi- tory bulla, are subject to variation in the different groups into which the genus may be divided. The Rusine group is characteristic of the Oriental region, where it is typically represented by the Sambur (C. aristotelis) of India, Burma, and China. The antlers are rounded, and often strongly grooved, without a bez tine, and with the beam simply forked at the extremity, upright, and but slightly curved; the angle formed by the brow tine, which rises close to the burr, being acute. The molars are markedly hypsodont, with small accessory columns. The lachrymal fossa is deep and the vacuity large; the auditory bulla is slightly inflated and rugose. Tail moderate ; neck maned. The Sambur, which is abundant in hilly districts, is a fine animal, standing nearly 5 feet in height, and of massive build; the general colour being deep brown. C. equinus, of Borneo, Sumatra, and Singapore, C. swinhoei, of Formosa, C. philippinus, and C. alfredi of the Philippines, are closely allied species, of which the two latter are of smaller dimensions. The Indian Hog Deer (C. porcinus) is a still smaller form, not larger than the Roe. C. hippelaphus of Java, C. timoriensis, and C. moluccensis are distinguished by the posterior branch of the beam of the antler being considerably larger than the anterior. The Rucervine group is another strictly Oriental one, and is represented by the Swamp Deer (C. duvaucelli) of India, the closely allied C. schomburgki of Siam, of which the antlers are shown in Fig. 119 (p. 309), and C. edi of Burma and Hainan. The beam of the antler is somewhat flattened, and more curved than in the Rusine group ; the large brow tine is given off from the beam at an obtuse angle and curves upwards ; the beam bifurcates into two branches, which again divide. Skull as in the Rusine group, but relatively narrower. ‘Tail short; neck maned. The Swamp Deer is somewhat smaller than the Sambur, and of a full yellowish colour. Fossil representatives of this group occur in the Pliocene of India. The Elaphurine group is represented only by the very aberrant C. davidianus of Northern China. In size and proportions this species approximates to the Swamp Deer, but the antlers are peculiar in rising straight from the brow and then giving off a long and straight back tine (correlated by Sir V. Brooke with the posterior branch of the Rusine antler); the summit of the beam is forked and in old individuals the two tines of the fork may again branch. Nasals long, and much expanded between the lachrymal vacuities, of which they form the inner border; lachrymal fossa large and deep. Tail long; neck maned. The .4aine group includes only the well-known Axis of India readily distinguished by the white spots with which the body is CERVIDE 321 marked. Antlers of a. Rusine type, the beam being much curved, and the brow tine usually given off at an acute or right angle. Molars very hypsodont. The coloration of the Axis is more brilliant than that of any other member of the family. Here may be noticed a group of Deer mainly characteristic of the eastern Palearctic region, frequently known as the Pseudaxine group, which appears to connect the Axine with the Hlaphine type. Well-known representatives of this group are C. sika (Fig. 128) of Japan, C. mantchwricus of China, and C. taévanus of Formosa. Fic. 128,—The Japanese Deer (Cervus sika). From Lord Powerscourt, Proc. Zool. Soc. 1884, p. 209. The antlers have a brow and tres tine, and then a forked beam, of which the posterior tine is the smaller. The lachrymal vacuity and fossa are of moderate size; and the auditory bulla is only moderately inflated, and quite smooth externally. Tail moderate ; neck maned. In summer the coat is spotted, but is plain in winter. A herd of C.. sika have been acclimatised in Ireland by Viscount Powerscourt, at Powerscourt, County Wicklow. A number of Deer from the Pliocene of Europe, such as C. perrieri and C. etueriarum, appear to be allied both to the Pseudaxine and Axine groups. The Llaphine or typical group is at once characterised by the presence of a bez tine to the antlers (Fig. 129), in which the beam is rounded, and splits up near the summit into a larger or smaller 21 322 UNGULATA number of snags, often arranged in a cup-like manner. Skull as in the preceding group. All the species large. The Red Deer, C. elaphus, which is dark brown in colour, with a light patch on the rump, inhabits Europe, Western Asia, and Northern Africa—the so-called Barbary Deer not being specifically distinct. A full-grown Scotch Stag is fully 4 feet in height at the withers. The antlers are shed between the end of February and the early part of April; old animals shedding earlier than younger ones. The young, which (as in all the members of the genus except some of the Rusine species) are spotted, are born at the end of May or the beginning Fic. 129.—Head of the Wapiti (Cervus canadensis). of June. The points on the antlers increase in number with the age of the creature, and when twelve are present it is known in Scotland as a “royal stag.” This number, however, is sometimes exceeded, as in the case of a pair of antlers, weighing 74 Ibs., from a stag killed in Transylvania, which had forty-five points. The antlers during the second year consist of a simple unbranched stem, to which a tine or branch is added in each succeeding year, until the normal development is attained, after which their growth is somewhat irregular. Many of the antlers dug up in British peat- beds (as Fig. 118) are larger than those of living individuals, and in the cave-deposits of England and the Continent antlers are met with rivalling those of the Wapiti in size; these large fossil antlers CERVIDA 323 probably indicating the ancestral form from which the Red Deer and several of the allied species are descended. The North American Wapiti (Cervus canadensis, Fig. 129), the Persian Maral (C. maral), the Kashmir Stag (C. cashmeerianus), as well as C. affinis of Tibet, are all closely allied to the Red Deer, but are of larger size, this being especially the case with the first two. A fine example of the antlers of the Wapiti is shown in the accompanying woodcut, and exhibits the absence of a cup at the surroyals, by which this species is distinguished from the Red Deer. The last, or Damine group of existing Deer includes the Common and the Persian Fallow Deer. These are readily characterised by the palmation of the antlers in the region of the surroyals and the spotted coat. The Common Fallow Deer (C. dama) stands about three feet in height. The Persian Fallow Deer ((. mesopotamicus) is very closely allied, differing only in its slightly larger size and the form of the antlers, the two breeding together. The common species, although now kept in English parks, does not appear to be a native of this country, having probably been introduced from the regions bordering the Mediterranean. The fur is of a yellowish-brown colour (whence the name “ fallow ”), marked with white spots ; there is, however, a uniformly dark brown variety found in Britain. The bucks and does live apart, except during the pairing season; and the doe produces one or two, and sometimes three fawns at a birth. The Fallow Deer from the Pleistocene and Pliocene deposits of the East Coast described under the names of C. brownt and C. faulconerit appear to have been closely allied to the existing species. The remarkable C. verticornis, of the Norfolk Forest-bed, is regarded as an aberrant member of this group, in which the antlers are very short and thick, with the brow tine cylindrical and downwardly curved, and the beam expanded above the tres tine into a crown with two points. The extinct Irish Deer (Cervus giganteus), of which the skeleton is shown in the woodcut (Fig. 130), is the only representative of the Megacerotine group. The antlers, which may have a span of over 11 feet, are enormously palmated, and have a bifurcated brow tine, a small bez tine, and a third posterior tine. The skeleton measures upwards of 6 feet at the withers. Remains of this species are especially common in the peat-bogs of Ireland, but are also met with in Pleistocene deposits over a large part of Europe. In addition to the forms already mentioned there are many other fossil species of Cervus, some of which, like the English Pleistocene C. sedgewicki, cannot be included in any of the existing groups. There is no conclusive evidence of the existence of any species of Cervus before the Lower Pliocene period. Telemetacarpalia.—This section includes all the Deer of the New World, together with some Old World forms, and is charac- 324 UNGULATA terised by retaining the distal extremities of the lateral (second and fifth) metacarpals. With the exception of Alces, Capreolus, and Hydropotes (which are either partly or entirely Old World types), the vomer is so much ossified as to divide the posterior bony nares into two distinct orifices (Fig. 132). RFD wees yy Zz sa i) ot Pa "Fra. 130.—Skeleton of the Gigantic Irish Deer (Cervus giganteus). After Owen. Rangifer1—The Reindeer, or Caribou as it is termed in North America, is the sole representative of the genus Rangifer, which is sufficiently distinguished from all its allies by the presence of antlers in both sexes. The lachrymal vacuity is small. This animal is distributed over the northern parts of Europe, Asia, and America; the differences which may be observable in specimens ‘from different regions not being sufficient to allow of specific distinction. The Reindeer is a heavily built animal, with short limbs, in which ? Hamilton-Smith, in Grifith’s Animal Kingdom, vol. v. p. 804 (1827). CERVIDE 325 the lateral hoofs are well developed, and the cleft between the two main hoofs is very deep, so that these hoofs spread out as the animal traverses the snow-clad regions in which it dwells. The antlers (Fig. 131) are of very large relative size. There is a bez as well as a brow tine, which are peculiar in being either 4 branched or #f palmated. In the American \ race (Caribou), as well as in some of the specimens found fossil in the English Pleistocene (Fig. 131), one of the brow tines is gener- ally aborted to allow of the great develop- Fia. 131.—Skull and antlers of the Reindeer (Rangifer tarandus), ment of the from an English Pleistocene deposit. br, Brow tine; bz, bez tine. other. The (After Owen.) dentition of the Reindeer is frequently remarkable for the very small size of the posterior lobe of the last lower molar. Vertebre : C7,D14,L5, 85, C11. The Reindeer has long been domesticated in Scandinavia, and is of especial value to the Laplanders, whom it serves as a substitute for the Horse, Cow, Sheep, and Goat. It is capable of drawing a weight of 300 lbs., and its fleetness and endurance are remarkable. Harnessed to a sledge it will travel without difficulty 100 miles a day over the frozen snow, on which its broad and deeply cleft hoofs are admirably adapted for travelling. During the summer the Lapland Reindeer feeds chiefly on the young shoots of the willow and birch; and since at this season migration to the coast seems necessary to the well-being of this animal, the Laplander, with his herds, sojourns for several months in the neighbourhood of the sea. In winter its food consists chiefly of the so-called reindeer-moss and other lichens which the animal makes use of its hoofs in seeking 326 UNGULATA for beneath the snow. The wild Reindeer grows to a much greater size than the tame breed; but in Northern Europe the former are being gradually reduced through the natives entrapping and domesticating them. The tame breed found in Northern Asia is much larger than the Lapland form, and is there used to ride on. Remains referable to the existing species are found in the cavern and other Pleistocene deposits of Europe. A lees.i—The Elk or Moose (Alces machlis) has the same general distribution as the Reindeer, and is like- wise the single existing representative of its genus. Itis the largest existing member of the Frio, 132.—Hinder part of the base of the cranium of the family, attaining some- Virginian Deer (Cariacus virginianus). From Garrod, Proc. times a height of 8 feet ACG SOD ATs BTS, at the withers. The antlers (Fig. 133) have neither brow nor bez tine, but form an enormous basin-shaped palmation, primarily composed of an anterior and a posterior branch; their weight may be as much as 60 lbs. The nasal bones are very short, and the narial aperture of great size. The Elk is covered with a thick coarse fur of a brownish colour, longest on the neck and throat. Its legs are long and its neck short, and as it is thus unable to feed close to the ground, it browses on the tops of low plants, the leaves of trees, and the tender shoots of the willow and birch. Its antlers attain their full length by the fifth year, but in after years they increase in breadth and in the number of snags, until fourteen of these are produced. Although spending a large part of their lives in forests, Elks do not suffer much inconvenience from the ereat expanse of their antlers, as in making their way among ‘trees they are carried horizontally to prevent entanglement with the branches. Their usual pace is a shambling trot, but when frightened they break into a gallop. The natural timidity of the Elk forsakes the male at the rutting season, and he will then attack whatever animal comes in his way. The antlers and hoofs are his 1 Hamilton-Smith, in Griffith's dnimal Kingdom, vol. v. p. 303 (1827). CERVIDA 327 principal weapons, and with a single blow from the latter he has been known to kill a wolf. The female often gives birth to two fawns, and with these she retires into the deepest recesses of the forest, the young remaining with her till their third year. The Elk ranges, but in scanty numbers, over the whole of Northern Europe and Asia, as far south as East Prussia, the Caucasus, and North China, and over North America from the New England States westward to British Columbia. Fossil species are found in the Pleistocene deposits of Europe. Cervalces..—A remarkable extinct Deer from the Pleistocene of North America, described as Cervalees, appears in some respects Fic. 133.—Head of Elk (Alces machlis). (although a true Telemetacarpalian) to connect Alces with Cervus. Thus the palmated antlers are divided into anterior and posterior branches, but below this division there are two tines apparently corresponding to the bez and posterior tines of Cervus giganteus (Fig. 130). Capreolus.2— Antlers (in the existing species) less than twice the length of the head, usually with three tines on each. Brow tine developed from the anterior surface of the upper half of the antler, and directed upwards. Lachrymal vacuity small. Premaxille not always articulating with nasals. Auditory bulle slightly inflated, rugose externally. Vertebrae: C7,D13,L6,86,C8. Tail very short. Glands in fore feet rudimentary ; large in hind feet. The Roe, or Roe Deer (Capreolus caprea), is a small form dis- 1 Scott, Proc. de. Nat. Sct. Philad, 1885, p. 181. 2 Hamilton-Smith, in Griffith's Animal Kingdom, vol. v. p. 818 (1827). 328 UNGULATA tributed over Europe and Western Asia, being one of the species found in the British Isles. The male is somewhat over two feet in height at the withers, of a dark reddish-brown colour in summer, with a white patch on the rump. The small antlers are approxi- mated at their bases, and consist of a rugged beam rising vertically for some distance, then bifurcating, and the posterior branch again dividing. The Roe dates from the Pleistocene period. Extinct Deer from the Continental Pliocene have been provisionally referred to Capreolus. Hydropotes.1—No antlers in either sex. Lachrymal fossa deep and short (Fig. 134); lachrymal vacuity of moderate size. Orbits Fic. 134.—The left lateral view of the skull of a male Chinese Water Deer (Hydropotes imermis), with the wall of the maxilla cut away to show the root of the canine. 3 natural size. (From Sir V. Brooke, Proc. Zool. Soc. 1872, p. 524.) small and but slightly prominent. Auditory bulla much inflated. Angle of mandible much produced backwardly (Fig. 134); alveolar margins of mandible in diastema sharp and everted. Canines of male very large, and slightly convergent. Vertebre: C 7,D 12, L6,84,C10. No tutts on metatarsals. Foot glands small in fore feet, deep in hind ones. The Chinese Water Deer (H. inermis) is the sole representative of this genus. In the absence of antlers and the large can- ines of the male it resem- bles Moschus, although very Fig. 135.—Upper surface of the brain of Hydropotes different in other respects. inermis. (From Garrod, Proc. Zool. Soc. 1877, p. 792.) Thus the brain (Fig. 135) has the hemispheres much convoluted, as in other Cervine, and approximates to that of Pudua ; 1 Swinhoe, Proc. Zool. Soc. 1870, p. 90. CERVIDE 329 while the placenta and viscera likewise agree with those of the true Deer. In the total absence of any ossification of the vomer to divide the posterior nares Hydropotes resembles Capreolus and differs from all the following genera. The Chinese Water-Deer is nearly of the same size as the Indian Muntjac. It has short legs and a long body, the hair covering the latter being of a light reddish- brown. It is a remarkably prolific animal, differing from all other Deer in producing five or six young at a time. The mandible of a ruminant from the Middle Miocene of Gers in France, described under the name of Platyprosopus, presents such a marked remblance to Hydropotes in the form of the angle as to suggest a more or less intimate affinity. Cariacus1—Skull (Fig. 132) with the vomer dividing the posterior nares into two distinct chambers; premaxillee not reach- ing nasals. Antlers never greatly exceeding the length of the head. Lachrymal vacuity very large, and lachrymal fossa small. Auditory bulle slightly inflated. Vertebre: C7,D13,L6,54,C13. Tail long or short. Colour uniform in adult. This genus, which agrees with the Reindeer in the division of the posterior nares by the ossified vomer, comprises a number of species confined to the New World, none of which attain very large dimensions, and the antlers of which are relatively smaller than in the existing species of Cervus. The genus may be divided into groups. The typical Cariacine group, as represented by C. virginianus, has well-developed antlers, with a short brow tine rising from the inner side of the beam, and directed upwards, and several branches ; a long tail; and no upper canines. In this species, as well as in C. mexicanus and other forms, the antlers do not divide dichotomously, and the lachrymal fossa is of moderate depth. The Mule Deer (C. macrotis) of North America is distinguished by the dichotomous branching of the antlers and the deeper lachrymal fossa. The Virginian Deer is somewhat smaller than the Fallow Deer, and of a uniform reddish-yellow colour in summer, and light gray in winter. The Blastocerine group of South America is represented by C. paludosus and C. campestris, and has dichotomous antlers, with no brow tine, and the posterior branch the larger, a short tail, and no upper canines. The Furciferine group includes C. chilensis and C. antisiensis, confined to western South America. The antlers are not longer than the head, with a large anterior tine curving forwards at right angles to the simple posterior one. Auditory bulle slightly inflated, and rugose. Upper canines may be present. The species are of medium size. C. clavatus, of Central America, while resem- bling this group in the characters of the skull and the arrangement 1 Gray, Proc. Zool. Soc. 1850, p. 237. 330 UNGULATA of the hair on the face, agrees with the next one in having simple spike-hke antlers. The South American Coassine group comprises the small forms known as Brockets, in which the antlers form simple spikes not exceeding half the length of the head. Some six species are known. Remains of Cariacus, mostly or entirely referable to existing species, are of common occurrence in the Brazilian cave-deposits. Blastomeryx, of the Pliocene of North America, is believed to be an allied type. Puduat—aAntlers in the form of minute simple spikes. Distinguished from the Coassine group of Cariacus by the articulation of the premaxille with the nasals (as in the Fwrciferine group), and the coalescence of the ectocuneiform with the naviculo-cuboid. as well as by various external characters. No upper canines. Re- presented only by the very small P. humilis of the Chilian Andes. Extinct Genera—In the European and other Tertiary deposits several genera of extinct (¢rvide occur, of which the more important may be briefly mentioned. -4mphitragulus, of the Lower Miocene of the Continent, has four lower premolars, brachydont molars, and no antlers; the largest species being somewhat bigger than the Musk-Deer. The closely allied Palcomeryx (Dremotherium or Micro- meryz) generally has but three lower premolars, and the brachydont upper molars (Fig. 122), like those of {mphitraaulus, want the small accessory inner column” found in modern Deer. In P. feianouzi, of the Lower Miocene, the lateral metacarpals, although slender, were complete, and the males had large canines, but no antlers. P. furcatus, of the Middle Miocene, had small antlers, and the canines appear to have been reduced in size. This genus, besides being repre- sented in the European Miocene, also occurs in the Pliocene of India and China; some of the species being as large as the Red Deer. Family GIRAFFID-£. In the existing genus the frontal appendages consist of a pair of short, erect, permanent bony processes placed over the union of the frontal and the parietal bones, ossified from distinct centres. though afterwards ankrlosed to the skull, covered externally with a hairy skin, present in both sexes, and even in the new-born animal. Anterior to these is a median protuberance on the frontal and contiguous parts of the nasal bones, which increases with age, and is sometimes spoken of as a third horn. Skull with a lachrymal vacuity. No upper canines. Molars brachydont, with rugose 1 Gray, Proce. Zool. Soe. 1850, p. 242. * This accessory column is shown in the figure of the molar of Bosclayhus on p. 311. GIRAF FIDE 331 enamel ; the upper ones having no inner accessory column. Lateral digits entirely absent on both fore and hind feet, even the hoofs not developed. Humerus with double bicipital groove. Vertebra : HEATH SE SS aint SS epyensenth RC SN aut Fic. 136.—The Giraffe (Giraffa camelopardalis). C7,D14,L5,83,C20. Gall-bladder generally absent. Male reproductive organs and placenta of a Bovine type. Dentition: i909 p Bm. Giraffa.1—The Giraffe (G. camelopardalis) is the sole existing representative of the genus, now confined to the Ethiopian region. 1 Zimmermann, Geograph. Geschichte, vol. ii. p. 125 (1780). 332 . UNGULATA In addition to the characters noticed above, the Giraffe is characterised by its great size and peculiar proportions; the neck and limbs being of great length, and the back inclining upwards from the loins to the withers. To produce the extremely elongated neck the seven cervical vertebra are proportionately long, which gives a somewhat stiff and awkward motion to the neck. The ears are large, the lips long and thin, the nostrils closable at the will of the animal, the tongue very long and extensile, and the tail of considerable length, with a large terminal tuft. An adult male may have a total height of 16 feet. The coloration consists of large blotches of darker or lighter chestnut- brown on a paler ground, the lower limbs and under parts being of a uniform pale colour. The Giraffe feeds almost exclusively on the foliage of trees, showing a preference for certain varieties of mimosa, and for the young shoots of the prickly acacia, for browsing on which its prehensile tongue and large free lips are specially adapted. It is gregarious in its habits, living in small herds of about twenty individuals, although Sir 8. Baker, who hunted it in Abyssinia, states that he has seen as many as a hundred together. Fossil species of Giraffa occur in Pliocene deposits over Greece, Persia, India, and China, thus affording one of many striking instances of the former wide distribution of the generic types now confined to the Ethiopian region. Allied Extinet Types.—The Pliocene deposits of many parts of the Old World yield remains of a number of large Ruminants which show such evident signs of affinity with the Giraffe that it is difficult to draw up a definition by which they can be separated in characters of family value from that genus. On the other hand, some of these forms approximate in the characters of the skull to some of the brachydont members of the Bovide, although it is quite clear from the nature of the cranial appendages that they cannot be included in that family. All these forms have brachydont molars, with rugose enamel, like those of the Giraffe ; while several of them have limb- bones approximating to those of the latter—the humerus, when known, having a double bicipital groove. The nature of the cranial appendages (when present) is not fully understood, but it appears that in some cases these approximated more to the type of an antler than to that of a horn ; although, from the absence of a “burr,” they appear never to have been shed. A gradual diminution in the length of the limbs and neck can be traced from the more Giraffoid to the more Bovoid forms of this extinct group; and it is manifest that if these animals be included in the Girafide the definition of that family as given above must be somewhat modified. Only brief mention can be made of the more important genera. The imperfectly known Vishnutheriwm, of the Pliocene of India and Burma, seems to make the nearest approach to the Giraffe, but ANTILOCAPRIDA 333 the limbs and cervical vertebrae were decidedly shorter, although of a similar slender type. Helladotherium, of the Pliocene of Greece and India, is represented by a species of considerably larger size than the Giraffe, with no appendages or lachrymal vacuity to the skull, and with shorter and stouter limbs and neck. Hydaspitherium, Bramatherium, and Sivatherium are Indian genera, characterised by the presence of large palmated and antler-like cranial appendages, varying considerably in arrangement. The former genus has a large lachrymal vacuity which is absent in the two latter. In the first and second genera all the appendages rise from a common base; but in Sivatherium there is a pair of simple horn-like projections on the orbits in addition to the posterior palmated antlers. Stvatheriwm was an animal of huge bulk, being the largest known representative of the Pecora. Another apparently allied type is Samotherium, of the Pliocene of the Isle of Samos, which appears also to have some affinity with the Antelopes. The skull is nearly as large as that of the Giraffe, and is of the same elongated shape, although depressed between the conical horn-cores, which rise vertically above the orbits, and without a median bony prominence on the frontals. The horn-cores form mere processes of the frontals. The diastema and the mandibular symphysis are shorter than in the Giraffe, and the latter is less deflected. The teeth, although larger, are almost indistinguishable from those of the Giraffe, the only well-marked difference being that the last lower premolar has a double in place of a single postero- internal column. Family ANTILOCAPRIDA. Closely allied to the Bovide, but the horns deciduous and branched. Antilocapra1—The Prongbuck, or Prong-horned Antelope (Antilocapra americana), as the single existing member of this family is called, is an animal of nearly the same size as the Fallow Deer, but of a lighter and more graceful build. It is an inhabitant of the prairies of North America, where it is one of the few representa- tives of the Cavicorn Pecora. The bony horn-cores are unbranched, and form vertical, blade-like projections immediately above the orbit. The horns themselves are compressed, and nearly one foot in length, having a gentle backward curvature, the short branch arising somewhat above the middle of its height, and inclining forwards. When the horn is about to be cast off it becomes loosened, and a new one is formed upon the bony core beneath it. The ears are long and pointed, and the tail is short. The neck has a thick mane of long chestnut-coloured hair, and there is a white patch on the rump. 1 Ord. Journ. de Physique, vol. Ixxxvii. p. 149 (1818). 334 UNGULATA Family BovIDa. Frontal appendages, when present, in the form of non-deciduous horns. Molars frequently hypsodont. Usually only one orifice to the lachrymal canal, situated inside the rim of the orbit. Lachrymal bone almost always articulating with the nasal. Canines absent in both sexes. The lateral toes may be completely absent, but more often they are represented by the hoofs alone, supported sometimes by a very rudimentary skeleton, consisting of mere irregular nodules of bone. Distal ends of the lateral metapodials never present. Gall-bladder almost always present. The number of cotyledons in the placenta generally varies from 60 to 100; whereas in the Cervide the number is usually from 5 to 12, Capreolus and Hydropotes having the fewest. In Giraffu the number is upwards of 180. The nature of the horns and horn-cores has been already explained ; in the majority of genera these appendages are present in both sexes, although much larger in the male (see p. 310). The Bovide, or hollow-horned Ruminants (Cavicornia), form a most extensive family, with members widely distributed through- out the Old World, with the exception of the Australian region ; but in America they are less numerous, and confined to the Arctic and northern temperate regions, no species being indigenous either to South or Central America. There is scarcely any natural and well-defined group in the whole class which presents greater difficulties of subdivision than this; consequently zoologists are as yet very little agreed as to the extent and boundaries of the genera into which it should be divided. For the present the genera provisionally adopted may be arranged under a number of sections or groups, which some writers regard as subfamilies. The series may be commenced with the Antelopes, the greater number of which are now characteristic of the Ethiopian region. Alcelaphine Section.—Includes large African Antelopes, of which the type genus ranges into Syria; generally characterised by their great height at the withers as compared with the rump. Skull with large frontal sinuses, extending into the horn-cores, and the horns lyre- shaped or recurved, and more or less approximated at the base. No large pits at apertures of supraorbital foramina in frontals; upper molars hypsodont and narrow. Horns in both sexes. General colour mostly uniform. Alcelaphus..lf Damalis be included, this genus is represented by some nine or ten living species. Head more or less long and narrow, with the muffle moderately broad and naked. Nostrils approximated, edged with stiff hairs. Horns compressed and ringed at the base, more or less lyrate, and bent back at the tips. Hoofs small. Tail of moderate length, and heavy. Two mamme. ? Blainville, Budd. Soc. Philom. 1816, p. 75. BOVID.E 335 In the typical forms, such as the Bubaline Antelope (4. buba- linus), the Harte-beest (f. cama, Fig. 137), and the Tora Antelope (4. tora, Fig. 138), the horns, which present the peculiar curvature shown in the figures, are situated on a crest at the vertex of the skull, and the facial portion of the cranium is greatly elongated. The Harte- beest, which is found throughout Central and Southern Africa, stands nearly 5 feet high at the withers, and is a somewhat ungainly looking animal, with short hair, which is grayish-brown above and newly white beneath. In the Pliocene of the Siwalik Hills in Northern India there occur remains of an -f{leeluphus (4. paleindicus) Fig, 137. —The Harte-beest (Aleelaphus cacmea). in which the skull had the long facial portion characteristic of the typical group, while the horns approximate to those of the Bontebok. The Blessbok (4. a/bifrons) and Bontebok (4. pygargus), belonging to the genus Jamalis of many authors, have the facial portion of the skull shorter, the horns situated more in advance of the plane of the occiput, and inclining regularly backwards. Of the Blessbok My. C. J. Anderson observes that “it is of a beautiful violet colour, and is found in company with black Wildebeests and Springboks in countless thousands on the vast. green plains of short crisp, sour grass occupying a central position in South Africa, Cattle and horses refuse to pasture on the grassy products of these plains, which afford sustenance to myriads of this Antelope, whose skin emits a most delicious and powerful perfume of flowers and sweet- 336 UNGULATA smelling herbs.” Since the time this was written these Antelopes have been greatly reduced in number. 4. (Damalis) hunteri, from East Africa, appears to be allied to A. senegalensis, but in the more elongated facial portion of the skull approximates to the Harte-beest, and thus confirms the view that Damalis should not form a distinct genus. Fic. 138.—Head of Alcelaphus tora. From Sclater, Proc. Zool. Soc. 1873, p. 762. Connochetes.1—Head short and massive, with the mufile very broad and bristly. Nostrils widely separated, hairy within. Horns on the vertex of the skull, immediately over the occiput, approxi- mated at base, cylindrical, bent outwards, and recurving upwards at the tip. Extremities of premaxille much expanded laterally, and firmly ankylosed. Vertebre: C7, D 14, L 6, $4, C 16. Hoofs very narrow. Tail very long, covered throughout with long hairs. Four mamme. Two species, C. taurina and C. gnu (Fig. 139), 1 Lichtenstein, Berlin Ges. Natuforsch. Freunde Magazin, vol. vi. pp. 152, 165 (1814). BOVID 337 hoth from South Africa. The former, or Brindled Gnu, is distin- guished by the absence of long hair on the face, the black (instead of white) tail, and the presence of dark vertical streaks on the shoulders ; it is never found to the south of the Orange River. The White-tailed Gnu stands about 4 feet 6 inches at the withers. These animals were formerly found in large herds, and are remarkable not only on account of their peculiar form, but also for their grotesque actions when alarmed. Some interesting observations have recently been published upon the mode of ae ze eA, 09d SS 3 aan a SN) Fia. 139.—The White-tailed Gnu (Connochutes gnu). development of the horns of the Gnu,! from which it appears that in very young individuals the horns are straight and divergent, situated some distance below the vertex of the head, and separated by a wide hairy interval. These young horns form the straight tips of those of the adult, the basal downwardly curved portion being subsequently developed. In the fully adult animal the hase of the horns forms a helmet-like mass on the forehead which completely obliterates the hairy frontal space of the young. Cephalophine Section. —Small or medium-sized African and Indian Antelopes, with simple horns present only in the males, a more or less elongated suborbital gland, a lachrymal depression in the skull, and square-crowned upper molars (Fig. 140). Lateral hoofs well developed. ' Ff, E. Blaauw, Proc. Zovl. Soc. 1889, p. 2. 22 338 UNGULATA Cephalophus.1—One pair of horns, arising far back on the frontals, conical, short, angulated at the base, and erect or recurved. Sub- orbital gland opening in the form of a slit, or as a row of pores. Auditory bulla divided by a distinct septum. Muffle large and moist. Tail very short. Head tufted. Upper molars of larger species with an accessory internal column. Dorsal vertebre fourteen in number. Some sixteen species, confined to southern and tropical Africa. The Duikerboks, as the members of this genus are called, are among the most graceful of the African Antelopes, the smallest species not being larger than a rabbit. The West African C. sylvicultor and C. longiceps are the largest species. Tetraceros.2-—Two pairs of conical horns, of which the anterior are much the smaller. Suborbital gland elongated, and lachrymal fossa very large. Upper molars (Fig. 140) without accessory internal column. One existing Indian species (TL. quadricornis). The Four-horned Antelope is found throughout the peninsula of India in jungle. The general colour is brown, lighter beneath and on the inside of the limbs. Remains of this species are found fossil in the cave-deposits of Madras, and a small Ruminant from the Pliocene of the Siwalik Hills has Fic. 140.—Palatal and outeraspects of D€€2 provisionally referred to this the three right upper premolars and first QeNus. inolar of the Four-horned Antelope (Tetra- Cervicaprine Section, Small or ceros quadricornis). From the Paleon- S : ‘alagiiae Indices large Antelopes now confined to the Ethiopian region, with horns present only in the males, lachrymal vacuity generally large, more or less distinct pits at the apertures of the supraorbital foramina in the frontals, and narrow upper molars in which there is no accessory internal column. Neotragus.*—Distinguished from the next genus by having the crown of the head tufted, muzzle hairy, premaxill long and reaching the lachrymals, nasals very short, mesethmoid much ossified, third lobe of last lower molar either absent or very small, and the hinder lobe of the corresponding upper molar much reduced. Three species, Salt’s Antelope (N. sultianus), from Abyssinia, and also N. kirki and N. damarensis ; the two latter having a small third lobe to the last molar. Writing of the first-named species, 1 Hamilton-Smith, in Grifith’s Animal Kingdom, vol. iv. p. 258 (1827). Taken to include Grimmia, Terphone, etc., of Gray. * Leach, Zrans. Linn. Soc. vol. xiv. p. 524 (1823). * Hamilton-Smith, in @rifith’s Animal Kingdom, vol. iv. p. 269 (1827). BOVIDE 339 Mr. W. T. Blanford! observes that “the Beni-Israel, or Om-dig-dig, one of the smallest Antelopes known, abounds on the shores of the Red Sea and throughout the tropical and subtropical regions of Abyssinia. It is occasionally, but rarely, found at higher eleva- tions ; I heard of instances of its being shot both at Serafie and Dildi, but it is not often seen above about 6000 feet. It inhabits bushes, keeping much to heavy jungle on the banks of water-courses, and is usually single, or in pairs, either a male and female or a female and young being found together; less often the female is accompanied by two young ones, which remain with her until full grown.” Nanotragus.2-—Horns small, parallel with frontals, and rising immediately above postorbital process of frontals, in front of the fronto-parietal suture. Lachrymal fossa very large, suddenly descending in front of the orbit, and extending on to the maxilla; lachrymal vacuity small. Auditory bulla large and smooth, without internal septum. Nasals of moderate length. Crown of the head smooth; naked part of muffle small; aperture of suborbital gland small. Lateral hoofs small or absent. Nine species.® The typical species is the Royal Antelope (V. pygmeus) of Guinea, the smallest existing representative of the Pecora. This species, together with N. moschatus and N. tragulus have no lateral hoofs, or tufts on the knees. In the Scopophorine group, comprising NV. scoparia, N. montanus, and N. hastatus, both these appendages are present; while in the Oreotragine group (N. melunotis and NV. oreotragus) the former are present and the latter absent. Pelea.t—Horns rather small, compressed, upright, scarcely diverging, and placed immediately over the orbits. No suborbital gland, nor lachrymal fossa; premaxille not reaching nasals. Tail short and bushy. Colour uniform. One species—the Rehbok (P. capreola), South Africa, is nearly of the size of a Fallow Deer, although more resembling a Chamois in build and habits. The colour is of a uniform light gray. This animal inhabits bare rocky districts, and thus differs widely from the Water-buck and its allies. Cobus.°—Large Antelopes, with the horns large, elongate, sub- lyrate, and ringed at the base, and with rudimentary suborbital glands. Skull with a deep frontal hollow, no lachrymal depression, 1 Geology and Zoology of Abyssinia, p. 268. 2 Sundevall, Kongl. Vetensk. Akad. Handi. for 1844, p. 191. Taken to include Calotragus, Scopophorus, Nesotragus, Pediotragus, and Oreotragus of Gray. 3 See V. Brooke, Proc, Zool. Soc. 1872, pp. 642 and 875. 4 Gray, Cat. Ungulate Mamm. Brit. Mus. p. 90 (1852). ®> Andrew Smith, Idlustrations of Zoology of South Africa, No. 12 (1840), “Kobus.” Is taken to include Adenota and Onotragus of Gray. 340 UNGULATA large lachrymal vacuity, and the premaxillz reaching the very long nasals. Tail long, with a ridge of hair above, and slightly tufted at the end. Colour uniform. Six species, African. The Antelopes of this genus are water-loving animals, the Water-buck (C. ellipsiprymnus) and the Singsing (C. defassus) being well-known examples. Both these species are much alike, standing as much as 4 feet 6 inches at the withers. The Water-buck of South and Eastern Africa is characterised by the coarseness of its long hair ; while in the Singsing of West and Central Africa the hair is remarkably fine and soft. Fossil Antelopes from the Pliocene of India are referred to Cobus. Helicophora, from the Lower Pliocene of Attica, is regarded as allied to Cobus, but it has no distinct supraorbital pits. Cervicapra.iman allied South African genus in which the tail is short and bushy and the premaxille do not reach the nasals. Three species. The Reitbok (C. arundinewm) is of a grizzly ochre colour ; it stands nearly 3 feet in height, and has horns about 1 foot in length. The Nagor (C. redunca) is about 6 inches shorter, with horns of half the length, and fulvous brown above and white below ; the West African C. bohor being rather larger. Antilopine Section. —A large group of moderate-sized or small Antelopes, most abundant in the deserts bordering the Palearctic, Oriental, and Ethiopian regions. Horns generally compressed and lyrate, or recurved, or cylindrical and _ spiral, ringed at base, sometimes present in both sexes. Skull with large pits at apertures of supraorbital foramina of frontals, and generally a distinct lachrymal fossa. Molars of upper jaw narrow, without inner accessory column, and resembling those of the Sheep and Goats. Tail moderate, compressed, hairy above. Antilope.-—Horns, present only in the male, long, cylindrical, subspiral, and diverging. Suborbital gland large, with a somewhat linear opening ; lachrymal depression of skull very large, and a small lachrymal fissure. Glands in the feet ; lateral hoofs present. One species, India. The well-known Black-buck (A. cervicapra) is found on open plains all over India, except in lower Bengal and Malabar. Old males are deep blackish-brown in colour on the back and sides and the outer surfaces of the limbs, the under parts and inner surfaces of the limbs white, and the back of the head, nape, and neck yellowish. Young males and females are fawn-coloured above. Very large herds are seen in the plains about Dehli and Mattra, which are said in some instances to reach to thousands. Horn- cores are found in the Pleistocene deposits of the valley of the * De Blainville, Bull. Soc. Philom. 1816, p. 75. Syn. Eleotragus. * Pallas, Spicilegia Zoologica, vol. i. p. 3 (1767). BOVIDA 341 Jumna which cannot be distinguished from those of the existing species. Aipyceros..— Horns compressed, lyrate, and wide-spreading ; present only in male. No suborbital gland, or lachrymal depression in the skull., No lateral hoofs. Two species; one from South and the other from West Africa. The Palla (4. melampus) is a large Antelope standing over 3 feet high at the withers, and readily distinguished by its dark red colour, gradually shading to white below. It is usually found on or near hills in herds of from twenty to thirty. 4. petersi is from the Congo. Saiga.2—Nose very large, convex, and inflated. Supraorbital gland present. Lachrymal fossa of skull small, and fissure absent ; narial aperture very large; nasals extremely short; supraorbital pits rather small. Horns yellow, lyrate, of moderate length ; present only in male. Vertebre: C7, D13,L6,84,C10. One species, Eastern Europe and Western Asia. The Saiga (S. tartarica) is a clumsily built and somewhat sheep-like Antelope inhabiting the steppes; it occurs fossil in the Pleistocene of France and England. Pantholops.’—Allied in the characters of the head and skull to Saiga, but the nose less convex, the nostrils of the male more swollen, and the horns of that sex black, very long, compressed, and lyrate ; those of female very. short. One species, Central Asia. The Chiru (P. hodgsoni) inhabits the highlands of Western Tibet and Turkestan. In the former area it generally goes in small herds of from three to six, and in the summer may be found grazing in early morning on the level spaces frequently found in the river valleys at elevations of about 15,000 feet. It is excessively shy and difficult to approach. The large size of the narial aperture in the skull of Chiru is suggestive of a connection with respiration at a high altitude, but this appears to be negatived by the occurrence of the same feature in the Saiga. Gazella.-—Delicately built and sandy-coloured Antelopes, with lyrate or recurved horns, which may be absent in the female, and are always smaller and simpler in that sex than in the male. Skull with moderate lachrymal fossa, and a distinct lachrymal fissure. Vertebre: C7, D13,L6,84,C14. Suborbital gland frequently small, and covered with hair. Face with a white streak running from the outer side of the base of each horn nearly down to the upper end of each nostril, cutting off a dark triangular central 1 Sundevall, Kongl. Vetensk. Akad. Handl, for 1845, p. 271. 2 Gray,’ List Mamm, Brit. Mus. p. 160 (1843). 3 Hodgson, Proc. Zool. Soc. 1834, p. 81. + De Blainville, Buli. Soc. Philom. 1816, p. 75. Is taken to include Procapra and J'ragops. 342 UNGULATA patch, and bordered externally by a diffused dark line (see Fig. 121, p. 310). The Gazelles, of which there are some twenty- four existing species, are typically Palearctic desert forms, the Springbok (G. euchore) being an outlying South African species. G. picticaudata and G. gutturosa are respectively found in Western Tibet and Mongolia, the former at great elevations. The majority of the Gazelles do not exceed 30 inches in height, although G. mohr is 36. Sir Victor Brooke classifies! the Gazelles as follows :— - A. No stripe on back; three lower premolars. a. White of rump not encroaching on the fawn of the haunches, I. Female with horns. 1. Horns lyrate or sublyrate——G. dorcas, G. isabella, » G. rufifrons, G. levipes, G. tilonura, G. naso. 2. Horns non-lyrate—G. cuvieri, G. leptoceros, G. spekei, G. arabica, G. bennetti, G. fuscifrons, G. muscatensis. II. Female without horns. G. subgutturosa, G. gutturosa, G. picticaudata. b. White of rump projecting forwards in an angle into the fawn colour of the haunches. Horns in both sexes. G. dama, G. mohr, G. soemmerring?, G. granti (Fig. 121), G. thomsont, B. A white stripe down the back, two lower premolars. Horns in both sexes.—G. euchore. The East African G. walleri is an aberrant species, in which the females are hornless, which has been made the type of the genus Lithocranius. It is characterised by the extreme density of the horns and skull, the slenderness of the mandible, and the small size of the cheek-teeth, the upper molars being relatively broader and lower than usual. The cranium is remarkable for the short- ness of its facial portion, the large size and production backwards of the supraoccipital, and for the circumstance that the long basicranial axis is nearly parallel with the plane of the palate. Fossil species of Gazella are found in the Pliocene and Pleistocene deposits of Europe and India. G. deperditu (brevicornis), of the Lower Pliocene of France and Greece, appears to be a generalised species in which the lower molars frequently have accessory columns, traces of which are found in some of the existing forms. Hippotragine Section.—Includes very large African Antelopes, with long horns, present in both sexes, which are placed over or behind the orbit, and are either recurved, straight, or subspiral. Skull with no distinct pits at apertures of supraorbital foramina in frontals, no lachrymal fossa, and only a small lachrymal fissure. No suborbital gland. Tail long, cylindrical, and tufted at the end. * Proc. Zool. Soc. 1873, p. 537. Three species subsequently described are here added to the list. BOVIDA ; 343 Upper molars extremely hypsodont, very broad, and with large accessory columns, thus closely resembling those of the Oxen. Some authorities divide this section into two. In the Pliocene it occurs in India and Europe. Hippotragus..Horns stout, rising vertically from a crest over the orbit at an obtuse angle to the plane of the nasals, then recurved ; lachrymal fissure in some instances almost obliterated. Neck with an erect recurved mane. Tail very distinctly tufted. Four species, tropical Africa and south to the Cape. The Sable Antelope (H. niger) is one of the best-known examples of this genus, occurring in South and East Africa. It stands upwards of 44 feet in height at the withers, and, except for some white streaks on the face and the whole of the under surface of the body, is of a black colour. The, Blaubok (ZZ. leuco- plueus) is distinguished by the glaucous hue of the hair. The other species are the Equine Antelope (4. equinus) and Baker’s Antelope (H. bukeri) from the Sudan, both closely allied, but the latter distinguished by its pale fulvous colour, pencilled ears, and black stripes on the shoulder. Skulls of fossil Antelopes from the Pliocene of India have been referred to Hippotragus (H. sivalensis), and Sir V. Brooke suggests that the European Pliocene Antilope recticornis is not generically separable. Oryx.°—Horns long, slender, nearly straight or somewhat recurved, rising behind the orbit, and inclining backwards in the plane of the nasals; lachrymal fossa distinct. Nape maned; tail long, and more haired than in Hippotragus. Four species, ranging over allthe African deserts to Arabia and Syria. The Gemsbok (0. gazella, Fig. 141), is a South African species characterised by its straight horns, the presence of a tuft of hair on the throat, as well as by the large patches and stripes of black on the head, back, limbs, and flanks. It stands nearly 4 feet in height at the shoulder, and the horns are 2 feet 9 inches in length. The colow of the upper part of the body is a rusty gray, and of the under part white, while these are separ- ated from each other by a well-defined black band on either side. These bands unite on the breast, and are continued as a single black band until reaching the lower jaw, where they again divide and form two transverse bands on the head, terminating at the base of the horns. The head otherwise is white, as also are the limbs, with the exception of the thighs, which are black. The Gemsbok generally goes in pairs, or in small herds of three or four. The Beisa (0. beise) of Abyssinia is distinguished by the absence of the tuft of hair on the throat. Writing of this 1 Sundevall, Kongl. Vetensk. Akad. Handl. for 1844, p. 196. 2 De Blainville, Bud/, Soe. Philom, 1816, p. 75. 344 UNGULATA species in his Geology and Zoology of Abyssinia, Mr. W. T. Blanford observes that “the appearance of a herd of Oryx is very imposing. They are some of the most elegant and symmetrical of animals, the motions being those of a wild Horse rather than of an Antelope. Their favourite pace appears to be either a steady quick walk or a trot; they rarely break into a gallop unless greatly alarmed. When frightened they dash off, sometimes snorting and putting Fic. 141.—The Gemsbok (Oryx gazella). their heads down as if charging, raising their long tails, and look- ing very formidable. They are wary animals, though far less so than some other Antelopes. It is said that they frequently attack when wounded, and their long straight horns are most deadly weapons.” The Arabian Beatrix Antelope (0. beatriz) is a much smaller animal, with the black markings confined to the head, fore limbs, and flanks, Finally, the Leucoryx (0. lewcoryx) of North Africa, while agreeing in size with the Beatrix, differs hy its curved horns and uniform coloration. ; The extinct Puleorys, of the Lower Pliocene of Europe and the Isle of Samos, appears to have been an ancestral form of Oryx, said to show some signs of affinity with Hippotrayus. , BOVIDE 345 Addaz.4—Horns with the same inclination as in Oryx, but with a slight spiral twist. No mane on nape, but a slight one on the throat. Hoofs rounded. One species (4. nusumaculatus), from North Africa and Arabia, the colour of which is nearly white. Tragelaphine Section.—Includes large, so-called Bovine, Ante- lopes now mainly characteristic of the Ethiopian region, but with one Oriental genus. Horns usually present in the male only (if developed in the female smaller), with a more or less distinct ridge in front, and usually twisted spirally, the front ridge twisting outwards from the base of the horn. Skull without lachrymal fossa, but with a large or small lachrymal fissure ; usually large pits at the apertures of the supraorbital foramina on the frontals ; premaxille reaching nasals. Muffle large and moist; nostrils approximated. Molars hypsodont or brachydont. Vertical white stripes frequently present on the body. a. LHind limbs much shorter than the fore. Horns behind the orbit, short, conical, faintly angulated. Nose bovine. Body without vertical stripes. Molars (Fig. 123, p. 311) hypsodont, with a large accessory column im those of the upper jaw. One Oriental genus. Boselaphus.2—The one genus of this subsection is represented only by the well-known Nilghai (B. tragocamelus) of India. The male stands over 4 feet in height at the shoulder, with horns about 8 inches in length; the hornless female being about one third smaller. Both sexes have a short erect mane, and the male has also a tuft of hair upon the throat. When adult the sexes are very different in colour, the male being of a dark iron gray or slate colour, approaching black on the head and legs, while the female and young are of a bright light brown or fawn colour. In both male and female at all ages the lips, chin, and under parts, as well as two transverse stripes on the iriner sides of the ears and rings on the fetlocks, are white, and the mane and tip of the tail black. The Nilghai is one of the few Antelopes occurring in India, where it is found from near the foot of the Himalaya to the south of Mysore, though rare to the north of the Ganges and also in the extreme south. It is most abundant in Central India, and does not occur in Assam or the countries to the east of the Bay of Bengal. It frequents forests and low jungles, though often found in toler- ably open plains, associating in small herds. One, or very often two, young produced at a birth. Fossil remains of species of this genus occur in the Pleistocene and Pliocene deposits of India. b. Fore and hind limbs equal. Horns long, and spirally twisted. Nose cervine, and aperture of suborbital gland very small. 1 Rafinesque, nal. Nat. 1815, p. 56. 2 De Blainville, Bull, Soc. Philom. 1816, p. 75. Syn. Portax, Hamilton- Smith. 346 UNGULATA Body generally striped. Molars brachydont, those of the upper jaw in existing forms with a small inner accessory column. Three existing Ethioman genera. Tragelaphus..Female hornless. Horns of males (Fig. 142) over il ponrea ns Fic. 142.—Head of Tragelaphus gratus. Fyrom Sclater, Proc. Zool. Soc. 1883, p. 36. orbit, with one or two spiral turns, obscurely ridged, the posterior ridge being more developed than the anterior. Skull with small supraorbital pits, very small lachrymal fissure, and no deep inter- cornual depression in the frontals. Neck maned or smooth. Hoofs short or long. Coloration usually brilliant, differing markedly in the two sexes, and the white bands on the body, when present numerous and distinct. Seven species. : 1 De Blainville, Bull. Soc. Philom. 1816, p. 75. Includes Euryceros, Gray. BOVIDA 347 The Harnessed Antelopes are among the handsomest of the whole group. The small Guib (Tf. scriptus) is not larger than a Goat, but 7. angusi is 3 feet 4 inches in height at the shoulder. In T. scriptus, T. angasi, and T. euryceros, the two sexes differ in colour, the body is marked by white stripes descending from a white dorsal streak, and the hoofs are short ; the third species differing from the others by the absence of a mane on the neck, back, and belly. 7. gratus agrees with this group in coloration (the mane being Fic. 143.—The Kudu (Strepsiceros kudu). From Sclater, List of Animals in Zoological Society's Gardens, 1883, p. 136. absent), but differs in the extreme elongation of its hoofs. The Nakong, 7. spekei, while having the long hoofs of J. gratus, has a perfectly plain body coloration, with a mane on the neck. The two species with elongated hoofs inhabit swampy districts, for which this peculiar structure is admirably adapted ; and the Nakong, when frightened, will rush into the water and leave only its nostrils and the tips of the horns above the surface. The small Bushbuck (ZL. sylvaticus) of South Africa has no stripes, and short hoofs. Strepsiceros.'—Females hornless. Horns (Fig. 143) more twisted than in Yragelaphus, forming an open spiral, with the anterior ridge 1 Gray, List. Mamm. Brit. Mus. p. 155 (1843). 348 UNGULATA very strongly developed, and rising at an obtuse angle to the plane of the nasals. Skull with large supraorbital pits, large lachrymal fissure, and deep intercornual depression. Hoofs short. Body with white vertical stripes descending from a longitudinal dorsal streak. Two existing species. . The Kudu (S. kudu, Fig. 143) extends from South Africa to Abyssinia, and is only inferior in size to the Eland. The horns are about 4 feet in length, and form a very open spiral, and there is a fringe of long hair down the front of the neck. The Lesser Kudu (S. imberbis), of Somaliland is a much smaller form, without the fringe of hair on the neck, and with a much smaller axis formed by the spiral of the horns. An imperfect skull from the Pliocene of Northern India has been referred to Strepsiceros. Oreas.1\—Females horned. Horns twisted on their own axis, with very strong ridges, inclining upwards and outwards in the plane of the nasals. General characters of skull as in preceding genus. Stripes on body, if present, very faintly marked. One existing species. The Eland (0. canna) is the largest of all the Antelopes, the males standing nearly 6 feet at the withers. One variety from South Africa is of a uniform pale fawn colour, while the Central African form is of a bright tan colour, marked by a number of thin pale vertical stripes descending from a dark dorsal ridge—these . markings fading more or less in the adults. The males have a large dewlap, a tuft of brown hair on the forehead, and a small mane on the neck. The straight black horns of the male are usually about 18 inches long. Elands were formerly extremely abundant in Southern and Eastern Africa, but their destruction has been so relentless that they have totally disappeared from extensive areas, and are daily becoming scarcer, Portions of upper jaws from the Pliocene deposits of India appear to indicate the former existence in that area of large Antelopes closely allied to the Eland, but distinguished from the living species by the greater size of the inner accessory column in the upper molars. Allied Extinct Types.—Large Antelopes with spirally twisted horns appear to have been common over Southern Europe in Pliocene times, but their exact affinity is in many cases difficult to determine. Of these, Palworeas, which occurs in the Lower Pliocene of Europe and Algeria, appears to present affinities both to Orews and Strepsiceros, and may have been the ancestral type from which these two genera are derived ; the upper molars have well-developed accessory columns. The so-called .Lntilope torticornis, of the French Pliocene, * Desmarest, Mummatogie, p. 471 (1822). BOVIDA 349 resembles Tragelaphus in the greater development of the posterior as compared with the anterior ridge of the horn-cores, and has accordingly been referred to that genus. Protragelaphus, of the Lower Pliocene of Attica, differs from all the other types in the absence of the anterior ridge on the horn-cores and of the supraorbital pits, while it has a distinct lachrymal fossa. In this place it will be convenient to notice certain fossil forms which do not accord with any of the existing sections of the family, and for the reception of which the Paleotragine section has been formed. In these types the horn-cores are laterally compressed like those of the modern Goats, but the upper molars resemble those of the brachydont Antelopes. The earliest of these genera, and the first representative of the Antelopes yet known, is Protragoceros, of the Middle Miocene of France, first described as Antilope clavata ; Paleotragoceros and Tragoceros, of the Lower Pliocene, are distin- guished by their larger horns and wider molars. A remarkable large Antelope from the Lower Pliocene of the Isle of Samos, in the Turkish Archipelago, proposed to be described as Criotherium, appears to be unlike any other form. The horns, which are placed on the extreme vertex of the skull, are very short, tightly twisted, and project in front of the forehead. The upper molars have short and broad crowns, with no accessory column on the inner side. Rupicaprine Section—The Caprine Antelopes, as the typical members of this section may be termed, appear to connect the true Antelopes with the Goats. They are mostly small or medium- sized forms, inhabiting portions of the Palearctic and Oriental regions, with one outlying North American genus. The typical forms present the following features. Horns present, and of nearly equal size in both sexes, rising behind the orbits, short, ringed at the base, conical or somewhat compressed, and recurved. Sub- orbital gland generally present, in some cases small. Build clumsy ; hoofs large ; tail short, tapering, hairy above. Skull with lachrymal fossa, but no fissure. Molars as in the Caprine section. Rupicapra.' Horns short and cylindrical, rising perpendicularly from the forehead for some distance, then bending sharply back- wards and downwards, forming hooks with pointed tips. Premaxille not reaching the nasals. One species, Palearctic. The Gemse, or Alpine Chamois (£2. tragus), inhabits the high mountains of Europe from the Pyrenees to the Caucasus. It stands about 2 feet in height at the withers. The body is covered in winter with long hair of a chestnut-brown colour, that of the head being paler, with a dark brown streak on each side. At other seasons the colour is somewhat lighter, in spring approaching to gray. Underneath the external covering the body is further 1 De Blainville, Bull. Soc. Philom. 1816, p. 75. 350 UNGULATA protected from cold by a coat of short thick wool of a grayish colour. The tail is black ; the ears are pointed and erect ; the hoofs have the outer edges higher than the soles, and are thus admirably adapted for laying hold of the slightest projection or roughness on the face of the rocky precipices it frequents. The Chamois is gregarious, living in herds of fifteen or twenty, and feeding generally in the morning or evening. The old males, however, live alone, except in the rutting season, which occurs in October, when they join the herds, driving off the young males, and engaging in contests with Fic. 144.—Nemorhewdus crispus, From Sclater, List of Animals in Zoological Society's Gardens, 1883, p. 151. each other that often end fatally. The period of gestation is twenty weeks, when the female, beneath the shelter of a projecting rock, produces one and sometimes two young. In summer the Chamois ascends to the limits of perpetual snow, being only out- stripped in the loftiness of its haunts by the Ibex; and during that season it shows its intolerance of heat by choosing such browsing grounds as have a northern exposure. Nemorhedus..—Horns rounded, gradually recurving, without distinct hook at the end. Suborbital gland small or wanting ; ears large; skull with a large lachrymal depression, and the premaxille not quite reaching the nasals. Some nine species, ranging from the Eastern Himalayas to North China and J. apan, and southwards 1 Hamilton-Smith, in Grifith’s Anima? Kingdom, vol. v. p. 352 (1827), BOVIDAE 351 to Formosa, the Malay Peninsula, and Sumatra. The smallest species is the Himalayan Goral (.V. geru/). Of the larger forms we may mention the Himalayan Serow (V. bubalinus) the Cambing- Utan (V. sumatrensis) of Sumatra, and the Japanese V. crispus (Fig. 144). Of the Serow, Colonel Kinloch remarks that “it is a large and powerful beast. The body is covered with very coarse hair, which assumes the form of a bristly mane on the head and shoulders, and gives the beast a ferocious appearance, which does not belie its disposition. The colour is a dull black on the back, bright red on the sides, and white underneath, the legs also being dirty white. The ears are very large, the muzzle is coarse. The Serow has an awkward gait, but in spite of this can go over the worst ground ; and it has perhaps no superior in going down steep hills. It is a solitary animal, and nowhere numerous.” Hupleceros.1—The Rocky-Mountain Goat (Huploceros montanus), inhabiting the northern parts of Califormia, appears to be very closely allied to Vemorhedus. The horns are somewhat compressed at the base; there is no suborbital gland; and the ears are small. The hair, which is whitish in colour, is very long, and especially abundant in the region of the throat, shoulders, flanks, and tail. The animal is about the size of a large Sheep. Budoreas?2—The Takin (B. taxicoler) of the Mishmi Hills in Assam, and an allied species from Eastern Tibet, are larger forms apparently related to Vemorhedus, but with a much greater develop- ment of the horns. The horns of what is considered to be the male® arise from the vertex of the skull, and are nearly in con- tact in the middle line; they first bend outwards and downwards, and then suddenly upwards and backwards. Those regarded by Mr. Hume as referable to the female are directed at first outwards, and then gradually curve upwards and backwards, without any down- ward flexure or angulation. The horns of the male may be 2 feet in length, with a basal diameter of 13 inches. The muzzle is hairy, with a small naked mufile. There appear to be considerable seasonal and sexual variations in colour; the body being in some cases of a yellow dun, while in others it is a dusky, reddish-brown, with much black intermingled. The heads of large males are blackish. Searcely anything is known of the habits of the Takin, which never appears to have been seen alive by Europeans. “‘aprine Section.—Both sexes with horns, but those of the female small. Horns usually compressed, triangular, with transverse ridges, and either curving backwards or spiral. Muzzle hairy, without naked muffle. Suborbital gland small or absent ; lachrymal 1 Hamilton-Smith, in Grijith’s Animal Kingdoin, vol. v. p. 354 (1827). Amended from ‘* Aplocerus.” 2 Hodgson, Journ. As. Soe. Benga’, vol. xix. p. 65 (1850). * See A. O. Hume, Proe. Zool. Soe, 1887, pp. 483-486. 352 UNGULATA fossa of skull present or absent. Tail short and flattened. Foot- glands frequently present. Molars very hypsodont; those of the upper jaw being narrow, without an accessory internal column. Mainly Palearctic, but with some outlying forms. This section includes the Goats and Sheep, which are so closely connected that it is difficult to give well-marked generic characters that will hold good for all the species. They seem to be one of Fic. 145.—The Alpine Ibex (Capra ibex). the latest developments of the Bovidw, since they are unknown before the Pliocene period ; and are essentially mountain forms. Capra.i—Horns flattened from side to side, and either curving backwards (Fig. 145) or spirally twisted. No suborhital gland and no lachrymal fossa in the skull. Foot-glands, if present, only in the fore feet. Chin more or less bearded. Males with a strong odour. Vertebree: C 7,D13,L6,8 4,C 9-13. Some dozen species, ranging over all the higher mountains of Southern Europe, from Spain to the Caucasus; also found in Abyssinia, Persia, Sind, and Baluchistan, thence through the higher Himalaya, and so on to Tibet and Northern China. One outlying species occurs in the Nilgherries of Southern India. ' Linn. Syst. Nut. 12th ed. vol. i. p. 94 (1766). BOVIDA 353 The European Ibex or Steinbok (Fig. 145), which may be taken as a typical Goat, stands about 24 feet in height at the shoulder. In summer the hair is short and smooth, and of an ashy-gray colour, but a long coat is developed in winter. The horns of the male rise in a bold backward sweep from the forehead, and are characterised by the strong transverse ridges on the broad and flat anterior surface. They are said to be not more than some 2 feet in length, but these dimensions are greatly exceeded by the horns of the Himalayan Ibex. The Alpine Ibex lives at a greater height than the Chamois, spending the day just at the limit of perpetual snow, and descending at night to graze at lower levels. Both this and the Himalayan species generally live in small herds of from five to fifteen or more; they are wary animals, although not so much so as many of the wild Sheep. The following list, mainly taken from two papers by Mr. Sclater,! gives the distribution of the various species of Goats, with some remarks on their peculiarities :-— (1) C. tbex, confined to the Alps of Switzerland, Savoy, and the Tyrol, and now nearly extinct, except where artificially pre- served. (2) C. sibiricu, closely allied to the preceding, but with larger horns, occurs in the Altai Mountains, and throughout the Himalaya from Kashmir to Nipal, and northward towards Turke- stan. (3) C. sinaitica, of the mountains of Upper Egypt, the Sinaitic Peninsula, and Palestine, is allied to the two preceding species, but has the horns somewhat more compressed, with a difference in the ridges on the front. (4) C. caucasica, a very distinct species, confined to the Caucasus, where it inhabits the western part of the Great Caucasus; with thick horns curving backwards and outwards in one plane, with the exception of their tips, which incline inwards.2 (5) C. pallasi is an allied species from the Eastern Caucasus, distinguished, among other features, by the curvature of the horns, which lie flatter and twist more outward from the forehead, with a greater terminal inward bend. (6) C. pyrenaica, of the Pyrenees, and the higher ranges of Central Spain, Andalusia, and Portugal, is another nearly related species. (7) C. wgagrus, formerly abundant over the Grecian Archipelago, but now restricted in Europe to Crete and some of the Cyclades, is found throughout the mountains of Asia Minor and Persia, and thence to Baluchistan and Sind. The horns are thinner and sharper in front than in the Ibexes, and this species is generally regarded as the ancestral stock of the various breeds of domestic Goats. (8) C. dorcas, a Goat from the island of Jura, near Eubcea, has been described under this name, and is apparently nearly allied 1 Proc. Zool, Soe, 1886, p. 314; and 1887, p. 552. ? Specimens referred by Dinnik to C. caucasica have been made the types of another species—C”. severtzovi. 23 354 UNGULATA to C. egagrus. (9) C. walie, an apparently well - characterised species from the highest ranges of Abyssinia. (10) C. falconeri ; the Markhoor differs from all the preceding species by the spiral twisting of its horns, which attain enormous dimensions. It occurs in the Pir-Panjal range south of Kashmir, and thence into Afghanistan and the Suleiman range, and northwards to Astor, Gilgit, and Scardo (Baltistan). The specimens from the Suleiman range have the spiral of the horns very close, somewhat as in the Eland ; while in those from Astor, Gilgit, and Scardo it is very open, as in the Kudu. The Pir-Panjal race occupies a somewhat inter- mediate position in this respect. (11) C. jemlaica, the Thar, inhabits suitable regions along the whole range of the Himalaya from Kashmir to Bhutan. Together with the next species, it differs from the more typical Goats in its short, thick, and much compressed horns, the anterior border of which is keeled, and the moist naked muffle. There are no glands in the fore feet. It was generically separated by Gray as Hemitragus. (12) C. hylocrius, the so-called Ibex of the Nilgherries, Anamallays, and other adjoin- ing ranges of Southern India, is an outlying species, apparently allied to the preceding, but with somewhat different horns, in which the external angle in front is much rounded off. Of fossil Goats we have but little knowledge. Remains of C. pyrenaica are found in cave-deposits at Gibraltar ; and it is not improbable that the genus is represented in the Upper Pliocene of France. Several species occur in the Pliocene of India, C. siralensis being apparently closely allied to C. jemlaica, while another has horns resembling those of C. falconeri, and it is possible that a third may be more nearly related to the Ibexes. Ovis.\—Horns curving backwards and downwards in a bold sweep, with the tips everted, generally with more or less prominent transverse ridges, and brownish in colour. Suborbital gland and lachrymal fossa usually present, but generally small. Foot-glands in all the feet. Chin not bearded ;2 males without a strong odour. Vertebre: C7, D 13, L 6,84, C1014. Some twelve species, mainly Palearctic, but extending into the adjacent portions of the Oriental region, and with one outlying species in North America. The more typical Sheep are closely connected with the Goats by the Himalayan Bharal (0. nahwra) and the Aoudad (0. tragelaphus) of Northern Africa, both these species having no suborbital gland and no lachrymal fossa, while their comparatively smooth and olive- coloured horns show a decided approximation to those of the Goats. Both present, however, the ovine character of glands in all the feet. In the typical Sheep the basioccipital of the skull is wider in front than behind, with the anterior pair of tubercles * Linn. Syst. Nat. 12th ed. vol. i. p. 97 (1766). ? There may be a beard on the throat, as in Q, cycloceros, BOVIDE 355 widely separated and much larger than the posterior pair. The Bharal, however, resembles the Goats in having an oblong basi- occipital, with the posterior tubercles larger and more prominent than the anterior ones, both being situated in the same antero- posterior line. These transitions towards the caprine type are, however, not sufficient to support the view that the Bharal should form the type of a distinct genus (Pseudois), more especially since some of the typical Sheep, like 0. canadensis, have the lachrymal fossa of the skull very much reduced in size. The distinction of the various permanent modifications under which wild Sheep occur is a matter of considerable difficulty. Trivial characters, such as size, slight variations in colour, and especially the form and curvature of the horns, are relied upon by different zoologists who have given attention to the subject in the discrimina- tion of species, but no complete accord has yet been established. The most generally recognised forms are enumerated below. The geographical distribution of wild Sheep is interesting. The immense mountain ranges of Central Asia, the Pamir and Thian- Shan of Turkestan, may be looked upon as the centre of their habitat. Here, at an elevation of 16,000 feet above the sea-level, is the home of the magnificent Ovis polt, named after the celebrated Venetian traveller Marco Polo, who met with it in his adventurous travels through this region in the thirteenth century. It is remark- able for the great size of the horns of the old rams and the wide open sweep of their curve, so that the points stand boldly out on each side, far away from the animal’s head, instead of curling round nearly in the same plane, as in most of the other species. A Sheep from the same region, in which the horns retain their more normal development, has received the name of 0. karelini, but, according to Mr. W. T. Blanford,! is not distinct specifically from 0. poli. East- ward and northward is found the Argali (0. argali), with a wide and not very well determined range; it formerly occurred in the Altai, but is now found in Northern Mongolia. Still farther north, in the Stanovoi Mountains and Kamschatka, is 0. nivicola, and away on the other side of Behring’s Strait, in the Rocky Mountains and adjacent highlands of western North America, is the “Bighorn” or Mountain Sheep (0. canadensis), the only member of the genus found in that continent, and indeed—except the Bison, Musk-Ox, Mountain Goat (Haploceros), and the Prong-buck (Antilocapra)— the only hollow-horned Ruminant, being like the rest obviously a straggler from the cradle of its race. The two last-named species are nearly allied, and are characterised by the slight development of the ridges on their horns and the very shallow lachrymal fossa. Turning southward from the point from which we started, and still a little to the east, in Nipal and Western Tibet, 1 Proe, Zool. Soc. 1884, p. 326. 356 UNGULATA is the Himalayan Argali (0. hodgsoni), having massive and strongly curved horns, with bold ridges, like those of the true Argali. Indeed, were it not for their isolated areas there would appear to be no grounds for distinguishing these two closely allied forms, and it is not improbable that they are really identical. 0. brookei appears to have been founded on a hybrid between 0. hodgsoni and O. vignet. In the same districts, and also in Southern Ladak, there occurs the Bharal (0. nahura), with smaller, smoother, and more spreading horns. Passing in a south-westerly direction we find a series of smaller forms, 0. vignei of Ladak, 0. cycloceros of Northern 4 ae Fic. 146.—The Moufilon (Ovis musimon). From a living animal in the London Zoological Gardens. India, Persia, and Baluchistan, 0. gmelini of Asia Minor and Persia O. ophion, confined to the elevated pine-clad Troodos Mountains of the island of Cyprus, and said at the time of the British occupa- tion in 1878 to have been reduced to a flock of about twenty-five individuals, and O. musimon, the Moufflon of Corsica and Sardinia (see Fig. 146), believed to have been formerly also a native of Spain. In the three latter species the females are hornless. Lastly. we have the somewhat aberrant, Goat-like Aoudad (0. tragelaphus), of the great mountain ranges of North Africa, in which, as already Stare | the te and horns resemble those of the Bharal although the tail is longer, and there is a thick fri tee on the throat, chest, and fore legs, neee one BOVID.E 357 We thus find that Sheep are essentially inhabitants of high mountainous parts of the world, for dwelling among which their wonderful powers of climbing and leaping give them special advantages. No species frequent by choice either level deserts, open plains, dense forests, or swamps. By far the greater number of species are inhabitants of the continent of Asia, one extending into North America, one into Southern Europe, and one into North Africa. No wild Sheep exist in any other part of the world, unless the so-called Musk-Ox of the Arctic regions, the nearest existing ally to the true Sheep, may be considered as one. Geo- logically speaking, Sheep appear to be very modern animals, or perhaps it would be safer to say that no remains that can be with certainty referred to the genus have been met with in the hitherto explored true Tertiary beds, which have yielded such abundant modifications of Antelopes and Deer. They are generally con- sidered not to be indigenous in the British Isles, but to have been introduced by man from the East in prehistoric times. A fossil Sheep (Oris savigni), apparently allied to the Argali, has, however, been deseribed from the so-called Forest-bed of the Norfolk coast. The Sheep was a domestic animal in Asia and Europe betore the dawn of history, though quite unknown as such in the New World until after the Spanish conquest. It has now been intro- duced by man into almost all parts of the world where settled agri- cultural operations are carried on, but flourishes especially in the temperate regions of both hemispheres. Whether our well-known and useful animal is derived from any one of the existing wild species, or from the crossing of several, or from some now extinct species, is quite a matter of conjecture. The variations of external characters seen in the different domestic breeds are very great. They are chiefly manifested in the form and number of the horns, which may be increased from the normal two to four or even eight, or may be altogether absent in the female alone, or in both sexes ; in the form and length of the ears, which often hang pendent by the side of the head: in the peculiar elevation or arching of the nasal bones in some Eastern races; in the length of the tail, and the development of great masses of fat at each side of its root, or in the tail itself; and in the colour and quality of the fleece. Ovibos1—This genus is generally considered to be a connecting link between the Caprine and Bovine sections, but should rather be regarded as an aberrant type of the former. Horns of adult male rounded, smooth, and closely approximated at their bases, where they are depressed and rugose ; curving downwards, and then upwards and forwards. Muzzle eaprine; no suborbital gland, no lachrymal fossa or fissure in skull ; orbits tubular: a large narial aperture and very short nasals; premaxille not reaching nasals. 1 De Blainville, Bull. See. PAP. 1816, p. 78. 358 UNGULATA Tail short, and molar teeth caprine. One existing and two fossil species, Palearctic and Nearctic. The animal commonly known as the Musk-Ox (Ovibos moschatus), though approaching in size the smaller varieties of Oxen, is in structure and habits closely allied to the Sheep, its affinities being well expressed by the generic name Ovibos bestowed upon it by De Blainville. The specific name, as also the common English appellatives “ Musk-Ox,” “ Musk-Buffalo,” or ‘“‘ Musk-Sheep,” applied to it by various authors, refer to the musky odour which the animal exhales. This does not appear to be due to the secretion of a special gland, as in the case of the Musk-Deer; but it must be Fic. 147.—The Musk-Ox (Ovibos moschatus). observed that, except as regards the osteology, very little is known of the anatomy of this species. It about equals in size the small Welsh and Scotch cattle. The head is large and broad. The horns in the old males have extremely broad bases, meeting in the median line, and covering the brow and whole crown of the head. They are directed at first downwards by the side of the face and then turn upwards and forwards, ending in the same plane as the eye. Their basal halves are of a dull white colour, oval in section and coarsely fibrous; their middle part smooth, shining, and round ; their tips black. In the females and young males the horns are smaller. and their bases are separated from each other by a space in the middle of the forehead. The ears are small, erect, and pointed, and nearly concealed in the hair. The space between the nostrils and the upper lip is covered with short close hair, as in Sheep and Goats without any trace of the bare muffle of the Oxen. The greater part BOVIDA 359 of the animal is covered with long brown hair, thick, matted, and curly on the shoulders, so as to give the appearance of a hump, but elsewhere straight and hanging down,—that of the sides, back, and haunches reaching as far as the middle of the legs and entirely concealing the very short tail. There is also a thick woolly under- fur, shed in the summer. The hair on the lower jaw, throat, and chest is long and straight, and hangs down like a beard or dewlap, though there is no loose fold of skin in this situation as in Oxen. The limbs are stout and short, terminating in unsymmetrical hoofs, the external one being rounded, the internal pointed, and the sole partially covered with hair. The Musk-Ox is at the present day confined to the most northern parts of North America, where it ranges over the rocky barren grounds between the 60th parallel and the shores of the Arctic Sea. Its southern range is gradually contracting, and it appears that it is no longer met with west of the Mackenzie River, though formerly abundant as far as Eschscholtz Bay. Northwards and eastwards it extends through the Parry Islands and Grinnell Land to North Greenland, reaching on the west coast as far south as Melville Bay; and it was also met with in abundance by the German polar expedition of 1869-70 at Sabine Island on the east coast. No trace of it has been found in Spitzbergen or Franz Joseph Land. As proved by the discovery of fossil remains, it ranged during the Pleistocene period over northern Siberia and the plains of Germany and France, its bones occurring very generally in river deposits along with those of the Reindeer, Mammoth, and Woolly Rhinoceros. It has also been found in Pleistocene gravels in several parts of England, as Maidenhead, Bromley, Freshfield near Bath, Barnwood near Gloucester, and also in the lower brick- earth of the Thames valley at Crayford, Kent. It is gregarious in habit, assembling in herds of twenty or thirty head, or, according to Hearne, sometimes eighty or a hundred, in which there are seldom more than two or three full-grown males. The Musk-Ox runs with considerable speed, notwithstanding the shortness of its legs. Major H. W. Feilden, naturalist to the Arctic expedition of 1875, says: “No person watching this animal in a state of nature could fail to see how essentially ovine are its actions. When alarmed they gather together like a flock of sheep herded by a collie dog, and the way in which they pack closely together and follow blindly the vacillating leadership of the old ram is unquestion- ably sheep-like. When thoroughly frightened they take to the hills, ascending precipitous slopes and scaling rocks with great agility.” They feed chiefly on grass, but also on moss, lichens, and tender shoots of the willow and pine. The female brings forth a single young one in the end of May or beginning of June after a gestation of nine months. According to Sir J. Richardson, “ when this animal 360 UNGULATA is fat its flesh is well tasted, and resembles that of the Caribou, but has a coarser grain. The flesh of the bulls is highly flavoured, and both bulls and cows when lean smell strongly of musk, their flesh at the same time being very dark and tough, and certainly far inferior to that of any other ruminating animal existing in North America.” The carcase of a Musk-Ox weighs, exclusive of fat, above 3 cwt. On this subject, Major Feilden? says: ‘The cause of the disagreeable odour which frequently taints the flesh of these animals has received no elucidation from my observations. It does not appear to be confined to either sex, or to any particular season of the year ; for a young unweaned animal, killed at its mother’s side and transferred within an hour to the stew-pans, was as rank and objectionable as any. The flesh of some of these animals of which I have partaken was dark, tender, and as well flavoured as that of four-year old Southdown mutton.” Remains of two fossil species of this genus (0. bombifrons and O. cavifrons) have been described from Pleistocene beds in the United States, the one from Kentucky and the other from the Arkansas River. Both (if indeed they be valid species) appear closely allied to the living form. Bovine Section.—Horns present and of nearly equal size in both sexes ; in form rounded or angulated, placed on or near the vertex of the skull, extending more or less outwards, and curving upwards near the extremities; external surface comparatively smooth and never marked by prominent transverse ridges or knobs. Muzzle broad, with large naked muffle ; nostrils lateral; no suborbital gland. Skull without any trace of lachrymal fossa or fissure. Tail long and cylindrical ; generally tufted at the extremity, rarely hairy throughout. Males usually with a dew-lap on the throat. No foot-glands. Molar teeth extremely hypsodont ; those of the upper jaw with a nearly square cross-section, and a large accessory inner column. The section is abundantly represented in the Palearctic, Oriental, and Ethiopian regions, with one Nearctic species and an outlying and aberrant species in Celebes. os.2—The whole of the species of Oxen were included by Linneus in the single genus Sos, and although the species have been distributed by modern zoologists in several genera—such as Anoa, Bubalus, Bison, Poéphagus, Bibos, and Bos—the characters by which they are separated are so slight that it seems, on the whole, preferable to retain the old genus in its original wide sense. Using then the term Bos in this sense, it will include all the representatives of the section—about a dozen in number—and may be divided into several groups. 1 Zoologist, September 1877. * Linn. Syst, Nad. 12th ed. vol. i. p. 98 (1766). BOVIDE 361 The first group includes the Buffaloes (genus Bubalus), chiefly characterised by their more or less flattened and angulated horns, which incline upwards and backwards, with an inward curve towards their tips, and are placed below the plane of the occiput, or vertex of the skull. The premaxille reach to the nasals, and the vomer is peculiar in being so much ossified as to join the posterior border of the palate. The back has a distinct ridge in the region of the withers; and the forehead is frequently convex. Oriental and Ethiopian region, and Celebes. The most generalised representative of this group is the small Anoa (B. depressicornis) of Celebes, the type of the genus Anoa or Probubalus, which has the same cranial structure as in the more typical Buffaloes, to the young of which (as was pointed out by the late Professor Garrod) it presents a striking resemblance. Its colour is black ; and the short and prismatic horns are directed upwards from the forehead. In the Pliocene Siwaliks of India there occur the remains of larger Buffaloes (B. occipitalis and B. acuticornis) closely allied to the Anoa, but with longer and more distinctly angulated horns. The still larger B. platyceros of the last-named deposits, in which the horns are wide-spreading and much flattened, appears to be in some respects intermediate between the preceding and following forms. The typical Indian Buffalo (Bos buffelus), which has been domesticated over South-East Asia, Egypt, and Southern Europe, is, in the wild state, a gigantic animal with enormous horns. These horns are longer, more slender, and more outwardly directed in the female than in the male; and in the former sex may have a length of more than 6 feet from base to tip. They are widely separated at their bases, the forehead is very convex, and the ears are not excessively large, and have no distinct fringe. These Buffaloes frequent swampy and moist dis- tricts in several parts of India, but it is in many instances difficult to decide whether they belong to really wild or to feral races. Very large skulls, specifically indistinguishable from those of the existing form, occur in the Pleistocene deposits of the Narbada valley in India; while an allied, if not specifically identical form, occurs in the Pliocene of the same country. There is some doubt whether B. antiquus of the Pleistocene of Algeria is most nearly related to the Indian or to the African species. In Africa two species of Buffalo are recognised by Sir Victor Brooke, namely the large B. caffer, occurring typically at the Cape, but said by this writer to range to Abyssinia, and the smaller B. pwmilus, which seems to have a very wide distribution. The skulls of both these forms are shorter than in the Indian species, while the horns are also shorter, much more curved inwardly, and more approximated on the forehead. In the large typical form of 1 Proc. Zool. Soc. 1878, p. 474. 362 UNGULATA L. caffer from South Africa the colour is black, the horns of the male are very thick, much reflected, and closely approximated on the forehead, where they form a helmet-like mass.1_ The large northern form described as B. wquinoctialis has the horns somewhat less thick, and thus approximates to the so-called B. pumilus. The latter occurs typically in Western Africa, where it has also been described as B. brachyceros. In the typical form the horns are thinner and less reflected than in B. caffer, and in some specimens they are more widely separated on the forehead, and are marked at their bases by distinct ruge. The colour is ruddy brown, inclining to rufous in one specimen. The skulls of Buffaloes from West Africa, probably referable to the form described as B. centralis, appear to connect B. pumilus with B. caffer, as shown by their larger size and the form of their horns; so that further observations are required to show whether the smaller form is really entitled to rank as a distinct species, or merely as a well-marked local race. The second group comprises the Bisons, which are more nearly allied to the true Oxen, having similar rounded horns, but the skull being less massive, with a longer and more tapering frontal region, and a wider frontal diameter. The superior part of the forehead is transversely arched, the intercornual space elevated in the middle, the horns situated below the plane of the occiput, and the orbits more or less prominent. The premaxillae do not extend upwards to reach the nasals. The Bisons (Fig. 148) have the body covered with short, crisp, woolly hair, while on the head and neck there is an abundance of much longer and darker hair, which forms a mane concealing the eyes, ears, and the bases of the horns. There is also a long beard beneath the chin; while a line of long hair extends from the head nearly to the tail, the latter being tufted at the extremity. The withers are much higher than the hind quarters, so that there is a kind of hump at the shoulders. The group is represented by two species—the European and the American Bison. The former is the Bos bonasus of Linnezus, and is also identical with the Bos bison of Ray. The German name Wisent is the equivalent of the Greek Bison. The American species is the Bos americanus of Gmelin. Both species are closely allied, but the American Bison is slightly the smaller animal of the two, and is shorter and weaker in the hind quarters, with a smaller pelvis; its body is, however, more massive in front ; and the hair on the head, neck, and fore quarters is longer and more luxuriant. A large bull American Bison, preserved in the Museum at Washington, stands 5 feet 8 inches in height at the withers. The European Bison appears to have been formerly ' Sir V. Brooke states that this species is distinguished from B. pumilus by the absence of a fringe to the ears, but specimens in the British Museum show that this is not the case. BOVIDE 363 abundant over a large portion of Europe in the Pleistocene period —the fossil race described as B. priscus not being specifically dis- tinct ; but at the present day it exists only in the primeval forests of Lithuania, Moldavia, Wallachia, and the Caucasus, where it is artificially preserved. The American Bison formerly ranged over about one-third of the North American continent. Thus, to quote from Mr. Horna- day,! “starting almost at tide-water on the Atlantic coast, it ex- tended across the Alleghany mountain system to the prairies along the Mississippi, and southward to the delta of that great system. Fic. 148.—The American Bison (Bos americanus). After Hornaday. Although the great plain country of the West was the natural home of the species, where it flourished most abundantly, it also wandered south across Texas to the burning plains of North-Eastern Mexico, westward across the Rocky Mountains into New Mexico, Utah, and Idaho, and northward across a vast treeless waste to the bleak and inhospitable shores of the Great Slave Lake itself.” In consequence of the settlement of the country by Europeans the area inhabited by the Bison was gradually contracted, till about 1840 one mighty herd occupied the centre of its former range. The completion of the Union Pacific Railway in 1869 divided this great herd into a southern and a northern division, the former comprising a number of individuals estimated at nearly four millions, while the latter contained about a million and a half. Before 1880 the southern herd had, however, practically ceased to exist ; while the same fate overtook the northern one in 1883. In 1889 some twenty stragglers in Texas represented the last of the southern herd ; while there were a few others in Colorado, Wyoming, Montana, 1 The Extirpation of the American Bison, 1889. 364 UNGULATA and Dakota. A herd of some two hundred wild individuals, derived from the northern herd, is preserved by the United States Government in the Yellowstone National Park; and it is believed that some five hundred of the race known as Wood-Bison exist in British territory ; but with these exceptions this magnificent species is exterminated. The multitudes in which the American Bison formerly existed are almost incredible ; the prairies being absolutely black with them as far as the eye could reach, and the numbers in the herds being, as we have said, reckoned by millions. Mr. Hornaday even considers that the whole of the game in South Ghee SR Na YN J iy SSAA Fic. 149.—The Yak (Bos grunniens), domestic variety. Africa was never equal to the number of Bison on an equal area of the American prairies. An extinct Bison from the Pleistocene of Texas, known as Bos latifrons, was probably the ancestor of the recent American species. The Yak (Bos .grunniens) appears to be allied both to the Bisons and the true Oxen, being distinguished from the former by the different position occupied by the long hair, which forms a fringe investing the shoulders, flanks, and thighs, and grows over the whole of the tail. In the skull the orbits are less tubular, the fore- head flatter, and the premaxille less widely separated from the nasals. There is no distinct dewlap. Wild Yaks inhabit the higher regions of Chinese Tibet and the region of the Karakoram as well as the more outlying parts of Ladak, such as the Chang- chemo valley. Owing, however, to incessant pursuit those now found within the territories of the Maharaja of Kashmir are stragglers BOVIDE 365 from Chinese Tibet. The height of the Yak is somewhat lower than that of the larger domestic cattle. The colour of the wild race is black, tending to brown on the flanks; but many of the tame breeds which have been crossed with ordinary cattle have more or less white (Fig. 149), and it is the white tails of these half-breeds that are so esteemed in India as “chowries.” Yaks are exceedingly intolerant of heat, and the wild ones always live at very great elevations. Tame Yaks are extensively used as beasts of burden in Tibet, where they are extremely valuable in crossing the high and desolate wastes of that region; they have, however, the great drawback that they refuse to eat corn, so that in districts where there is no grass it is frequently necessary to make forced marches with wearied beasts in order to prevent them (and thus the whole party) perishing from starvation. The skull of an extinct species from the Pliocene of Northern India, described as Bos sivalensis, appears to indicate a species allied to the Yak. With the Bibovine group we come to the consideration of three Oriental species which connect the preceding forms with the typical Oxen. The three species are the Gaur (B. gaurus) the Gayal (B. frontalis, Fig. 150) of India, and the Banteng (B. sondaicus) of Burma, Java, Bali, and Lambok. In this group, as in the true Oxen, there are thirteen pairs of ribs, against fourteen in the Bisons. All the three species are characterised by the great height of the spines of the anterior dorsal vertebra, causing a promi- nent ridge down the back. The horns, which are of a greenish colour in the Gaur, are somewhat flattened, and after running out- wards are directed upwards instead of backwards; they occupy the vertex of the skull. The frontals are more or less concave, the premaxille do not join the nasals, and the occipital aspect of the skull is characterised by the deep incisions made by the temporal fosse. The lower part of the legs is white (Fig. 150), and the hoofs are comparatively small and pointed. The Gaur (b. gaurus) is the largest of the three species, and inhabits all the large forests of India from near Cape Comorin to the foot of the Himalaya; it is commonly known to sportsmen as the Indian Bison. It stands fully 6 feet in height at the withers, which are much elevated ; and since the whole back is arched the line from the nose to the root of the tail forms an almost continuous curve. The most characteristic feature of the animal is, however, the large and convex intercornual frontal crest, which curves forward, and thus gives a concave profile to this part of the skull. Asa rule the Gaur prefers hilly regions, although it is sometimes met with on the flat. It is very shy and readily frightened ; and it has never been domesticated. The Gayal, or Mithan, of which a figure is given in woodcut 150, is at once dis- tinguished from the Gaur by the straight line between the horns 366 UNGULATA (which are black in colour), owing to the absence of the intercor- nual crest of the latter. The horns are also shorter, more rounded, and less curved. In the Indian Museum, Calcutta, there are, how- ever, skulls which are to a great extent intermediate between those of typical Gaurs and those of typical Gayals, but these may belong to hybrids. The Gayal occurs in Assam, Chittagong, and adjacent districts, but it appears that these animals exist in a semi-domestic- ated condition, no wild race being known to Europeans, although it is probable that such may exist in the unexplored Mishmi Hills. Fic. 150,—The Gayal (Bos frontalis). From Sclater, List of Animals in Zoological Society's Gardens, 1883. The Banteng (B. sondaicus) is a smaller and lighter built animal than either of the preceding, with a longer and sharper head, and more rounded and slender horns. The dorsal ridge is, moreover but slightly developed ; while the bright dun colour of the body of the female readily distinguishes it from the darker hue of the Gaur and Gayal. A. fossil skull from the Pleistocene deposits of the Narbada valley, India, described as Bos palwogaurus, is believed to indicate a species nearly allied to the Gaur, if indeed it be specifically distinct. BOVIDE 367 The true Oxen, or Taurine group, are now represented solely by Bos taurus and Bos indicus. Both of these species are now known only by domesticated races, unless the herds of the former preserved at Chillingham and some other British parks are the survivors of an original wild race. The dorsal ridge of the Bibovine group is here wanting; the horns are rounded, with their extremities directed backwards, and are placed at the extreme vertex of the skull; while the long frontal region is nearly flat; the temporal fossee scarcely intrude upon the occipital aspect of the skull; and the premaxille reach the nasals. The hoofs are large and rounded. It is known that wild Oxen were abundant in the forests of Europe at the time of Julius Cesar, by whom they were described as the Urus, equal to the German Aurochs ; and the large skulls found in turbary and Pleistocene deposits, and described under the name of Bos primigentus, can only be regarded as having belonged to the large original race of B. taurus, of which it has been thought the Chillingham cattle are smaller descendants! The subfossil skulls described as B. longifrons and B. frontosus must also be looked upon as referable to smaller races of the same species. That the domestic cattle of Europe are descendants from the various races of the same original species there can be no doubt, but in the case of the humped cattle of India (B. indicus) it is quite probable that their origin may be, at least in part, different. The extinct Bos namadicus, of the Pleistocene deposits of India, was a species with the general characters of the Taurine group, but with an inclination to a flattening of the horns, and with an approximation to a Bibovine type of occiput, as well as with the separation of the premaxille from the nasals. The earliest representatives of this group occur in the Pliocene of the Siwalik Hills in Northern India. One of these species (B. planifrons) appears to be allied to B. namadicus ; but the other (Bb. acutifrons) was a gigantic species characterised by the sharp median angulation of the frontal region, and the pyriform section of the enormous horn-cores. The extinct B. elatus, from the Upper Pliocene of France and Italy, is the representative of a generalised type, which may be known as the Leptobovine group. The males had rounded horn- cores widely separated at their bases, and placed low down on the forehead. The females (which have been described as Leptobos) were often or always hornless. The limbs were unusually slender. This group also occurs in the Pliocene of the Siwalik Hills. 1 The late Mr. Alston, Fauna of Scotland, ‘‘ Mammalia” (Glasgow, 1880), p. 25, considers that the Chillingham cattle are descendants of a race which had escaped from domestication. 368 UNGULATA Suborder PERISSODACTYLA This is a perfectly well-defined group of Ungulate mammals, represented in the actual fauna of the world by only three distinct types or families—the Tapirs, the Rhinoceroses, and the Horses— poor in genera and species, and (except in the case of the two domesticated species of Equus, which have been largely multiplied and diffused by man’s agency) not generally numerous in individuals, though widely scattered over the earth’s surface. Paleontological Fic. 151.—Bones of right fore foot of existing Perissodactyles. A, Tapir (Tapirus indicus), x4; B, Rhinoceros (Rhinoceros sumatrensis), x}; C, Horse (Equus caballus), x2. U, ulna; R, radius ; c, cuneiform ; 7, lunar; s, seaphoid; u, unciform; m, Magnum ; td, trapezoid ; tm trapezium.—From Flower, Osteology of Mammalia. records, however, show very clearly that these are but the surviving remnants of a very extensive and much-varied assemblage of animals, which flourished upon the earth through the Tertiary geological period, and which, if it could be reconstructed in its entirety, would not only show members filling up structurally the intervals between the existing apparently isolated forms, but would also show several marked lines of specialisation which have become extinct without leaving any direct successors. The following are the principal characters distinguishing them from the Artiodactyla. Premolar and molar teeth in continuous series, with massive, quadrate, transversely ridged or complex crowns,—the posterior premolars often resembling the true molars PERISSODACTYLA 369 in size and structure. Crown of the last lower molar commonly bilobed, and if a third lobe is present in this tooth it is wanting in the last lower milk-molar. Dorso-lumbar vertebrae never fewer than twenty-two, usually twenty-three in the existing species. Nasal bones expanded posteriorly. An alisphenoid canal. Femur with a third trochanter! The middle or third digit on both fore and hind feet larger than any of the others, and symmetrical in itself, the free border of the ungual phalanx being evenly rounded (see Fig. 151). This may be the only functional toe, or the second and fourth may be subequally developed on each side of it. In the Tapirs and many extinct forms, the fifth toe also remains on the fore limb, but its presence does not interfere with the symmetrical arrangement of the remainder of the foot around the median line of the third or middle digit. Traces of a hallux have only been found in some extremely ancient and primitive forms. The astragalus has a pulley-like surface above for articulation with the tibia, but its distal surface is flattened and unites to a much greater extent with the navicular than with the cuboid, which bone is of comparatively less importance than in the Artiodactyla. The calcaneum does not articulate with the lower or distal extremity of the fibula. The stomach is always simple, the cecum is large and capacious, the placenta diffused, and the mamme are inguinal. The gall-bladder is invariably absent. As regards the dentition, the whole of the premolar series may be preceded by milk-teeth ; and it has been demonstrated in Rhinoceros that when there is no displacement of the first cheek- tooth that tooth is a persistent milk-molar; the same condition apparently holding good in Paleotherium. This feature indicates considerable dental specialisation, the milk-molars, according to the theory generally accepted by the leading English zoologists, being the acquired, and the premolars the original series. Another peculiar feature of the dentition of the Perissodactyla, very rarely met with among the Artiodactyla, is that the premolars tend to resemble the true molars ; this feature occurring in all the existing genera, although not found in the earlier generalised types. The cheek-teeth of all the members of the suborder are primarily con- structed on some modification of what is known as the lophodont plan. Thus the upper molars (Fig. 155, p. 375) have an outer antero- posterior wall from which proceed two transverse ridges, formed by the coalescence of the primitive inner and outer columns, towards the inner aspect of the crown; while in the lower molars there may be either two simple transverse ridges, or these ridges may be curved into crescents, coming into contact with one another at their extremities. Those forms having brachydont teeth show this plan of structure in its simplest modification; but in cases, as in the 1 Wanting in the aberrant Chalicotherium. 24 370 UNGULATA Horse, where the teeth assume an extremely hypsodont form, the original plan is so obscured by infoldings of the enamel that it can only be traced with difficulty. At the present day the Perissodactyla are sharply differ- entiated into Horses, Tapirs, and Rhinoceroses, but the knowledge already gained of the extinct representatives of the suborder shows such a close alliance between these groups that it is exceedingly difficult to make any satisfactory classification of the whole. This is of course exactly what might have been expected ; and the same would doubtless be the case with all other groups if we knew as much of their past history as we do of that of the Perissodactyles. The detailed account of the anatomy of the Horse given in the sequel will afford much information as to the general structure of the members of the suborder. Family TAPIRIDE. Both upper and lower cheek-teeth brachydont and simply bilophodont; hinder premolars as complex as the molars ; last lower molar without third lobe; first upper cheek-tooth with a milk- predecessor.t Outer columns of upper molars conical. Four digits in the manus, and three in the pes. Tapirus.2—Dentition i $, ¢ 1, p 4, m 3; total 42. Of the upper incisors, the first and second are nearly equal, with short, broad crowns; the third is large and conical, considerably larger than the canine, which is separated from it by an interval. Lower incisors diminishing in size from the first to the third ; the canine, which is in contact with the third incisor, large and conical, working against (and behind) the canine-like third upper incisor. In both jaws there is a diastema between the canines and the commence- ment of the teeth of the cheek-series, which are all in contact. First upper premolar with a triangular crown, narrow in front owing to the absence of the anterior inner cusp. The other upper premolars and molars all formed on the same plan and of nearly the same size, with four roots and quadrate crowns, rather wider transversely than from before backwards, each having four cusps, connected by a pair of transverse ridges, anterior and posterior. The first lower premolar compressed in front ; the others composed of a simple pair of transverse crests, with a small anterior and posterior cingular ridge. Skull elevated and compressed. Orbit and temporal fossa widely continuous, there being no true postorbital process from the frontal bone. Anterior narial apertures very large, and extend- ing high on the face between the orbits; nasal bones short, elevated, 1 See W. N. Parker, Proc. Zool. Soc. 1882, Pe 075: * Cuvier, Tableaw Elément, de UV Hist. Nat. p. 152 (1798) ; ex Brisson. TAPIRIDE 371 triangular, and pointed in front. Vertebre: C 7, D 18, L 5, S 6, C about 12. Limbs short and stout. Fore feet with four toes, having distinct hoofs: the first is absent, the third the longest, the second and fourth nearly equal, the fifth the shortest and scarcely reaching the ground in the ordinary standing position. Hind feet with the typical Perissodactyle arrangement of three toes,—the middle one being the largest, the two others nearly equal. Nose and upper lip elongated into a flexible, mobile snout or short pro- boscis, near the end of which the nostrils are situated. Eyes rather small. Ears of moderate size, ovate, erect. Tail very short. Skin thick and but scantily covered with hair. The existing species of Tapir may be grouped into two sections, the distinctive characters of which are only recognisable in the skeleton. (A) With a great anterior prolongation of the ossifica- tion of the nasal septum (mesethmoid), extending in the adult far beyond the nasal bones, and supported and embraced at the base by ascending plates from the maxille (genus Elasmognathus, Gill). Two species, both from Central America, Tapirus bairdi and T. dowi. The former is found in Mexico, Honduras, Nicaragua, Costa Rica, and Panama; the latter in Guatemala, Nicaragua, and Costa Rica. (B) With ossification of the septum not extending farther forward than the nasal bones (Zapirus proper). Three species, 7. indicus, the largest of the genus, from the Malay Peninsula (as far north as Tavoy and Mergui), Sumatra, and Borneo, distinguished by its peculiar coloration, the head, neck, fore and hind limhs, being glossy black, and the intermediate part of the body white; 7. americanus (T. terrestris, Linn.), the common Tapir of the forests and lowlands of Brazil and Paraguay (Fig. 152); and 7. roulini, the Pinchaque Tapir of the high regions of the Andes. All the American species are of a nearly uniform dark brown or blackish colour when adult ; but it is a curious circumstance that when young (and in this the Malay species conforms with the others) they are conspicuously marked with spots and longitudinal stripes of white or fawn colour on a darker ground. The habits of all the kinds of Tapirs appear to be very similar. They are solitary, nocturnal, shy, and inoffensive, chiefly frequent- ing the depths of shady forests and the neighbourhood of water, to which they frequently resort for the purpose of bathing, and in which they often take refuge when pursued. They feed on various vegetable substances, as shoots of trees and bushes, buds, and leaves. They are hunted by the natives of the lands in which oie live for the sake of their hides and flesh. The singular fact of the existence of so closely allied animals. as the Malayan and the American Tapirs in such distant regions of the earth, and in no intervening places, is accounted for by what is known of the geological history of the race; for the Tapirs must 372 UNGULATA once have had a very wide distribution. There is no proof of their having lived in the Eocene epoch, but in deposits of Miocene and Pliocene date remains undistinguishable generically from the modern Tapirs, and described as 7. priscus, T. arvernensis, etc., have been found in France, Germany, and in the Red Crag of Suffolk. Tapirs appear, however, to have become extinct in Europe before the Pleistocene period, since none of their bones or teeth have been found in any of the caverns or alluvial deposits in which those of Elephants, Rhinoceroses, and Hippopotamuses occur in abundance; but in other regions their distribution at this age was far wider than at present, Fia. 152.—The American Tapir (Tapirus americanus). as they are known to have extended eastward to China (Z. sinensis, Owen) and westwards over the greater part of the southern United States of America, from South Carolina to California. Lund also distinguished two species or varieties from the caves of Brazil, one of which appears identical with 7. americanus. Thus we have no difficulty in tracing the common origin in the Miocene Tapirs of Europe of the now widely separated American and Asiatic species. It is, moreover, interesting to observe how very slight an amount of variation has taken place in forms isolated durin enormous period of time. . The anatomy of the soft parts of the Tapirs! conforms to the g such an 1 See J. Murie, Journ. Anat. and Physiol. vol. vi. p. 131, 1871 ; W.N,. Parker Proce, Zool. Soc, 1882, p. 768; and F. E. Beddard, Proc. Zool. Soc. 1889, p. 252, ; LOPHIODONTIDAZ 373 general Perissodactyle type, as exemplified in the Rhinoceros and the Horse, although on the whole (as might have been expected) presenting a closer resemblance to the former. 7. americanus differs from 7. indicus by the absence, or at any rate the less development, of the intestinal valvule conniventes, the presence of a moderator band in the heart, the shape of the glans penis, and the more elongated cecum, which is sacculated by four dis- tinct longitudinal fibrous bands. The convolutions of the hemi- spheres of the brain of the Tapirs are simpler than in other Perisso- dactyles, thus tending to confirm the inferences which may be drawn from the skeleton and teeth as to the comparatively low or general- ised organisation of these animals. Palceotapirus—This name has been applied to an imperfectly known form from the Upper Eocene Phosphorites of Central France, which is regarded by Dr. Filhol as referable to this family. Family LOPHIODONTID&. Molars brachydont and bilophodont, those of the lower jaw with either straight or imperfectly crescentoid ridges ; premolars smaller and usually simpler than the molars; last lower molar generally with a third lobe. Outer columns of upper molars conical or flattened. Digits usually as in the preceding family. This family includes a number of more or less imperfectly known forms, all of which are extinct and apparently confined to the Eocene period, and ranging from the size of a Rabbit to that of a Rhinoceros. Although some .of these appear to have died out without giving rise to more specialised forms, it is probable that this family contained the ancestral types from which most or all of the modern Perissodactyles have been derived. Only very brief mention can be made here of some of the leading genera. Lophiodon, of the Middle and Upper Eocene of Europe, with the dental formula, i 3,¢4, p 3, m 3, includes the largest representatives of the family, and is generally regarded as a stock which has died out without giving rise to later forms. The ridges of the lower molars are straight, and the last of these teeth has a third lobe; while the second transverse ridge of the last upper premolar is usually incom- plete ; the outer columns of the upper molars are flattened, as in the next genus. Hyrachyus, of the Upper Eocene of the United States, and probably also occurring in the French Eocenes, is an allied genus, with four premolars and no third lobe to the last lower molar; the fourth upper premolar having the two ridges uniting internally to form a crescent. This genus has been regarded as the ancestor of the Rhinocerotic Hyracodon. The genus Hyracotherium was established in 1839 by Owen for a small animal no larger than a Hare, the skull of which was found in the London Clay at Herne 374 UNGULATA Bay. A more nearly perfect specimen, apparently of the same species, was afterwards (in 1857) described under the name of Pliolophus vulpn- ceps, of which the skull is figured in the accompanying woodcut. Other forms referable to the same genus have been obtained from the Wasatch Eocene of the United States, and were described by Professor Marsh under the name of Hohippus. There were four premolars, the fourth being unlike the molars, and in the upper jaw having only one inner cusp. The upper molars are of the general type of those of Lophiodon, but have conical outer columns, and the anterior transverse ridge imperfect, while the ridges of the lower molars are crescentoid. Systemodon differs from Hyracotherium Fic. 153.—Right side of skull of Hyracotherinum leporinum, from the London Clay. }natural size. (After Owen.) 3, Occiput; 7, sagittal crest; 11, frontals ; 15, nasals ; 21, maxilla; 22, premaxilla ; d, mandibular condyle ; a, aperture of facial nerve ; 1-4, premolars ; m 1-3, molars. by the absence of a diastema between the first and second pre- molars; it occurs in the Wasatch Lower Eocene of the United States. In Pachynolophus (Lophiotherium, Orotherium, or Orohippus), which is common to the Middle and Upper Eocene of Europe and the Bridger Eocene of North America, the outer columns of the upper molars are flattened, and in some cases, at least, the last premolar resembles the molars, that of the upper jaw having two inner cusps.1 This genus, indeed, so closely connects Hyracotherium with the genera Epihippus and Anchilophus as to show that the distinction between the Lophiodontide and Palwotheriide is really an arbitrary one. Lpihippus, of the Upper Eocene of the United States, has both the third and fourth upper premolars as complex in the molars, and is distinguished from Anchilophus by the lower cusps and more imperfect transverse ridges of these teeth. The so-called Orohippus agilis belongs to this genus. Isectolophus is another American Eocene genus which may be provisionally placed in this family ; it is regarded by Professors Scott and Osborn as connecting Systemodon ’ The Swiss P. siderolithicus has only one cusp in the last upper premolar, PALELOTHERHDA 375 with the Tapiride; the fourth and probably the third upper pre- molar approximating in structure to the molars; the upper molars have conical outer columns. Helaletes is another closely allied form, with similar premolars, but with the outer columns of the upper molars flattened. Family PALZOTHERIIDE. __ Molars (Fig. 155) brachydont, with the valleys between the ridges never filled with cement ; upper premolars either simpler than Fic. 154.—Restoration of Palceotherium (Upper Eocene). After Cuvier. or as complex as the molars ; lower molars with crescentoid ridges, and the last of the series with or without a third lobe. Outer columns of upper molars flattened. f f Orbit (at least usually) confluent a : with temporal fossa. Three digits on each foot. This family in- cludes extinct genera ranging from ( the Middle and Upper Eocene to ¢ {Mf the Miocene, and passes so gradu- ~ } ally into the following one that the maintenance of the two can only be supported on the ground of convenience. The typical genus, Paleotherium, was made known to Fic. 155.—A half-worn right upper molar of science in the early part of the present century by Cuvier, who restored the skeleton (Fig. 154) with a short neck like that of the Tapirs, although it has been sub- sequently found that the neck was considerably longer. This Palcotherium magnum. (After Owen.) fF External surfaces of outer columns ; @, postero- external column (metacone); b, antero-ex- ternal column (paracone); ¢, postero-internal column (hypocone) ; @, antero-internal column (protocone); 4, anterior intermediate column (protoconule); ¢, median valley; g, posterior valley. genus (which may be taken to include Paloplotherium) ranges from 376 UNGULATA the Middle to the Upper Eocene of Europe, and usually has the full typical dentition, although the first premolar may disappear. The last lower molar has a third lobe ; and in the typical forms the last premolar is as complex as the molars, the diastema is short, and the canines are not large. In other forms, however, the hinder ridge of the fourth upper premolar may be aborted. The first upper cheek- tooth is generally a well-developed tooth, which may have a deciduous predecessor. Anchilophus, of the Upper Eocene of Europe, and Anchitherium, of the Miocene of Europe and North America, connect the preceding forms with the Zquide. In the latter genus there is the full number of teeth, the last lower molar has almost completely lost the third lobe of Anchilophus, and the surfaces of the two outer lobes of the upper molars (Figs. 157, 158) lack the median vertical ridges of that genus. In the American species of Anchitherium (which have been described as Mesohippus and Miohippus) the lateral digits are larger than in the European Middle Miocene Anchitheriwm aurelianense ; a mere splint represents the fifth metacarpal, and the meso- and ento-cuneiform of the tarsus do not unite as they do in the latter. Family EQuip. Molars hypsodont, with the outer columns of the upper ones flattened, the valleys completely filled with cement, and the enamel thrown into folds and plications ; upper premolars as complex as molars, which they slightly exceed in size; ridges of lower molars crescentoid, and complicated by enamel-foldings ; no distinct third lobe to last lower molar; summits of incisors with a central infold- ing of enamel. Orbit completely surrounded by bone. Digits three or one, but in the former case the median one is alone of functional importance ; ulna and fibula incomplete ; meso- and ento- cuneiform of tarsus united. Such are the leading characters which serve to distinguish the existing Horses and their nearest fossil allies from the Palwotheriide. The Horse, as being the best known of the Perissodactyle Ungu- lates, is selected for a somewhat detailed description ; but before proceeding to this it will be advisable to take a brief survey of the relations of the Hquide to the extinct forms already noticed, and also of the modifications of the family at present existing. The earliest form which can be certainly included in this line of descent is the American Lower Eocene genus Phenacodus (noticed - below under the head of the suborder Condylarthra), in which there were five complete digits to the feet. From this form there is but a step to Systemodon and Hyracotherium, in which the func- tional digits of the manus were reduced to four, as in Pachynolophus EQUIDAE B77. (Fig. 156, a), although one species retained a rudiment of the metacarpal of the pollex. The transition from these animals of the Eocene period to the Horses of modern times has been accompanied by a gradual increase in size. The diminutive Hyracotherium of the Lower, and Pachy- nolophus of the Middle and Upper Eocene were succeeded in the Miocene period by the forms to which the name of Anchitherium has been given, of the size of sheep; these again in Pliocene times by Hipparion and Protohippus, as large as the modern donkeys ; and it is mainly in the Pleistocene period that Equide occur which approach in size the existing Horse. Important structural modi- fications have also taken place, with corresponding changes in the Fic. 156.—Successive stages of modification of the feet of extinct forms of Horse -like animals (chiefly from Marsh), showing gradual reduction of the outer and enlargement of the middle toe (111). a, Pachynolophus (Eocene) ; b, Anchitheriwm (Early Miocene) ; c, Anchitherium (Late Miocene); , Hipparion (Pliocene); e, Equus (Pleistocene). mode of life of the animal. Thus the neck has become elongated, the skull altered in form, the teeth greatly modified, and the limbs have undergone remarkable changes. The last two require to be described more in detail. The teeth in the Eocene forms had, as mentioned above, the characteristic number of forty-four. This number has been retained throughout the series, at least theoretically; but one tooth on either side of each jaw, the anterior premolar, which in all the Eocene and Miocene species was well developed, persisting through the lifetime of the animal, is in all modern Horses rudimentary, functionless, and generally lost at an early period of life, evidently passing through a stage which must soon lead to its complete dis- appearance. The canines have also greatly diminished in size, and are rarely present in the female sex, so that practically a very large number of adult Horses of the present day have eight teeth less than the number possessed by their predecessors. The diastema or interval between the incisor and premolar teeth (of essential 378 UNGULATA importance in the domesticated Horse to his master, as without it there would be no room for inserting the special instrument of subjugation to his commands, the bit) already existed in the earliest known forms, but has gradually increased in length. The incisors have undergone in comparatively recent times that curious change producing the structure more fully described hereafter, which distinguishes the Horse’s incisors from those of all other known animals, with the exception of the extinct Muacrauchenia. Lastly, the molars have undergone a remarkable series of modi- fications, much resembling in principle those that have taken place in several other groups of herbivorous animals. Distinctions in form which existed between the premolars, at least in the anterior part of the series, and the true molars have gradually dis- appeared, the teeth becoming all very uniform in the shape and structure of their grinding surface. The crowns of all these teeth a b ie Fic. 157.—a, Grinding surface of unworn molar tooth of Anchitherium ; b, corresponding surface of unworn molar of young Horse; c, the same tooth after it has been some time in use. The uncoloured portions are the dentine or ivory, the shaded parts the cement filling the cavities and surrounding the exterior. The black line separating these two structures is the enamel or hardest constituent of the tooth. in the early forms were very short (see Fig. 158, a); there was a distinct constriction, or neck, between the crown and roots; and when the tooth was developing, as soon as the neck once rose fairly above the alveolar margin, the tooth remained permanently in this position. The term “brachydont” expresses this condition of teeth, the mode of growth of which may be illustrated by those of man. The free surface had two nearly transverse curved ridges, with valleys between (Fig. 157, a); but the valleys were shallow and had no deposit of cement filling them, the whole exposed surface of the unworn tooth. being formed of enamel. When the ridges became worn down the dentine of the interior was exposed, forming islands surrounded by enamel. With the progress of time the crowns of the teeth gradually became longer, the valleys deeper, and the ridges not only more elevated but more curved and com- plex in arrangement. To give support to these high ridges and save them from breaking in use, the valleys or cavities between them became filled up to the top with cement, and as the crown wore down an admirable grinding surface consisting of patches and EQUIDA 379 islands of the two softer substances, dentine and cement, separated by variously reduplicated and contorted lines of intensely hard enamel, resulted (Fig. 157, ¢). The crown continued lengthening until in the modern Horses it has assumed the form called “ hyps- odont” (Fig. 158, 3). Instead of contracting into a neck, and forming roots, its sides continue parallel for a considerable depth in the socket, and as the surface wears away, the whole tooth slowly pushes up, and maintains the grinding edge constantly at the same level above the alveolus, much as in the perpetually growing Rodent’s teeth. But in existing Horses there is still a limit to the growth of the molar. After a length is attained which in normal conditions supplies sufficient grind- ing surface for the lifetime of the animal, a neck and roots are formed, and the tooth is reduced to the condition of that of the brachydont ancestor. It is per- fectly clear that this lengthening of the crown adds greatly to the power of the teeth as organs of mastication, and en- sD ables the animals in which it has taken Fic. 158.—a, Outer view of second place to find their sustenance among the Upper molar tooth of Anchitheriwm, . (brachydont form); b, corresponding comparatively dry and harsh herbage tooth of Horse (hypsodont form). of the open plains, instead of being limited to the more succulent vegetable productions of the marshes and forests in which their predecessors probably dwelt. The modifications of the limbs which took place pari passu with those of the teeth must have been associated with increased speed, especially over firm and unyielding ground. Short, stout legs, and broad feet, with numerous toes, spreading apart from each other when the weight of the creature is borne on them, are sufficiently well adapted for plodding deliberately over marshy and yielding surfaces, and the Tapirs and the Rhinoceroses, which in the structure of the limbs have altered but little from the primitive Eocene forms, still haunt the borders of streams and lakes and the shady depths of the forests, as was probably the habit of their ancient representatives, while the Horses are all inhabitants of the open plains, for life in which their whole organisation is in the most eminent degree adapted. The length and mobility of the neck, position of the eye and ear, and great development of the organ of smell, give them ample means of becoming aware of the approach of enemies, while the length of their limbs, the angles the different segments form with each other, and especially the combination of firmness, stability, and lightness in the reduction of all the toes to a single one, upon which the whole weight of the body and all the muscular power are concentrated, give them speed a 380 UNGULATA and endurance surpassing that of almost any other animal. When surprised, however, they are by no means helpless, both fore and hind feet becoming at need powerful weapons of defence. If we were not so habituated to the sight of the Horse as hardly ever to consider its structure, we should greatly marvel at being told of a mammal so strangely constructed that it had but a single toe on each extremity, on the end of the nail of which it walked or galloped. Such a conformation is without a parallel in the vertebrate series, and is one of the most remarkable instances of specialisation, or deviation from the usual type, in accordance with particular conditions of life. It is clear, both from the structure of the foot itself, and also by an examination of the intermediate forms, that this toe corresponds to the middle or third digit of the complete typical or pentadactyle foot; and there is very strong evidence to show that by a gradual concentration of all the power of the limb upon this toe, and the concurrent dwindling away and final dis- appearance of all the others, the present condition of the Horse’s foot has been produced. Protohippus.1—tIn this Lower Pliocene North American genus (also described as Merychippus) the cheek-teeth resemble those of the generalised species of Equus, but have shorter crowns; while the milk-molars approximate to the permanent molars of Anchi- theriwm. Each foot has three digits. Hipparion.2— Upper cheek-teeth (Fig. 159), with the antero- Fic. 159.—Three right upper cheek-teeth of Hipparion. «a, Antero-external column ; b, postero-external column; ¢, postero-internal column, or posterior pillar; d, antero-internal column, or anterior pillar; f, posterior intermediate column; i, anterior intermediate column. (From the Palcontologia Indica.) internal column, or anterior pillar as it may be conveniently termed in this family, detached throughout the greater part of its height from the adjacent column. Lither a single or three digits in each foot. First upper premolar large and persistent. This genus was very widely distributed in the Pliocene, occurring in Europe, Asia, and North America. In the typical European forms, and also in those 1 Leidy, Proc. Ac. Nat. Sci. Philad. 1858, p. 26. * Christol, Ann. Set. Indust. Mid. France, vol. i. p. 180 (1832). EQUIDE 381 of North America, there were three digits in the feet (Fig. 156, d); but in the Indian H. antilopinum (separated by Cope as Hippo- dactylus) the lateral digits seem to have disappeared. There is some doubt whether or no Hipparion should occupy a place in the direct ancestry of the Horse, and Professor Cope suggests that while in America the intermediate place between Anchitherium and Equus was held by Protohippus, in Europe the same position was occupied by Hipparion—a view which involves the dual origin of the Horses of the New and Old Worlds. Equus1—Upper cheek-teeth with the anterior pillar (except in a very early stage of wear) joined by a narrow neck to the adjacent column (Fig. 157, c). Each foot with a single complete digit, but with remnants of the proximal portions of the second and fourth metapodials (Fig. 156, ¢); some extinct forms having claw-like rudiments of the terminal phalangeals of the lateral digits. First upper premolar very small or altogether absent in existing species, but in some fossil species larger and persistent; first lower premolar only occasionally developed in some fossil forms. Ears long. Tail long, with long hairs either at the end or throughout. F. Cuvier, Dents des Mammiferes, p. 256 (1825) HYSTRICIDE 485 (E. dorsatus), a stout heavily-built animal, with long hairs almost or quite hiding the spines; four anterior and five posterior toes ; and a short stumpy tail. It is a native of the greater part of Canada and the United States where there is any remnant of the original forest left. Remains of Hrethizon occur in cavern-deposits in Pennsylvania. Synetheres1—This genus contains some eight or ten species, known as Tree Porcupines (Fig. 214), found throughout the tropical ni WAN \\\ \ \ AY . WS Nyt WP Fic. 214.—The Tree Porcupine (Synetheres prehensitlis). parts of South America, and one of them extending northwards into Mexico. They are of a lighter build than the Ground Porcupines, are covered with short, close, many-coloured spines, often mixed with hairs, and their tails are always prehensile. Their hind feet have only four toes, owing to the suppression of the hallux; but they have a peculiar fleshy pad on the inner side of the foot, between which and the toes boughs and other objects can be firmly grasped as with ahand. Vertebre: C7,D17,L5,8 3, C 36. An extinct species of this genus has been described from the cavern-deposits of Brazil. 1B. Cuvier, A/ém. du Muséum, vol. ix. p. 413 (1822). ‘‘ Sinéthére.” 486 RODENTIA Chetomys.1—Distinguished by the shape of its skull and the greater complexity of its teeth. It contains only one species (C. subspinosus), a native of the hottest parts of Brazil. Subfamily Hystricinzee.—Molars semi-rooted; clavicles incom- plete; soles smooth ; a rudimentary pollex; six mamme ; tail not prehensile. Now confined to the Old World, where they occur in Southern Europe, Africa, India, and the Malay Archipelago as far eastwards as Borneo. Habits terrestrial and nocturnal. Three genera, Hystriz.2—This genus is readily characterised by the inflated skull, in which the nasal chamber is often considerably larger than Fig. 215.—The Common Porcupine (Hystrix cristata). the brain-case, and by the short tail, tipped with numerous slender stalked open quills, which make a loud rattling noise when the animal moves. Vertebre: C 7,D15,L4,8 4,012. The best- known member is the Common Porcupine (H. cristata, Fig. 215), which occurs throughout Southern Europe and North and West Africa, but is replaced in South Africa by H. africe-australis, and in India by the Hairy-nosed Porcupine (H. leucura). The following account of the habits of the last-named species is from Dr. Jerdon: “ Hystria leucwra is found over a great part of India, from the lower ranges of the Himalayas to the extreme south but does not occur in lower Bengal, where it is replaced by H. bengalensis. It forms extensive burrows, often in societies, in the sides of hills, banks of rivers and nullas, and very often in the 1 Gray, Proc. Zool. Soc. 1843, p. 21. * Linn. Syst, Nat. 12th ed. vol. i. p. 76 (1766). CHINCHILLIDA 487 dams of tanks, and in old mud walls, etc. In some parts of the country they are very destructive to various crops, potatoes, carrots, and other vegetables. They never issue forth till after dark, but now and then one will be found returning to his lair in daylight. Dogs take up the scent of the Porcupine very keenly, and on the Nilghiris I have killed many by the aid of dogs, tracking them to their dens. They charge backwards at their foes, erecting their spines at the same time, and dogs generally get seriously in- jured by their strong spines, which are sometimes driven deeply into the assailant. The Porcupine is not bad eating,—the meat, which is white, tasting something between pork and veal.” Besides these three large crested species of Hystriz, there are four or five smaller species without nuchal crests occurring in North-East India and in the Malay region, from Nipal to Borneo. Fossil species of Hystriz occur in the Pleistocene and Pliocene of India, and in Europe from the Upper Pliocene to the Middle Miocene, being perhaps also represented in the French Phosphorites. Remains from the Pliocene and Miocene of the United States have been referred to this genus, and if rightly determined are of especial interest from a distributional point of view. Atherura.1—The Brush-tailed Poreupines are much smaller animals than the last, characterised by their long tails tipped with bundles of peculiar flattened spines. Of the three species two are found in the Malay region and one in West Africa. A fossil species occurs in the cavern-deposits of Madras. Trichys.2— This genus contains but one Bornean species (7. guenthert), externally very like an Atherura, but differmg from the members of that genus in many important cranial characters. Family CHINCHILLIDA. Terrestrial forms, with elongated hind limbs, bushy tails, very soft fur, and complete clavicles. Jugal without an inferior angle, and extending forwards to the lachrymal ; palate contracted in front and deeply emarginate behind; incisors short, and the molars — divided by continuous enamel-folds into transverse lamine. Neo- tropical region. This family includes only three existing species, divided into as many genera. Chinchilla.2—In this genus the fore feet have five and the hind four digits, the tail is long and bushy, and the auditory bull are enormous, appearing on the top of the skull. The one species (C. lanigera) is restricted to the alpine zones of the Andes from the north of Peru to the south of Chili. It is a Squirrel-like Rodent, 1 Cuvier, Regne-Animal, 2d ed. vol. i. p. 215 (1829). ‘* Atherure.” 2 Giinther, Proc. Zool. Soc. 1876, p. 739. 3 Bennett, Gardens, etc. Zool. Soc. pt. i. p. i. (1829). 488 RODENTIA about 10 inches in length, the tail somewhat exceeding 5 inches, and the ears very large. Its fur is greatly valued on account of its extreme softness and delicate gray colour. Lagidium+ and Lagostomus.°—Lagidium has four digits in both fore and hind feet, and Lagostomus three only in the hind feet, and the auditory bulle are much smaller than in the preceding genus. Lagidivim has the same distribution as Chinchilla - while Lagestomus, as represented by the Viscacha (L. trichodactilus), is found in the Pampas from the Uruguay River to the Rio Negro. The Viscachas live in burrows, generally in large numbers, and are nocturnal in their habits. Remains referable to the existing species, as well as others which appear to belong to extinct forms, occur in the Pleistocene deposits of South America. Extinct Genera. —Several Rodents from the South American Tertiaries more or less closely allied to Lagastomus have been described by Dr. Ameghino under the names of Prolagostoimus. Plislagostemus, ete. The huge Megamys (Potamarchus), from the infra-Pampean deposits of Parana and Patagonia, is referred to this family, and has dimensions approximating to those of an Ox. Other fossil genera have received the names of Eyillema and Tetra- stylus. Family CASTOROIDID. Castoroides.°—The large Beaver-like Rodent with the dimensions of a Bear from the Pleistocene of the United States described under this name is regarded by Dr. Coues as the type of a family. Its dentition is nearest to that of Chinchilla and Hydrocherus, but some of the cranial characters are like those of the Custoride. The genera Amblyrhiza and Loxomilus, from the Pleistocene of the Antilles, appear to be allied types. Family DASYPROCTIDE. Terrestrial forms with subequal limbs, hoof-like claws, short or obsolete tail, and rudimentary clavicles. Mandibular masseteric ridge obsolete ; palate broad; incisors long: molars semi - rooted, with external and internal enamel-folds. Neotropical region. Dasyprocta.s—Includes several slender-limbed species, with three hind toes, commonly called Agoutis, inhabiting Central and South America, one (D. cristata) extending into the West-Indian Islands. Numerous fossil remains of this genus occur in the cavern-deposits of Brazil. 1 Meyer, Vora deta Ac. Cars. Lrop.-Car. vol. xvi. p- 576 (1833). ? Brooks, Trans. Linn. Sor. yol. xvi. p. 102 (1828. ® Foster, Sccond Rep. Geol. of Ohio, p- $1 (183s . * Illiger, Prodromus Sust. Memin. p. 93 (1811). DINOM VID4E—CAVUDA 489 Celogenys.i—tThis genus is readily characterised by the presence of five hind toes, and the extraordinary development of its zygo- matic arches, which are enormously expanded vertically, forming great convex bony capsules on the sides of the face, enclosing on each side a large cavity lined with mucous membrane, and communicating by a small opening with the mouth. The Paca C. paca) is about 2 feet long, and, like the species of Dasyprocta, lives generally in the forests or along the banks of rivers. This species appears to date from the epoch of the Pleistocene deposits of the Brazilian caves. A smaller species from Ecuador, living at ele- vations of from 6000 to 10,000 feet, has been described as C. taczunowskti. Family DINOMYIDA. Distinguished from the Dasyproctide by the cleft upper lip, rather long and bushy tail, the presence of four digits in both fore and hind feet, and the complete clavicles. The manubrium is broad ; the optic foramina are confluent; the incisors broad ; and the molars rootless, with enamel-folds dividing them into transverse lamine. Dinonys.2—The sole representative of this family is the Rodent known as D. branicki, of which hitherto only a single specimen has been obtained. This was captured in Peru, where it was found at daybreak walking about a courtyard; the inhabitants of the dis- trict were previously unacquainted with the species, from which its extreme rarity may be inferred. Externally it resembles much the Paca, having similar S-like nostrils; but in the laminated molars, and many features of the skeleton, it differs from all the other Rodents with hoof-like nails. It is regarded by its describer, the late Professor Peters, as a connecting link between the Octodontide, Chinchillide, Dasyproctide, and Caviide. Family CAVUD®. Terrestrial or natatorial forms, with short incisors, strong man- dibular masseteric ridges, long and curved paroccipitals, and palate contracted in front. Fore feet with four digits, hind feet with three. Clavicles imperfect. Molars divided by enamel-folds into transverse laminz ; milk-teeth shed before birth. Other characters as in Dasyproctide. Neotropical region. Cavia.3—Limbs and ears short, subequal ; tail none. Vertebre : C7,D13,L6,84,C7. This genus includes several species widely 1 F. Cuvier, Ann. du Afuséwm, vol. x. p. 208 (1807). * Peters, Monatsber, Ak. Berlin, 1873, p. 551. 3 Pallas, Mise. Zool. p. 80 (1766) ; ex Klein. 490 RODENTIA distributed throughout South America, extending even to the Straits of Magellan. The Restless Cavy (C. porcellus), which is found throughout Uruguay and Brazil, has been very generally regarded as the ancestral form of the domesticated Guinea-Pig. It is about 10 inches long, and weighs a little over a pound; its fur is long and of a nearly uniform grayish-brown colour. This species is rarely found in dry sandy localities, preferring marshes covered with aquatic plants, among which it lies concealed, feeding in the early morning and after sunset in the evening; but when the soil is dry it forms burrows. It lives in societies of from six to eighteen individuals, breeding but once a year, with one, or at most only two, young at a birth. The Guinea-Pig (probably a misnomer of Guiana- Pig) is larger than C. porcellus, and is regarded by Dr. Nehring as descended from another species, C. cuéleri. It is white in colour, with irregular patches of reddish-brown and black. The Bolivian Cavy (C. boliviensis), found throughout the higher regions of Bolivia, usually at an elevation of 10,000 or 12,000 feet, is exceedingly shy, and lives in burrows, which in some districts are so numerous as to have completely undermined the soil. The Rock-Cavy (C. rupestris), distinguished by its short, blunt nails, is found in rocky situations throughout Brazil, and is much sought after for its flesh. The Southern Cavy (C. australis), common along the coast of Pata- gonia, forms deep burrows, with several outlets, in sandy declivities. Remains of existing species of Cavia are found in the cavern- deposits of Lagoa Santa, Brazil. Dolichotis.A—Characterised by the great length of the ears and the short tail. The palate is so much contracted in front that the premolars of opposite sides touch by their antero-internal edges. Vertebre: C7, D12, L8, 8 3, C10. The Patagonian Cavy (D. patuchonica)—the only living repre- sentative of the genus—is rather larger than a Hare, which it somewhat resembles in external appearance. It inhabits the dry sterile districts of Patagonia and La Plata, disappearing wherever the country becomes more humid. This animal burrows in the earth, although in districts where the Viscacha is found it is said to avail itself of the works of the latter. Unlike other cavies, its eyes are protected from the glare of the sun by prominent eyelashes. The body is covered with a long dense fur of a rusty colour. Two young are produced at a birth. Three species of Dolichotis have been described from the Brazilian cave-deposits, one of which is probably not really separable from the existing form. Hydrocherus.-—A. large aquatic form with all the feet fully webbed ; the skull (Fig. 213, p. 481) large, with enormous par- occipital processes ; and the molars very complex, the third upper 1 Desmarest, Mammalogie, p. 360 (1822). ° Erxleben, Syst. Reg. Animal, p. 191 (1777) ; ex Brisson. LAGOM VIDA 491 one having some twelve transverse lamine. Upper incisors grooved. Vertebre: C7, D 14, L6, 83, C8. The Capybara (H. capybara) is the largest existing Rodent, and the only living representative of the genus. It is a bulky and stoutly built animal, and attains a length of about 4 feet. The body is covered with long and coarse hair, reddish-brown above and brownish- yellow beneath. Capybaras are found over the whole of the eastern part of South America, and to the westward range into Bolivia and Peru. They frequent the borders of rivers and lakes, concealing themselves among reeds and other water plants. Remains of Hydrocherus are found in the cavern-deposits of Brazil, which are probably referable to the existing species ; one extinct species from the Pleistocene of Buenos Ayres is estimated to have attained a length of 5 feet, while H. magnus of the same deposits was of still larger dimensions. The genus is also represented in the Pleistocene of South Carolina and the infra-Pampean beds of Parana. Extinct Genera—A number of South American fossil Rodents have been referred to extinct genera of Caviide. Thus Plexocherus, fromthe Tertiary of Argentina, differs from Hydrocherus in having only nine lamine in the last upper molar ; Cardiomys, Cardiatheriwm, ete., from the infra-Pampeans are also stated to be allied to Hydrocherus, while Contracavia, of the same deposits, is related to Cavia, but of larger size. Microcavia, again, from the Pleistocene of Argentina, is regarded as connecting Cavia with Dolichotis. The Tertiary European genera Issiodoromys and Nesocerodon are apparently referable to the present family. Suborder DUPLICIDENTATA. . Two pairs of incisors in the upper jaw (the second very small, and placed directly behind the large first pair), the enamel of which extends round to their posterior surfaces. At birth there are three pairs of these incisors, but the outer one on each side is soon lost. Incisive foramina large and usually confiuent ; bony palate very narrow from before backwards ; no true alisphenoid canal ; fibula ankylosed to the tibia, and articulating with the calcaneum. Testes permanently external. This suborder includes the Picas, Hares, and Rabbits, all of which are strictly terrestrial. Family LAGOMYIDA. Complete clavicles, subequal limbs, no external tail, and short ears. Skull depressed, frontals contracted and without postorbital processes ; p+ or 3; molars rootless, with transverse enamel-folds. Palearctic and Nearctic. Lagomys.i—Represented by about a dozen species of small 1 Guvier, Tabl. Elément. de U Hist. Nat. p. 182 (1798). 492 RODENTIA Guinea-Pig-like animals, inhabiting chiefly the mountainous parts of Northern Asia (from 11,000 to 14,000 feet), one species only being known from South-East Europe, and another from the Rocky Mountains. The Picas, or Tailless Hares, live in holes among the rocks of their native mountains, and are agile and shy little creatures. The genus is well represented through the upper and middle Tertiaries. It has been proposed to separate those fossil forms with p 2 as Myolagus, and those with p 3 as Titanomys, but this seems scarcely advisable. Family LEPORIDZ. Imperfect clavicles, elongated hind limbs, short recurved tail, and long ears. Skull (Fig. 216) com- pressed, frontals with large wing- shaped post-orbital processes p23; molars as in the Lagomyide. Cosmopolitan (ex- cept Australasia). Vertebre: C 7, D 12,L 7,5 4, C 13- 15. Lepus. — The single genus Lepus includes about twenty species, all of which resemble one another in general external characters. In all the fore limbs have five and the hind only four digits, and the soles of the feet are densely clothed with hairs similar to those covering the legs; the inner surface of the cheeks is also hairy. Although the family has such a wide distribution, the greater number of the species are restricted to the Palzarctic and Nearctie regions, only a single species (L. brasiliensis) extending into South America, where it has existed since the date of the Pleistocene deposits of the Brazilian -caves. The Common Hare (L. timidus*) may be taken as a typical example of the genus, and is characterised by the great length of 1 Linn. Syst, Nat. 12th ed. vol. i. p. 77 (1766). ? From the absence of the Common Hare in Scandinavia it is considered probable that the name Z, timidus was really applied to the Mountain Hare, and some writers accordingly use the name ZL. ewropeus for the former. Fic, 216,—Skull of Hare (Lepus timidus). LEPORIDA 493 the ears and hind limbs. It is found in all parts of Europe except the north of Russia, the Scandinavian peninsula, and Ireland. Its fur is usually of a tawny gray colour above and white beneath, with the upper surface of the short tail and the tips of the ears black. The col- our of the fur differs, however, considerably in different lati tudes and at dif- ferent seasons of the year; show- ing a tendency Fic, 217.—The Common Hare (Lepus timidus). to become white ~ during winter in northern countries, while assuming a reddish- yellow hue in the more genial climate of southern Europe. The Hare isa nocturnal animal, remaining during the day on its “ form,” as the slight depression is called which it makes i in the open field, usually among grass. The Mountain Hare (L. variabilis) is found throughout the northern part of the Palearctic region, ranging from Ireland in the west to Japan in the east, and also occurring in several of the more southerly mountain ranges, such as the Pyrenees, the Alps, and the Caucasus. It is a Lo ees E ~ smaller than the WIGS eee eS common species, Fic. 218.—The Mountain Hare (Lepus variabilis). with a smaller and morerounded head, and shorter ears, tail, and hind limbs. In cold climates the colour of the whole animal changes in the winter to a pure white 494 RODENTIA (as in Fig. 218), with the exception of the tips of the ears, which remain black. In Ireland no winter change of colour takes place. The Rabbit (L. cuniculus), speaking of the wild race only, is distinguished from the Hare externally by its smaller size, shorter ears and feet, the absence or reduction of the black patch at the tip of the ears so characteristic of the Hare, and by its grayer colour. The skull is smaller and lighter, with a slenderer muzzle and a longer and narrower palate. Besides these characters, how- ever, the Rabbit is sharply separated from the Hare by the fact that it brings forth its young naked, blind, and helpless; to compensate Fic, 219.—The Rabbit (Lepus cuniculus). for this, it digs a deep burrow in the earth in which they are born and reared, while the young of the Hare are born fully clothed with fur, and able to take care of themselves in the “form” in which they are born. The weight of the Rabbit is from 24 to 3 Ibs., although individuals perfectly wild have been recorded up to more than 5 lbs. Its general habits are too well known to need a detailed description here. It breeds from four to eight times a year, bringing forth each time from three to eight young. Its period of gestation is about thirty days, and it begins to breed when six months old. Tt attains to an age of about seven or eight years. The geographical distribution of the Rabbit presents many most interesting peculiarities. It is believed to be originally a native of the western half of the Mediterranean basin only, and still abounds in Spain, Sardinia, Southern Italy, Sicily, Greece, Tunis, and Algeria ; and many of the Islands adjoining these countries are quite overrun LEPORID.E 495 with it. Thence it has spread, partly by man’s agency, northwards throughout temperate Western Europe, increasing rapidly wherever it gains a footing: and this extension is still going on, as is shown by the case of Scotland, in which sixty years ago Rabbits were little known, while they are now found in all suitable localities up to the extreme north. It has also gained admittance into Ireland, and now abounds there as much as in England. Out of Europe the same extension of range has been going on. In New Zealand and Australia Rabbits, introduced either for profit or sport, have increased to such an extent as to form one of the most serious pests that the farmers have to contend against, as the climate and soil seem to suit them perfectly, and their natural enemies are too few and too lowly organised to keep their numbers within reasonable hounds. In other cases Rabbits introduced into islands have become or remained more or less distinct from their parent stock; thus the Rabbits both of the Falkland Islands and of Jamaica still show traces of their descent from domesticated varieties, and have never reverted to the ordinary brownish-gray type. And again, as was pointed out by Mr. Darwin,! the Rabbits in the island of Porto Santo, near Maderia, whose ancestors were introduced from Spain in 1418 or 1419, have formed quite a distinct diminutive race, barely half the bulk or weight of English Rabbits, and differing in certain slight details of colour and habits. Bibliography of arene R. Waterhouse. “ Observations of the Rodentia,” Mag. Nat. Hist. iii. (1839); alan, Nat. Hist, viii. and x. (1839-42): Id. **On the Geographical ce. of the Rodentia,” Proc. Zool. See. 1839. pp. 162-174; Id. Natural History of the Maminalia, vol. ii. “ Rodentia” (1848) : Gervais, Die. Univ. @ Hist. Nat. xi. p. 202 (1848); Brandt, ‘‘ Untersuchungen iiber die craniologischen Entwickelungsstufen und Classification der Nager der Jetzwelt,” Mém. de UaAead. Impér. de St. Fetersbourg (1855): Lilljeborg, Sustematisk Erversight af de Gnagnde Digadjuren, Upsala, 18606; Alston, ‘‘On the Classification of the Order Glires.”” Proe. Zool. Sov. 1876, pp. 61-88 : Trouessart, Catal. de Rongeurs, Vivants et Fossiles,” Bullet. Sov. d'Etudes Scient. @ Angers, 1880-81; Cones and Allen, ‘* Monographs of North American Rodentia,” Caited States Geol. Surv. of Territories, vol. xi. (1817); Winge. ‘ Rodentia pa Lagos Santa. Brazil.” Was. Lund. vol. iii. (S87, ; various papers by Peters in Vonretster. Ak. Berlin, and by Alston, Anderson, Blanford, Dobson, Milne-Edwards. Thomas. and cthers, in Proce. Zool. Soc. Journ. Asia’, Soc. Beng., Ann. Mav. Nat. Hist., ete. 1 Variations of Aninals and Plants, 2d ed. vol. i. p. 119. CHAPTER AI THE ORDER CARNIVORA THOUGH the existing Carnivora as at present restricted! form a very natural and well-defined order among the Mammalia, it is difficult to find any important common diagnostic characters by which they can be absolutely separated ; so that, as in the case of so many other natural groups, it is by the possession of a combina- tion of various characters that they must be distinguished. Thus they are all unguiculate, and never have less than four well-developed toes on each foot, with nails more or less pointed, rarely rudimentary or absent. The pollex and hallux are never opposable to the other digits. They are regularly diphyodont and heterodont, and their teeth are always rooted.2_ Their dentition consists of small pointed incisors, usually three in number, on either side of each jaw, of which the first is always the smallest and the third the largest, the difference being most marked in the upper jaw; strong conical, pointed, recurved canines; cheek-teeth variable, but generally, especially in the anterior part of the series, more or less compressed, pointed, and trenchant; if the crowns are flat and tuberculated they are never complex or divided into lobes by deep inflexions of enamel. The condyle of the lower jaw is a transversely placed half - cylinder working in a deep glenoid fossa of corresponding form. The brain varies much in/relative size and form, but the hemispheres are never destitute of well-marked convolutions (Fig. 23,p. 71). The stomach (Fig. 234) is always simple and pyriform. The cecum is either absent or short and simple (Fig. 235), and the colon is not sacculated, or greatly wider than the small intestine. Vesicule seminales are never present. Cowper’s glands are present 1 The Fere of Linneus included all the then known species of the modern orders Carnivora, Insectivora, and Marsupialia. * The tusks of the Walrus, altogether so aberrant in its dentition, are partial exceptions to this statement, but in old individuals the pulp-cavity fills up, and they cease to grow. fi CARNIVORA VERA 497 in some, absent in other groups. The uterus is bicornuate. The mamme are abdominal, and very variable in number. The placenta is deciduate, and almost always zonary. The clavicle is often entirely absent, and when present is never complete. The humerus often has an entepicondylar foramen. The radius and ulna are distinct. The scaphoid and lunar bones are united into one, and there is never a distinct os centrale in the adult. The fibula is always a distinct slender bone. Several of these characters are, however, not applicable to all the members of the extinct group of Carnivores for which the name Creodonta has been proposed, as will be noticed in the sequel. The large majority of the species composing this order subsist chiefly upon some variety of animal food, though many are omnivorous, and some few chiefly, though not entirely, vegetable eaters. The more typical forms live altogether on recently-killed warm-blooded animals, and their whole organisation is thoroughly adapted to a predaceous mode of life. In conformity with this manner of obtaining their subsistence they are generally bold and savage in disposition, though some species are capable of being domesticated, and when placed under favourable circumstances for the development of such qualities exhibit a very high degree of intelligence and fidelity. The existing representatives of the order are naturally divided into two suborders, the members of the one being the more typical, and mainly terrestrial in their mode of life; while those of the other are aberrant, having the whole of their organisation specially modified for living habitually in water. These are called respectively the True, or Fissiped, and the Pinniped Carnivora. Suborder CARNIVORA VERA. Generally adapted for terrestrial progression and mode of life, though some may be partially aquatic in their habits. The fore limbs never have the first digit, or the hind limbs the first and fifth digits, longer than the others. Incisors 3 on each side, with very rare exceptions. Cerebral hemispheres more or less elongated ; always with three or four gyri on the outer surface forming arches above each other, the lowest surrounding the Sylvian fissure. The molar series of teeth have not the uniform characters of those of the Pinnipedia. There is always one tooth in each jaw which is specially modified, and to which the name of “sectorial” or “carnassial” tooth has been applied. The teeth in front of this are more or less sharp pointed and compressed ; while those behind it are broad and tuberculated. The characters of the carnassial teeth deserve special attention, as, though fundamentally the same 32 498 CARNIVORA throughout the suborder, they are greatly modified in different genera. The upper carnassial is the most posterior of the teeth which have predecessors, and is therefore reckoned as the last premolar (p 4 of the typical dentition). It consists essentially of a more or less compressed blade supported on two roots and an inner tubercle supported by a distinct root (see Fig. 220). The blade when fully developed has three cusps or lobes (1, 2, and 3), but the anterior is always small, and often absent. The middle lobe is conical, high, and pointed; the posterior lobe has a compressed straight knife-like edge. The inner tubercle (4) varies very much Fic. 220.—Left upper carnassial teeth of Carnivora. I, Felis; II, Canis; III, Ursus. 1, Anterior, 2, middle (paracone), and 3, posterior (metacone) cusp of blade; 4, inner tubercle (protocone) supported on distinct root; 5, inner cusp posterior in position, and without distinct root, characteristic of the Urside. in extent, but is generally placed near the anterior end of the blade, though sometimes it is median in position. In the Urside alone both the inner tubercle and root are wanting, and there is often a small internal and posterior cusp (5) without root. In this aberrant family also the carnassial is relatively to the other teeth much smaller than in the rest of the Carnivora. The lower carnassial (see Fig. 221) is the most anterior of the teeth without predecessors in the milk-series; it is therefore reckoned the first true molar (m 1). It has two roots supporting a crown, consisting when fully developed of a compressed bilobed blade (1 and 2), a heel, or talon (4), and an inner cusp (3). The lobes of the blade, of which the hinder (2) is the larger, are separated by a notch, generally prolonged into a linear fissure. In the most. specialised Carnivora, as the Felide (I), the blade alone is developed, both talon and inner cusp being absent or rudimentary. In others, as CARMTIORA TERA 499 Mites Vo and Ursus (V1), the heel is greatly developed, broad, and tuberculated. The blade im these cases is generally placed obliquely, irs flat or convex (outer) side looking forwards, so that the two lobes are almost side by side. instead of anterior and posterior. The inner cusp (3) is generally conical, pointed, and placed to the inner side of the hinder lobe of the blade. The special characters of these teeth are more disguised in the Sea Otter (Zafar) than in any other form, but even in it they can be traced. The homology of the various parts of the Carnivorous carnassial I, Felis ; u, Can said) of blade ; 2. po a iin (aypocenid) Ir will be seen that the to the development of the partiva of the crown Fre. 221.—Le® lower carmassis! teeth ef Car IV, Dutra: V, Meles: VI. Crsesx 2. Att lobe lobe of blade + 3, inner c= relative sie of the two roots varies sented chey Lave respectively to saprest ih E. I F uy az with the primitive titubercular type (p. 30) is indicated in the feures. It may be observed, however, that the anterior lobe of the three-lobed upper carnassial is an element added on to the more primitive two-lobed type. When the talon of the lower carnassial, as In Canis, consists of a large outer and small inner cusp, the latter imot seen in the figure) is the entoconid. The toes are nearly always armed with large, strong, curved. and telerabiy sharp claws, ensheathing the ungual phalanges, and held more firmly in their places bv broad lamine cf bone reflected over their attached ends trom the bases of the phalances. In some forms, most notably the Felidae, these claws are retractile ; that is to 500 CARNIVORA say, the ungual phalanx, with the claw attached, folds back in the fore foot into a sheath by the outer or ulnar side of the middle phalanx of the digit, being retained in this position when the animal is at rest by a strong elastic ligament. In the hind foot the ungual phalanx is retracted on to the top, and not the side of the middle phalanx. By the action of the deep flexor muscles, the ungual phalanges are straightened out, the claws protruded from their sheath, and the soft “velvety” paw becomes suddenly con- verted into a most formidable weapon of offence. The habitual retraction of the claws preserves their points from wear in ordinary progression. The skeleton of the Lion represented in Fig. 15 (p. 45) illus- trates the digitigrade mode of progression of the Fvlide, as well as the essential characters of the bony framework of a typical Carnivore. The Fissipedal Carnivora were divided by Cuvier into two groups, according to the position of the feet in walking,—the Plantigrada, or those that place the whole of the soles to the ground, and the Digitigrada, or those that walk only on the toes ; and the difference between these groups was considered of equal importance to that which separated the Pinnigrada or Seals from both of them. The distinction is, however, quite an artificial one, since every intermediate condition exists between the extreme typical plantigrade gait of the Bears and the truly digitigrade walk of the Cats and Dogs; in fact, the greater number of the Carnivora belong to neither one form nor the other, but may be called “subplantigrade”; often when at rest applying the whole of the sole to the ground, but keeping the heel raised to a greater or less extent when walking. An amended classification of the existing forms is into three distinct sections, of which the Cats, the Dogs, and the Bears may be respectively taken as representatives, and which are hence called Hluroidea, Cynoidea, and , Carnivora, 15 ype. (rom Mi much larger than in the Cat; it is characterised by the presence of some strongly marked internal folds near the pyloric extremity, which stop sud- denly at a point where the stomach makes an abrupt constriction and flexure. Beyond this point there are three other longitudinal folds; and the prloric valve is small. The allied genera present modifications from this form of stomach. The cecum (Fig. 235) is short, thick, and yointed. The liver :Fig. 236) uch resembles that of the Cat, but differs in that the lett lateral lobe is undivided, al- though having a small groove on its posterior or abdominal aspect, while the cystic fissure is less deep, and situated more to the right. The caudate lobe is relatively lonzer. has a deep Fic. eee images (After Mivart, coneavity, and runs uninter- i Seka, ruptedly into the Spigelian; the latter being relatively somewhat larger than in the Cai, with a deep groove dividing the proximal third from the distal twothirds. In Virerra the rizht lateral and right central lobes are nearly equal in size. The variations in the form ot the liver 54 ex bend where th: internal ° wart. Proc. Zool. S22. 1552, p. 36.) BD) | 530 CARNIVORA of the allied genera are detailed in Professor Mivart’s memoir. The brain of the Genet is shown in Fig. 23 (p. 71); the small depression d placed on the superior lateral gyrus appears to be the sole representative of the distinct crucial sulcus which dis- Fic. 236,—Abdominal aspect of the liver of the Genet. c, Caudal lobe ;. gb, gall-bladder ; ha, hepatic artery ; hd, hepatic duct ; LC, left central lobe; LL, left lateral lobe ; pv, portal vein ; RC, right central lobe; RL, right lateral lobe; Sp, Spigelian lobe; ve, vena cava, (From Mivart, Proc. Zool. Soc. 1882, p. 510.) tinguishes the brains of the Felide from those of all other members of the ALluroidea. Prionodon.1—This and the following genus comprise the beauti- ful Linsangs (Fig. 238), which are dis- tinguished from the preceding genera by the loss of the second upper molar, which is, however, very small in some of the Genets. In the present genus the ground colour is whitish or yellowish with brown or black markings, which may either form broad continuous patches across the hinder part of the body, or may be broken up into spots. The tail is very long, the limbs comparatively short, and the fur very short and close. The pollex and hallux are well developed ; the claws are almost completely retractile ; and the tarsus and metatarsus are com- Fia. ee cutann of Prionodon. pletely haired. The pupil is round. The (From Mivart, Proc. Zool. Soc, 1882, ‘ ‘ p. 508.) cecum (Fig. 237) is remarkably small. This genus is exclusively Oriental, and comprises ”. gracilis from Borneo, Java, and (?) Sumatra, P. pardi- ' Horsfield, Zool. Research, Java (1824).—Prionodontide. VIVERRIDA 531 color from Nipal, and P. maculosus from Tenasserim ; the head and body of the latter measuring from 18 to 20 inches in length. Speaking of P. pardicolor, Mr. Hodgson observes that it is “equally at home on trees-and on the ground; it dwells and breeds in the hollows of decayed trees. It is not gregarious at all, and preys chiefly upon small birds, which it is wont to pounce upon from the Fia. 238.—The African Linsang (Poiana potnsis). From Mivart, Proc. Zool. Soc. 1882, p. 160. cover of the grass. The times of breeding are said to be February and August, and the litter to consist of two young, there being urp litters each year.” Poiana.1—This African genus, represented solely by one species, P. potnsis (Fig. 238), from Fernando Po, is very closely allied to the preceding, but the spots are smaller, and show no tendency to run into transverse bands or stripes, except in the region of the head and shoulder; while the sole of the foot has a narrow bald band running up towards the tarsus, as in Genetta. The length 1 Gray, Proc. Zool. Soc. 1864, p. 520. 532 CARNIVORA of the head and body is 38 inches, and that of the tail about 40 inches. It is probable that this animal should really be regarded as a slightly aberrant species of the genus Prionodon. The five following genera differ in several important respects from all the preceding, and collectively constitute the Paradoxurine section of Professor Mivart. With the exception of one African form, they are mainly Oriental. In this section the auditory bulla is frequently in two portions, the posterior moiety in one case being unossified, and it is always much narrowed in front (Fig. 239). The palate (as in the figure) may be much produced behind the molars ; and the teeth are often but slightly sectorial, and may be very small. The long tail is in most cases not ringed. Paradoxwrus1—Dentition: 1 3, ¢4, p 4, m 2; total 40. The blunt and rounded form of the cusps of the hinder premolar and the molar teeth distinguishes this genus from most of the members of the family. Vertebre: C 7, D 13, L 7, 8 3, C 29-36. Head pointed in front. Ears small, rounded. Body long. Limbs moderate. Palms and soles almost entirely naked, and joining the foot-pads without the intervention of any hairy space. Claws com- pletely retractile. Pupil vertical. Tail long, non-prehensile ; in the Indian species without rings. The Paradoxures or Palm-Civets are less strictly carnivorous than the other members of the family. They are mostly about the size of the common Cat, or rather larger, and are partly arboreal in their habits. The species are rather numerous, and present considerable variations in the details of the form and size of their molar teeth ; in only a few does the bony palate extend behind the molars. They are restricted geographic- ally to Southern Asia and the Indo-Malayan archipelago. The best known species? are P. niger, P. hermaphroditus, P. jerdoni, P. aureus, P. grayt from India and Burma, P. philippinensis of the Philip- pines, P. larvatus of Southern China and Formosa, P. leucomystax of the Malay Peninsula, Sumatra, and Borneo, and P. musschenbroeki of Celebes. The name Paradoxurus was applied from the mistaken notion that the tail was prehensile. Mr. Blanford® gives the following account of the habits of P. niger: “The common Palm- Civet, Tree-Cat, or Toddy-Cat, is a familiar animal in most parts of India, though, being thoroughly nocturnal in its habits, it is but rarely seen in the daytime. It is arboreal, passing the day gener- ally in trees, either coiled up in the branches, or in a hole in the trunk, and in places where cocoa-nut palms are common it frequently selects one of them for a residence. Mango groves are also a favourite resort. It not unfrequently takes up its abode in the thatched roofs of houses ; Jerdon found a large colony 1 F. Cuvier, Hist. Nat. des Mammiferes, No. 186 (1821). * See W. T. Blanford, Proc. Zool. Sov. 1885, p. 780. * Fauna of British India, ‘‘ Mammalia,” p. 108 (1888). VIVERRID.E 533 established in the rafters of his own house in Tellicheri. It even occurs in large towns; I have known of one being caught in the middle of Calcutta.” ; -fretogale4—This genus—represented only by 4. trivirgata of Java, and 4. leucotis of Burma, Tenasserim, Sumatra, Java, etc.— is chiefly distinguished from Paradorurus by the extremely small size of the cheek-teeth (Fig. 239), which are often not in contact with one another; the upper carnassial being almost triangular in shape. Palate fre- quently convex longi- tudinally between the carnassials, and greatly produced behind the last molar, with a very narrow bony aperture of the posterior nares. The soles of the feet are still more naked than in Paradoxurus ; and the pollex and hallux are more diverg- ent. In A. leucotis the length of the head and body is 26°5 inches, and the tail 27 inches. In many specimens the three dorsal stripes are much less distinctly marked than in others, and tend to break up into spots; while the general coloration is Fic. 239.—Palatal aspect of the left side of the cranium considerably lighter. a eg se leucotis. a, ss opening i . 2 alisphenoid canal; 0, foramen ovale; ¢, carotid canal }. ie rE (From Mivart, Proc. Zool. Soc. 188, p. 165.) modifica 10n 0. e€ Paradoxure type, contains one species, H. herdwickci, from Borneo and Malacca, an elegant-looking animal, smaller and more slender than the Paradoxures, of light gray colour, with transverse broad dark bands across the back and loins; the proximal portion of the tail being ringed. The tarsus is hairy. The general cranial 1 Gray, Proc. Zool. Soc. 1864, p. 542, ex Petero. ? Jourdan, Comptes Rendus, vol. v. p. 442 (1837). Amended. 534 CARNIVORA characters are those of Paradoxurus, but the auditory bulla is ankylosed into a single piece. lretictis\—Dentition: 73, ¢4, p+, m2; total 40. The pos- terior upper molar and the first lower premolar very often absent. Cheek-teeth generally small and rounded, with a distinct interval between them, but formed generally on the same pattern as Paradoxurus. Vertebre: C 7, D 14, L 5, 8 3, C 34. Body elongated. Head broad behind, with a small pointed face. Whiskers long and numerous. Ears small, rounded, but clothed with a pencil of long hairs. Eyes small. Limbs short. Soles and palms broad, entirely naked. Tail very long and prehensile ; thickly covered with long hair. Fur long and harsh. Cecum extremely small. But one species is known, 4. binturong, the Binturong, an inhabitant of Southern Asia from Nipal through the Malay Peninsula to the islands of Sumatra and Java. Although structurally agreeing closely with the Paradoxures, its tufted ears, long, coarse, and dark hair, and prehensile tail give it a very different external appearance. It may be regarded as a very aberrant Paradoxure, connected, so far as dental characters are concerned, with Paradoxurus by means of Arctogale. The bony palate also extends considerably behind the last molar, as in the latter. The Binturong is slow and cautious in its movements, chiefly if not entirely arboreal, and appears to feed on vegetable as well as animal substances, Nandinia? contains one species, .V. binetata, a somewhat aberrant Paradoxure, from West Africa. It is rather smaller than the true Paradoxures, with smaller and more pointed molar teeth, and no cecum. The wall of the hinder chamber of the auditory bulla remains through life unossified. The dentition appears to be of a more decidedly carnivorous type than in the other members of the section. Cynogale.2—This remarkable genus is regarded by Professor Mivart as representing a third section of the Virerrine , it contains one species, C. bennetti (described by S. Miiller under the name of Potamophilus barbatus), from Borneo, Sumatra, and the Malay Peninsula. This is a curious Otter-like modification of the Viverrine type, having semi-aquatic habits, both swimming in the water and climbing trees, living upon fish, crustacea, small mammals, birds, and fruit. The number and general arrangement of its teeth are as in Paradoxurus, but the premolars are peculiarly elongated, compressed, pointed and recurved, somewhat as in the Seals, though the molars are tuberculated. The head is elongated, 1 Temminck, Prospectus de Monographies des ammiferes, March 1824; Monographies, vol. i. p. xxi. (1827). * Gray, List of Mamm. Brit. Mus. p. 54 (1843). 5 Gray, Proc. Zool. Soc. 1836, p. 88. VIVERRIDE 535 the muzzle broad and depressed. Whiskers very long and abundant. Ears small and rounded. Toes short and slightly webbed at the base. Tail short, cylindrical, covered with short hair. Fur very dense and soft, of a dark brown colour, mixed with black and gray. Humerus without entepicondylar foramen. Subfamily Herpestine.—Auditory bulla very prominent, and somewhat pear-shaped, the posterior chamber being large, rounded, and generally with its greatest prominence to the outer side. The anterior chamber considerably dilated, and produced into a short inferior wall to the auditory meatus, in which is a depression or vacuity just below the centre of the opening of the meatus. Sometimes this vacuity is continued into the meatus, forming a narrow fissure. The paroccipital process does not project beyond the bulla, but is spread out and lost (in adult animals) on its posterior surface. Toes straight; claws lengthened, exserted, non-retractile. No perineal glands. The dentition is always of a markedly sectorial type; and the orbit may be surrounded by bone. Very generally the anus opens into a sac-like depression. The majority of the genera are Ethiopian; the type genus alone extending into the Oriental and Palearctic regions. Herpestes.\— Dentition : i 3, ¢ 4, p 4, sometimes 3, m 2; total 40 or 36. Teeth of molar series generally with strongly developed, sharply-pointed cusps. Skull elongated, constricted behind the orbits. Face short and compressed. Frontal region broad and arched. Postorbital processes of frontal and jugal bones well developed, generally meeting so as to complete the circle of the orbit behind. Vertebre: C 7, D 13, L 7, 8 3, C 21-26. Head pointed in front. Ears short and rounded. Body very long and slender. Extremities short. Five toes on each foot, the first, especially that on the hind foot, very short. Toes free, or but slightly palmated. Palms generally naked. Distal portion of soles naked, under surface of tarsus and metatarsus usually clothed with hair, but considerable specific variation in this respect. Tail long or moderate, generally thick at the base, and sometimes covered with more or less elongated hair. The longer hairs covering the body and tail almost always annulated. This genus contains a very large number of animals commonly called Ichneumons, or in India Mungooses, varying in size from that of a large Cat down to a Weasel. They are widely distributed over the African continent and the southern parts of Asia, especially India and the Indo-Malayan archipelago, one species occurring also in Spain. They are mostly terrestrial in their habits, feeding on small mammals and birds, reptiles, especially snakes, eggs of birds and reptiles, and also insects. Some species are partially domesticated, being used to keep houses clear of rats, mice, and 1 Tlliger, Prodromus Syst. Mamm. p. 185 (1811). 536 CARNIVORA snakes. H. ichnewmon was a sacred animal to the ancient Egyptians. They vary considerably in appearance, some, as J. gulera and H. urva (Fig. 240), are larger and heavier, with stouter body, longer limbs, and stronger teeth. The common Indian Mungoose (H. mwngo) is considerably smaller than the Egyptian form; its fur is of a pale gray colour, the hairs being largely white ringed, while the cheeks and throat are more or less reddish. Like the Egyptian species, it is frequently domesticated, and put to a similar use. It is especially serviceable in India as a serpent- killer, destroying not only the eggs and young of these creatures, but attacking without hesitation and killing the most venomous Fia, 240.—The Crab-eating Mungoose (Herpestes urva). From Blanford, Mammalia of British India, p. 180. adult snakes. The fact that it invariably survives those en- counters has led to the belief that it either enjoys immunity from the effects of snake-poison, or that after being bitten it has recourse, as the natives maintain, to the root of a plant as an antidote. Neither of these suppositions has stood the test of scientific examination, for it has been found that when actually bitten it falls a victim to the poison as rapidly as other mammals, while there is no trustworthy evidence of its seeking a vegetable antidote. The truth seems to be that the Mungoose, by its exceeding agility and quickness of eye, avoids the fangs of the snake while fixing its own teeth in the back of the reptile’s neck. One large species, believed to be from Africa, recently described as H. grandis, is remarkable for the extreme complexity of the cusps on the molars, and also for the absence of an entepicondylar foramen to the humerus; the latter feature also occurring in the allied H. albicuudatus. The Oriental H. wrra (Fig. 246) is stated to be somewhat aquatic in habits, and to feed on frogs and crabs. VIVERRIDA 537 Remains of the small H. nipalensis occur in the cavern-deposits of Madras. Viverroids from the Miocene and Upper Eocene of Europe, which agree with Herpestes in the presence of an inner tubercle to the third upper premolar and of a hinder cusp to the fourth lower premolar, have been referred to the existing genus. The species which have been separated generically under the three following names are very closely allied to Herpestes. Helogale, premolars 3, without diastema between first and second; soles of feet completely naked. Contains two small South-African species, H. parvula and H. undulata. Bdeogale? contains also two small Ichneumon-like animals, B. crassicauda and puisa, differing from Herpestes proper in having only four toes on each foot, both pollex and hallux being absent. The orbit is nearly complete, the tail of moderate length and rather bushy. Cynictis.3—Pollex present, but hallux absent. Skull shorter and broader than in Herpestes, rather con- tracted behind the orbits, which are large and complete behind. Face short. An- terior chamber of the auditory bulla very large. Front claws elongated. C. peni- cillata, from South Africa. The cecum (Fig. 241) of this genus is longer than in any other member of the family. All the foregoing Herpestines have the nose short, with its under surface flat, bald, and with a median longitudinal groove. The remaining forms have the nose more or less produced, with its under side convex, and a space between the nostrils and the upper lip covered with close adpressed hairs, and without any . median groove. Fic. 241.— Cxecum 0 Cynictis coma pee 5-5. Claws of fore pg ore aia oo feet short, compressed, acute. Under sur- face of tarsus hairy. Palate flat. Founded on a single specimen from East Africa, 2. melleri. Crossarchus.-—Dentition : ¢ 3, ¢4, p 3, m 3; total 36. Snout elongated. Toes 5-5. Claws on fore feet long and curved. Hallux very short. Under surface of tarsus naked. ‘Tail shorter than the body, tapering. Palate flat. Fur harsh. Species: C. 1 Gray, Proc. Zool. Soc. 1861, p. 308. 2 Peters, Mith. Ges. Nat. Freunde Berlin, 19th November 1850. 3 Ogilby, Proc. Zool. Soc. 18338, p. 48. 4 Gray, Proc. Zool. Soc. 1864, p. 573. 5 FB. Cuvier, Hist. Nat. des Mammiferes, No. 199 (1825). 338 CARNIVORA olscurus, the Kusimanse, a small burrowing animal from West Africa, of uniform dark brown colour; (\ fasciatus ; C. zebra ; and C. gambianus. Suricata..—A more distinct genus than any of the above. The dental formula as in the last, but the teeth of the cheek-series remarkably short in the antero-posterior direction, corresponding with the shortness of the skull generally (Fig. 222). Orbits complete behind. Vertebre: C 7,D 15,L 6,8 3,C 20. Though the head is short and broad, the nose is pointed and rather produced and movable. Ears very short. Body shorter and limbs longer than in Herpestes. Toes 44, the pollex and hallux being absent. Claws on fore feet very long and narrow, arched, pointed, and subequal. Hind feet with much shorter claws, soles hairy. Tail rather shorter than the body. One species only is known, the Suricate, S. fetradactyla, a small gray-brown animal, with dark transverse stripes on the hinder part of the back, from South Africa. The cecum is short. Galidictis? Galidea,? and Hemigalidea* are names of three slight generic modifications of the Viverrine type, allied to the Herpestine, but placed by Mivart in a distinct subfamily, Galidictiin. They are all characterised by the absence of the alisphenoid canal in the skull, as well as of the entepicondylar foramen to the humerus ; and are inhabitants of Mada- gascar. The best known, Gualidea elegans, is a lively Squirrel-like little animal with soft fur and a long bushy tail, which climbs and jumps with agility. It is of a chestnut- brown colour, the tail being annulated with darker brown. The cecum (Fig. 242) is remarkable for its comparative length and pointed termination. Hemigalidea is dis- gee er tinguished by the absence of rings on the Geiahee Ew Wess ene, tail. Galidictis vittata and striata chiefly Zool. Sec, 1882, p. 308.) differ from the Ichneumons in their colora- tion, being gray with parallel longitudinal stripes of dark brown. Eupleres® is another form, also from Madagascar, which has been placed in a subfamily apart. It differs remarkably from all the other Vivcrride in the weak development of the jaws and the ? Desmarest, ‘‘Tabl. Méth. Mamm.” in Nour. Dict. @'Hist. Nat, vol. xxiv. (sod, ° Geoffroy, Comptes Rendus, 1837, p. 578, 3 Geoffroy, Mag. de Zool. 1839, pp. 27, 37. * Doyere, dan. Sei. Nat. vol. iv. p. 281 (1835), ° Jourdan, Comptes Rendus, 1837, p. 422. Amended. PROTELEIDL 539 small size of the teeth (Fig. 243), in consequence of which it was, when first discovered, placed in the order Insectivora. Dentition : 23,c4,p4, m2; total 40. Vertebre: C7, D13,L7,8 3, C 20. No alisphenoid canal ; an entepi- condylar _fora- men to the hum- erus. But one species is known, E. goudoti. Extinct Gen- erd.—The Ter- Fic. 243.—Skull of Eupleres goudoti. + natural size. Mus. Roy. Coll. Surgeons. tiaries of the Old World have yielded several genera allied to the existing Viverroids, some of which show decided signs of affinity with other families. Of these the Lower Miocene 4mphictis appears to be nearly related to Viverra, but is distinguished by the form of the second lower molar, which is longer and has two distinct roots. Paleoprionodon, of the French Phosphorites, has a dentition very like that of Prionodon, the molars being reduced to $; the skull has an alisphenoid canal and the general basal characters of the /iverride, but resembles the Justelide in the presence of a glenoid foramen and in the position of the condylar foramen. In Stensplesictis, of the same deposits, the dental formula is 7 3, ¢ 4, p 4, m2; and although the skull has a complete septum in the bulla, yet some of the cranial and dental features ap- proximate so decidedly towards those of the extinct J/ustelide, as to lead some authorities to refer the genus to that family. The most probable explanation of this resemblance is that the Musteloids have originated from generalised Viverroids allied to Stenoplesictis. The Lower Pliocene Ictitherium differs from all other Viverroids in the presence of three distinct lobes to the upper carnassial, and thereby connects the other members of the family so closely with the Hyenide that it is practically impossible to draw up a defini- tion which will distinguish the two families. The North American Eocene genera Jfiacis and Didymictis are generally regarded as representing a separate family—.lJiacide— with affinities both to the Viverride and Canide. Family PROTELEID-£. Skull with no alisphenoid canal; and the auditory bulla divided into two distinct chambers. Dorsal vertebre 15. Molars+. Pre- molar and molar teeth very small and simple in character. Proteles.\—This genus contains but a single species, P. cristatus, 1 Geoffroy, Wem. du Muséum, vol. xi. p. 354 (1824). 540 CARNIVORA the Aard-Wolf or Earth-Wolf of the Dutch colonists of the Cape, an animal nearly allied to the Hyznas, but remarkably modified in its dentition, the molar teeth being very small, placed far apart, and almost: rudi- mentary in char- acter (Fig. 244). The canines are long and rather slender. The dental formula is i3,c1,pand m a ~, ; total 30 or 32. Vertebree : SS OY, D 15, 15, Fic. 244.—Skull and Dentition of the Aard-Wolf (Proteles cristatus). § gi C 24. The 4 natural size. fore feet with five toes; the pollex though short, with a distinct claw. The hind feet with four subequal toes. Claws all strong, blunt, sub- compressed, and non-retractile. The general external appearance is very like that of a small Striped Hyzna, but the muzzle is more pointed and the ears larger. It has a copious mane of long hair, capable of being erected when the animal is excited, along the middle line of the neck and back. It is a native of South Africa, and is a burrowing nocturnal animal, feeding on decomposing animal substances, larve, and termites. Observations upon speci- mens in captivity indicate that it has neither inclination nor power to attack or feed upon living vertebrated animals. Some writers regard Proteles as representing a subfamily of the Hyenide. Family HY£ZNIDZ. Skull with no alisphenoid canal; and the auditory bulla not divided by a septum into two chambers. Dorsal vertebre 15. Molars usually 4, but in some fossil forms 4, or 2, the second lower molar being very small; upper carnassial with three distinct lobes; lower carnassial with a large blade and small talon. No entepicondylar foramen to the humerus.. This family is confined to the Old World, where it is now represented by a single genus, which, although evidently nearly related to the Viverride, is sufficiently distinct to be regarded as not referable to that family. The extinct Ictitherium, however, as already mentioned, connects the more generalised members of the Hycenide very closely with the Viverride. Hyena?—Dentition in existing forms usually 7 3, ¢ 3, p 4, m 1 For Anatomy of Proteles see Flower, Proc. Zool. Soc. 1869, p. 474. 2 Zimmermann, Specimen Zoologie: Geographice, p. 365 (1777). HVA NIDA 541 zy; total 34. Teeth, especially canines and premolars, very large, strong, and conical. Upper carnassial (Fig. 245) with a very large, distinctly trilobed blade and a moderately developed inner tubercle placed -at the anterior extremity of the blade. Molar very small, and placed transversely close to the hinder edge of the last, as in the Felide, Lower carnassial con- sisting of little more than the bilobed blade. Zygo- matic arches of cranium : very wide and strong. Fic. 245.—Outer (4) and palatal (B) aspects of the right Sagittal crest high, giving SPR coms wth ot te Set Hyena, ens attachment to very power- ful biting muscles. Orbits incomplete behind. Vertebre: C 7, D 15,L5,8 4,C 19. Limbs rather long, especially the anterior pair, digitigrade, four subequal toes on each, with stout non- ‘retractile claws. Pollex and hallux only represented by rudi- mentary metacarpal and metatarsal bones. Tail rather short. A large post-anal median glandular pouch, into which the largely developed anal scent glands pour their secretion. The three existing species of Hyzna are divisible into two sections, to which some zoologists assign generic rank, but fossil forms show such a transition between these two types as to render any such division impracticable. The typical or Huhycenine group presents the following dis- tinctive features. Upper molar moderately developed and three- rooted. An inner cusp and hind talon more or less developed on the lower molar. Ears large, pointed. Hair long, forming a mane on the back and shoulders. H. striata, the Striped Hyzena (Fig. 246) of Northern Africa and Southern Asia. H. brunnea, of South Africa, in some respects intermediate between this and the next group. The Striped Hyzna is dirty gray in colour, with narrow trans- verse tawny or blackish stripes on the body and legs ; the length of the head and body is 34 feet, and that of the tail, with its hair, 14 feet. It occurs throughout peninsular India, where it is most common in open hilly districts, and in North Africa. Mr. Blanford! gives the following account of its habits: “It is a nocturnal animal, and although an occasional individual may be met with returning to its den in the early morning, its rambles are usually commenced after sunset and ended before sunrise. During the night it roams far and wide, and no tracks of wild animals are more common in the countries where it is found than its unmistakable footprints, very like a dog’s 1 Fauna of British India, ‘‘ Mammalia,” p. 183 (1888). 542 CARNIVORA in shape, but with the marks of the hind feet conspicuously smaller than those of the fore feet. Unlike the Spotted Hyzna, the Striped species appears to be solitary in its habits, and it is rare to meet with more than two together. The principal food of the Hyena consists of the carcases of animals that have died of disease or been killed by beasts of prey, and very often it carries off portions of the body to its den. I once shot one that was carrying away the hind leg of a Nilghai. The powerful jaws and large teeth are admirably adapted for crushing bones, which are consumed by Veg: & = RU ENN : SSS RSE RONE Fic. 246.—The Striped Hyzna (Hyena striata). Hyzenas, after the flesh has been picked off by vultures and jackals. Occasionally sheep or goats, and more often dogs, are carried off by Hyenas, and the latter at all events are often taken alive to the animal’s den.” The Striped Hyena is essentially a cowardly animal, and one that is much more silent than H. crocuta. Remains of striata are found in the cavern-deposits of the south of France, and also in the Upper Pliocene of the Val d’Arno in Tuscany, and in the English Red Crag. The Crocutine group presents the following characters. Upper molar extremely small, two- or one-rooted, often deciduous. Lower molar without trace of inner cusp, and with an extremely small talon. Ears moderate, rounded. Hair not elongated to form amane. . crocuta, the Spotted Hyena (Fig. 247), from Africa south of the Sahara. In dental characters as well as in its visceral anatomy, especially as regards the reproductive organs of the female,! this species may be considered as by far the more 1 The anatomical peculiarities of Hywna crocuta have been fully elucidated in HVENIDE 543 specialised form. The Spotted Hyena isa larger and bolder animal than the Striped species, hunting in packs, and uttering very frequently its unearthly cry. The coloration consists of dark brown spots on a yellowish ground. It was formerly very common at the Cape. Remains of a large race of this species are exceedingly common in the cavern-deposits of Europe, where they were first described under the name of Hyena spelea ; teeth have also been met with in the Norfolk Forest-bed, and in cavern-deposits in Madras—the latter locality being exceedingly interesting from a distributional point of view. In addition to the remains of existing species, to which refer- Fia. 247.—The Spotted Hyzna (Hycna crocuta). ence has been already made, there were numerous extinct forms of Hyena in the upper Tertiaries of Europe, from the horizon of the Lower Pliocene Pikermi beds of Greece upwards. In the Crocutine group Z. colvini of the Pliocene of India (Fig. 248), and H. robusta of that of Italy, appear to have been ancestral forms allied to . crocuta ; the former being distinguished by the loss of the first upper pre- molar. H. eximia, of the Pikermi beds, is a more generalised form, in which the first lower premolar (lost in existing forms) is retained. In the typical group, H. arvernensis and H. perrieri, of the Upper Pliocene of the Continent, approximate to H. brunnea, although H. perriert makes a farther step towards the Crocutine group by the loss of the inner cusp in the lower carnassial. The extinct Hyenic- a series of papers by Morrison Watson in the Proceedings of the Zoological Society for 1877, 1878, 1879, and 1881, in which references to previous authors on the subject will be found. 544 CARNIVORA tine group, as represented by the Indian H. sivalensis and the Grecian H. greca, connects H. striata with Palhyena. Both are characterised by the presence of a small second lower molar behind the carnassial ; while H. greca also has four lower premolars. Still more generalised is the Lychyenine group, comprising H. macrostoma of India and H. cheretis of the Pikermi beds; in these forms the muzzle was longer, and the premolars much more compressed than in the existing species, thus making a very decided approach to the Vwerride. There were four lower premolars ; the lower carnassial had an inner cusp, and it is probable that there was a second lower molar ; while the first upper molar was placed partially behind the Fia. 248.—Outer view of part of the right ramus of the mandible of Hyena colvini, showing the third and fourth premolars and the carnassial. (From the Paleontologia Indica.) carnassial. The Lower Pliocene Palhyena hipparionum, in which the dental formula is i 3,¢ 4, p 4, m 2, is a smaller form with long jaws and compressed premolars which approaches so closely to the Viverroid genus Ictitherium as to show pretty clearly how the Hyznas have been gradually modified from that stock. Section CYNOIDEA. Family CANIDa. This section contains the single family of the Canide, or Dog- like animals, which appear to hold an intermediate position between the other two sections, retaining also many of the more generalised characters of the ancient members of the order. The structure of the auditory bulla and adjacent parts of the bones of the skull is intermediate between that of the AZluroid and Arctoid forms. In the number and arrangement of the teeth they more nearly approach the primitive heterodont type than any other existing Carnivora. CANIDA 545 A cecum is always present, sometimes short and simple, but when long it is folded upon itself in a characteristic manner. The characters of the base of the cranium are shown in Fig. 8 air /. we > pes Ke YS > Fic. 249.—Right lateral aspect of the skull of the Dog (Canis fumiliaris). (p. 38), where it will be seen that the auditory bulla is inflated, although it has only a rudimental internal septum ; the paroccipital process, although in contact with the bulla, is prominent, and there is a large glenoid foramen. In all the existing forms the humerus has lost the entepicondylar foramen; the crowns of the upper molars are triangular in shape (Fig. 251), and the blade of the upper carnassial consists of two lobes. In the alimentary canal the cecum (Fig. 250) is extremely characteristic. It is a simple appendage of nearly uniform width (about equal to that of the ileum) attached to the side of the canal, just beyond the ileo-cxecal valve, and with a rounded termination. In a Dog of average size it is 5 or 6 inches long if uncoiled, but it is normally folded by its mesenteric attachments backwards and forwards several times on itself by the side of the ileum, after the manner shown in the figure. The existing Dogs form a very compact group, with numerous species closely resembling each other in essential characters, though differing considerably externally. The most marked differences are slight variations in the number of the true molar teeth, which exceed the usual number in the Cape Long- eared Fox (Otocyon), and fall short of it in some other less aberrant forms to which the names of Icticyon and Cyon have been given, and a diminution in the number of toes in the Cape Hunting Dog (Lycaon), Fic. Cecum of the Arc- tic Fox (Canis lago- 250. — pus). %, Tleum; c, colon. In the nat- ural position the colon is upper- most. which has 4-4, instead of 5-4 as in the remainder of the family. 35 546 CARNIVORA After taking these away, there remain a great number of animals called Dogs, Wolves, Jackals, and Foxes, varying from one another only in the characters of the tail, ears, fur, form of the pupil, and some trifling peculiarities of skull and teeth, upon which some authors have divided them into many genera. ‘These divisions are, however, extremely difficult, if not impossible, to define, on account - of the numerous gradual transitions from one form to the other. Canis.1—It appears on the whole convenient to retain all the species, with the exception of Otocyon, Icticyon, and Lycaon, in the old genus Canis, the most prominent characters of which are the following. Teeth, usually i 3, ¢ 4, p 4, m 3; total 42. The absence of the last upper molar (m_ 3), alone distinguishes this from the generalised dentition of hetero- donts, and this tooth is occasionally present in one species (C. can- crworus). In certain ps pa m1 m2 — Asiatic species (C. pri- Pe annem mma enh ctam erin Wot sucrnus and its allies), which on this account have been separated to form the genus Cyon of Hodgson, the last lower molar (m 3) appears to be constantly absent. The milk- dentition is di 3, de 1, dm 2; total 28,—the first permanent pre- | molar having no predecessor. The teeth of both permanent and milk or temporary series are figured on p. 26, Fig. 3, from the outer aspect, while the woodcut 251 shows the palatal aspect of the hinder upper teeth. The upper carnassial (p 4) consists of a stout blade, of which the anterior lobe is almost obsolete, the middle lobe large, conical, and pointed backwards, and the posterior lobe in the form of a compressed ridge; the inner tubercle is very small, and placed quite at the fore part of the tooth. The first molar is more than half the antero-posterior length of the carnassial, and consider- ably wider than it is long; its crown consists of two prominent, conical cusps, of which the anterior is the larger, and a low broad inward prolongation, supporting two more or less distinct cusps and a raised inner border. The second molar resembles the first in general form, but is considerably smaller. The lower carnassial (m 1) is a very large tooth, with a strong compressed bilobed blade the hinder lobe being considerably the larger and more pointed a small but distinct inner cusp placed at the hinder margin of the posterior lobe of the blade, and a broad, low, tuberculated talon, * Linn. Syst, Nat. 12th ed. vol. i. p. 56 (1766). CANIDE 547 or heel, occupying about one-third of the whole length of the tooth. The second molar is less than half the length of the first, with a pair of cusps placed side by side anteriorly, and a less distinct posterior pair. The third is an extremely small and simple tooth, with a subcircular tuberculated crown and single root. The cranium (Fig. 249) is more or less elongated, the facial portion tapering forwards and compressed. The jaws are elongated, and the zygomata moderately strong. The postorbital processes of the frontal short, leaving the orbit widely open posteriorly. Verte- bre: C 7,D 13, L 7,8 3, C 17-22. Clavicles present, but very rudimentary. Limbs of moderate proportions, digitigrade. Feet short ; five toes on the fore foot, the pollex much shorter than the others, and not reaching to the ground. Four toes on the hind foot, the hallux being represented by a rudiment of the metatarsal.! All the toes are provided with exserted, non-retractile, slightly curved, and blunt claws, which, being exposed, become worn at the tips. Tail moderate, or rather long, generally somewhat bushy. The pupil of the eye, when contracted, is in some species round, in others elliptical and vertical. This extensive genus may be considered as truly cosmopolitan. One or more species occur in every part of the American continent from Greenland to Patagonia and the Falkland Isles; and similarly, in the Old World, Europe, Africa, and Asia, with most of the large islands adjacent, and even Australia, have their wild Dogs, though in the last case they may belong to a feral race, introduced origin- ally by man. They are generally sociable animals, hunting their prey in packs. Many species burrow in the ground; none habitu- ally climb trees. Though mostly carnivorous, feeding chiefly on animals they have chased and killed themselves, many, especially among the smaller species, eat garbage, carrion, insects, and also fruit, berries, and other vegetable substances. The species are very numerous, and, as in most other large genera, very ill-defined, few zoologists agreeing as to which of the many slightly different modifications should be considered as local varieties and which true species. Perhaps the best cranial character by which the different members of the genus can be distinguished is that pointed out by Burmeister, viz. that in the animals generally called Dogs, Wolves, and Jackals the postorbital process of the frontal bone is regularly smooth and convex above, with its extremity bent downwards, whereas in Foxes this process is hollowed above, with its outer margin (particularly of the anterior border) somewhat raised. This modification coincides in the main with that upon which Professor 1 In Domestic Dogs a hallux is frequently developed, though often in a rudi- mentary condition, the phalanges and claw being suspended loosely in the skin, without direct connection with the other bones of the foot ; it is called by dog- fanciers the ‘‘ dew claw.” Re CARI ORA Huxley? has basei bis division 2 atte group into two parallel series, the Thovids or Lupine ¢ and Alopec:ics or Vulpine is which he chamiectersss bY a Presence of rental air-sinuses in the former, which not only aifect the external conten but to a sv creater degree the shape of the anterior part of <2 cranial cavity, and Lichtenstein, Wiegmann’s Archiv. 1838, vol. i, p. 290. CANIDAL 555 of the four molar teeth as a survival of a condition of the dentition exhibited by the common ancestors of the existing Canide and the existing carnivorous Marsupials. There is, however, at present no paleontological proof of this, as none of the numerous fossil forms of Canide yet discovered have more than the normal number of molars. Extinct Genera.—A large number of fossil Carnivora have been described from various Tertiary deposits which are more or less closely allied to the existing Canide, although, as already men- tioned, connecting the latter so closely on the one hand with the Vwerride and on the other hand with the Urside, that it is almost, if not quite impossible to say where one family begins and the other ends. A few only of the more important of these annectant types will be mentioned here. Temnocyon, of the Miocene of the United States, is a true Dog, which agrees with Icticyon in having a secant hind talon to the lower carnassial, but preserves a generalised char- acter in having an entepicondylar foramen to the humerus. An extremely interesting form is Cynodictis, of the Middle Tertiaries of Europe and the United States, which (as now restricted by . Dr. Schlosser) includes a number of species mostly not larger than Foxes. The dental formula is generally the same as in Canis, but (as in that genus) the last lower molar may be absent. The teeth are very like those of the V’iwerride, the lower carnassial never being greatly elongated antero-posteriorly, and its inner cusp being situ- ated immediately on the inner side of the hinder lobe of the blade, instead of somewhat behind it, as is the case in most Dogs. In the skull the auditory bulla is inflated, but is said to have no distinct septum; while the humerus invariably has an entepicondylar foramen. It is suggested that Cynodictis is not far removed from the ancestral type of many of the Viverroids and Canoids, and may itself have been derived from the undermentioned genus Amphicyon. M. Boule considers, indeed, that from the resemblance of the Plio- cene Canis megamastoides (p. 553) to Cynodictis we ought to regard the Foxes and Jackals as the descendants of Cynodictis, while the Wolves have been derived directly from Amphicyon. The last named genus, which includes some species as large as a Bear, is found in the Upper Eocene and Lower Miocene of Europe, and is represented in the Miocene of the United States by the allied Daphenus. It is characterised by the presence of three upper molars—thus bringing up the dental formula to the full Eutherian number ; by the five digits on all the feet, which were plantigrade; and by the presence of a third trochanter to the femur and an entepicondylar foramen to the humerus. The teeth are essentially those of a dog, and the base of the skull is also dog-like, although it is highly probable that the auditory bulla had no trace of a septum. According, however, to Dr. Filhol! the minute foramina 1 Arch. Mus. Lyon. vol. iii. art. 1, p. 85 (1881). sc CARNE LA ti deseribed by Professor Cope® im the poscparictal and mastoid which aeexr in Ursus, but are said to be afcen: in amis, are present in lophicyen. S02 tar, however, as We can sce. the presemee cr atseme of those foramima cannot be regarded as dinemostie of (ws and Cams, although they sre generally more sronely developed im the former. Amphicgen many, Indesd, be sonciered Se 8 75 genemised Dog, with atiniies to the Bears in the closely as to render +: absolutely impossible to indicate any ekaracters af family importance by which they can be distinguished. The upper varmasial of Dinca is tke wn. For other cenera, see >. 562. Seva ARCTOMEA This seetion inchdes a consderable mumber of forms which agree in the essential charactermties of the scrne cures of the base of the cranium and repmadiccive onzans and im the absence of & excum fo the imtesiml eanal They have no Cowpers chnds bat there is a rudimentary prostate and a large cylindrical penial bene : while all the members of the coe) have ive the various forms. but the foJowns features are commun te all. Ths eavity oe _ of the auditory bulla is simple, and has Ist weg fhe GEE seats Radeon Do trace af a dividing septum: the ee inferior lip of the auditory meatus ma oS Gaa, PeS. ERE Ge considerably pO lon: zed : “the pereceipital process (%) a the exvevipital Is more er less triangular, directed backwank, cunwands and downwards. and sanding qtite sparc from the balla: the mastoid process (m) of the periocie is ‘always widely separated from the parvccipital, and generally very prominent : the carotid foramen (cur) is large, and placed on the inner margin ® Pree. tmer. Phil, Sco. val wei, p 452 158 - ORSIDA: 557 of the bulla, usually near the middle, but occasionally more posteriorly ; the condyloid foramen is distinct and exposed, and never sunk into a common opening with the foramen lacerum posticum ; and the glenoid foramen is always present, and usually conspicuous. The alisphenoid canal is absent except in Ursus, Melursus, and Aflurus. It has been already observed (p. 501) that the evidence of fossil forms, so far as it goes, is not in favour of the Arctoidea being a natural group ; so that its retention must be regarded as a some- what provisional measure, largely based on its convenience. The group may be divided into the three families, Urside, Procyonide, and Mustelide.+ Family Urstp&. In existing forms the true molars 4, with broad, flat tuber- culated crowns. Typically the three anterior premolars of both jaws rudimentary and often deciduous. Fourth upper premolar (carnassial) with no third or inner root. An alisphenoid canal (except in Atlwropus). Skull with the auditory bulla depressed, and scarcely at all inflated. Feet plantigrade. No entepicondylar foramen to the humerus. Kidneys conglomerate. Geographical distribution extensive. Ursus.2—Dentition: i 4,c4,p4,m4; total 42. The three anterior premolars above and below one-rooted, rudimentary, and frequently wanting. Usually the first (placed close to the canine) is present, and after a considerable interval the third, which is situated close to the other teeth of the molar series. The second is very rarely present in the adult state. The fourth (upper car- nassial) differs essentially from the corresponding tooth of other Carnivores in wanting the inner tubercle supported by a distinct root. Its sectorial characters are very slightly marked, and it is much smaller than the first molar. The crowns of both the true molars are longer than broad, with flattened, tuberculated, grinding surfaces. The second has a large backward prolongation or heel. The lower carnassial has a small and indistinct blade and greatly developed tubercular heel. The second molar is of about the same length, but with a broader and more flattened tubercular crown. The third is smaller. The milk-teeth are comparatively small, and shed at an early age. Skull more or less elongated. Orbits small and incomplete behind. Palate prolonged considerably behind the last molar tooth. Vertebre: C 7, D 14, L 6, 8 5, C 8-10. Body heavy. Feet broad, completely plantigrade ; the five toes on each foot all well developed, and armed with long compressed and 1 For full details of the Arctoidea see Mivart, Proc. Zool. Soc. 1885, p. 340. 2 Linn. Syst. Nat. 12th ed. vol. i. p. 69 (1766). 558 CARNIVORA moderately curved non-retractile claws. Palms and soles naked. Tail very short. Kars moderate, erect, rounded, hairy. Fur generally long, soft, and shaggy. The Bears are all animals of considerable bulk, and include among them the largest members of the order, Though the species are not numerous, they are widely spread over the carth’s surface (but absent from the Ethiopian and Australian regions, and only represented by one species in the Neotropical region), and differ much among themselves in their food and manner of life. They are. mostly omnivorous or vegetable feeders, and even the Polar HN AVN (nwt in ¥( lta, 255.—Head of the Brown Bear (Ursus actos). From Sclater, Proc. Zool, Soe, 1867, p. S17. Bear, usually purely carnivorous or piscivorous, devours grass with avidity in summer. Tho various species may be arranged in the following groups :— Thalassaretine Group.—lH cad comparatively small, molar teeth small and narrow. Soles more covered with hair than in the others. This group is represented only by the well-known Polar or White Bear (U. maritimus) of the Arctic regions, which is one of the fow mammals which are completely white at all seasons of the year. The typical, or Ursine, group includes a number of species, of which the Common Brown Bear (U. aretos) is the best known example. This species is an exceedingly variable one, and has a very wide range in the Palwaretic region; the Syrian form described as U. syriacus, ws well as the Tairy-cared Bear (UV. piscator, Fig. 255) of North-Hastern Asia, and the Snow-Bear (U/%. isabedlinus) of URSID AL 559 Kashmir and Nipal, not: being spocitically separable, ‘The Brown Bour hibernates in cold regions, and im the Himalaya keeps to comparatively high regions, emerging front its winter lair in March, April, ov May, according to the season and elovation, to feed on the numerous bulbous plants which abound in tho regions it inhabits. Both the Syrian and Himalayan varieties are generally of lighter colour and smaller size than the typieal European form. Bears Were at one time found in the British Isles, from which, how- over, they have boon long since exterminatod, They are still found in the Pyrenees, and are comparatively abundant ino parts of Norway, Hungary, and Russia. In the Kashmir Himalaya they were very abundant in some districts a few years ago, one of the present writers having in 1874 seon no less than seven examples at one time from the top of a mountain ridge; of late yoars their uumbers have, however, been greatly diminished. The Brown Boar, although with strong powers of smelling, is very slow of sight and hearing, and in the Himalaya it is easy to approach so neat that they may be shot with a smooth-bore gun, The Grizzly Roar (U0 horribilis) of North Amorica is so closely allied to the Brown Boar that some writers think it should only rauk as a very wellmatked local variety. The Black Bears of the Himalaya (U. torquatus), dapan (U. japonicus), aud North America (U7. americanus) bolong to this group, The Limalayan species ranges from Persia to Assam, and thence to China and Formosa. In the greater part of this area it is essontiilly a forest animal, and may be found in autumn in tho forests of the Kashmir valley feeding upon chestiuts and other fruits. It is also oxcoedingly fond of maize, mulberries, and walunts; and a fow years ago it was no very uncommon sight to see three or even tive of these bears up a single mulberry ev walunt tree in Kashmir, The Speetacled Boar (UL ornatus) of the Peruvian Andes is another member of this group. The fMelurctine group is represented only by the Malay Bear or Sun Bear (U. malayacis), IM which the head is short and broad ; the mokiw teeth ace comparatively broad (but the length still exceeding the breadth), the tougne is very long and extensile, and the fur short aud smooth, ‘This small species inhabits the Malay Peninsula, Sumatra, Java, Borneo, Tenasserim, Arakan, Chittagong, and the Gave hills of India; it inhabits forest districts, and is an expert climber, Tho earliest known ovcurrence of the genus is in the Lower Pliocone of the Indian Stwalik Hills; where it is represented by Oy thevbaldt, which was probably the ancestor of the existing Melursus. The geuns is represented in the Upper Phoecene of Europe by the small U) efruscus sand in the Pleistocene by the exist- ing Uv aredos, as welbas by the great extinet Cave- Bear (U7 spelcus), distinguished by the complexity of the crowns of the molars and 560 CARNIVORA the total loss of the three anterior premolars in the adult condition. Remains of Bears are also found in cavern-deposits in the north of Africa. The small U. namadicus, from the Pleistocene of the Narbada valley, India, may have been allied to U. malayanus. Melursus—This differs from the true Bears in the first upper incisor being absent or shed at a very early age, in the very small size of the other teeth, in the very large extensile lips, the deep concavity of the palate, and other minor characters. The one species, Jf. labiatus, the well-known Sloth-Bear of India, feeds chiefly on black ants, termites, beetles, fruit, honey, etc. This species inhabits peninsular India, from near the Himalaya to Cape Comorin and Ceylon, and its remains are found in the cavern-deposits of Madras. The black hair is very long and coarse; there is a light horse-shoe-shaped mark on the chest (as in Ursus torquatus), and the extremity of the muzzle is of an ashy gray. Fic. 255.—.Eluropus melanvleucus. (From Milne-Edwards.) Eluropus.2-—Dentition: i 3,¢4, p4,m2; total 40. Premolars large, increasing in size from first to last, and two-rooted except the first. First upper molar with quadrate crown, broader than long ; second larger than the first. Cranium with zygomatic arches and sagittal crest immensely developed, and ascending ramus of mandible very high, giving greater spaces for attachments of temporal muscle than in any other existing member of the order. Facial portion 1 Meyer, Uebersicht d. nev. Zool. Entdeckungen, ete. p. 155 (1793). 2 A. Milne-Edwards, Nouv. Arch. du Muséum, vol. vii. Bull. p- §8 (1871). Amended from ‘* Ailuropus.” CASIDE 361 short. Bony palate not extending behind the last molar tooth. No alisphenoid canal. Feet bear-like, but soles more hairy, and perhaps less completely plantigrade. Fur long and thick. Tail very short. One extremely rare species. 4. melanoleucus (Fig. 256), discovered by Pere David in 1889. in the most inaccessible mountains of Moupin in Eastern Tibet. Said to feed principally on roots, bamboos, and other vegetables. It is of the size of a small Brown Bear, of a white colour, with ears, spots round the eves, shoulders and limbs black. In the large size and complex crowns of the upper premolars this genus ditfers very markedly from the true Bears. The fourth upper premolar (car- nassial) makes no approach to the markedly seetorial type presented by the corresponding tooth of Hyvnarctus, its structure being, on the whole, more like that of 2% irs. Extinct Gonere—The genus frefotherium includes some very large Bear-like animals from the Pleistocene of South America and California, in which the dentition departs less widely from a nor- mal carnivorous type than in the true Bears. Thus the upper car- nassial (Fig. 257). is relatively larger than in Ursus: while the erowns of the upper molars are broader and shorter. The humerus is said to have an entepicondylar —— fora- men. Hyenarefas. ot the Miocene and Plio- eene of Europe and Southern Asia. has the erowns of the upper molars either square or triangular: the upper carnassial having three distinet. lobes to the blade, while the lower earnassial is practically indistinguishable from that of the Dog-like Dinero (p. 558). The proximal extremity of the ulna differs from that of Ursvs in having a long olecranon, and thereby re- sembles the corresponding bone of the Dezs. Indeed all the characters at present available tend to show a complete passage trom the Tertiary Dog-like animals. through Dinceyon, Hyenaretis, 30 237.— Palate of arciclerivm benariznse. Pleistoeene. A Ameriea—} natural size (Prom the Pulawciedoyia Indieu.) Py) 562 CARNIVORA and Arctuthertwm, to the true Bears. Most of the species of Hyce- nurctus were of very large dimensions, but smaller forms occur in the Miocene. Cephalogale, of the Continental Tertiaries, is a genus represented by several species of medium size showing evident signs of affinity with Hyenarctus. The upper molars have sub- triangular crowns, while the carnassial is short, and has two com- paratively low lobes. Here also may be mentioned several other genera, apparently more or less closely allied to the present group, some of which are regarded by Dr. Schlosser as showing marked signs of affinity to the Procyonide. Among these are Simocyon from the Pliocene of Europe, with p <7 m 2%; and Enhydrocyon of the North American Miocene, with p 3, m 2, a secant talon to the lower carnassial, and a very short skull. The Miocene lurodon comprises several large North American forms, having a trilobed upper carnassial like that of Hyewnarctus, and a dental formula similar to that of the latter and Canis Prohyena is founded upon a much-worn jaw of Alwrodon. Hycnocyon, of the Miocene of the United States, with p 2, m 1, appears to be an allied form, also having a trilobed upper carnassial. Family PROCYONID#. True molars 3, tuberculated or multicuspid ; upper carnassial short and broad. Alisphenoid canal absent, except in Zlurus. Feet plantigrade. Tail generally annulated. In some cases an entepicondylar foramen to the humerus. Typically American, but with the outlying Oriental genus Alurus. Alurus..—Dentition : ¢ 3, ¢ 4, p 2,m 2; total 38. First lower premolar very minute and deciduous. Molars (Fig. 259) remark- able for their great transverse breadth and the numerous cusps of their crowns. Vertebre: C 7,D14,L6,83,C18. Skull (Fig. 259) high and compressed, very convex, with the facial portion short, the palate convex antero-posteriorly, and the ascending ramus of mandible extremely high. Head round. Face short and broad. Ears large, erect, pointed. Limbs stout, with large sharp semi- retractile claws. Tail nearly as long as body, cylindrical, annulated, and clothed with long hairs. Fur long and thick. One existing species, 2. fulgens, the Panda (Fig. 258), an animal rather lazeer than a Cat, found in the South-East Himalaya, at heights of feo 7,000 to 12,000 feet above the sea, among rocks and trees, and chiefly feeding on fruits and other vegetable substances. Its fur is of a remarkably rich reddish-brown colour, darker below. The genus -£7urus has been made the type of a distinct family, 1 F. Cuvier, Hist. Nat. des Mammifercs (1825), Amended from “Ailurus,” For anatomy, see Flower, Proc. Zovl. Soc., 1870, p. 752. PROCYONIDE . 563 but its relationship to the Raccoons is regarded by Mr. W. T. Blanford ! as sufficiently close to admit of its being included in the same family. According to this zoologist the Panda often sleeps coiled up like a Cat, with the bushy tail over its head, but at other times resting on its legs with the head tucked under the chest and between the fore legs, after a manner said to be common with the Raccoons. Although by no means strictly nocturnal, these animals sleep much during the day, and roam out in search of food in the morning and evening. The young are born in a very helpless Fic, 258.—The Panda (lurus fulgens). The dark nasal stripe shown in this figure is generally absent. (From Sclater, Proc. Zool. Soc. 1869, p. 408.) condition, and remain for a long period concealed in the holes of trees or rocks. Fossil remains of a species of Alurus (42. anglicus) have been obtained from the English Pliocene Crag deposits which indicate an animal of about one and half times the size of &. fulgens. The first evidence of this fossil species was afforded by part of the mandible with the last molar in place, and the subsequent discovery of an entire first upper molar renders full confirmation of the generic determination. This distribution of 4lurus is very important, as showing how its area may have once approximated to that of the ancestors of the American representatives of the family. It is probable that the genus existed in India during the Siwalik period. 1 Fauna of British India, ‘‘ Mammalia,” p. 189 (1888). 564 CARNIVORA The whole of the undermentioned genera are American, and ave characterised by the absence of an alisphenoid canal in the skull. Proeyon3—Dentition : i #,¢4, p 4, m #2; total 40. The molar teeth broad and tuberculated (Fig. 259). The upper carnassial with three cusps along the outer margin, and a very broad bicuspid inner tubercle, giving an almost quadrate form to the crown. First molar with a large tuberculated crown, rather broader than long ; Fig. 259.—Lateral view of skull and right half of palate of .klurus Julgens, (Prom Blanford, Mammatia of British India, p. 190.) second considerably smaller, with transversely oblong crown. Lower carnassial with an extremely small and ill-defined blade, placed transversely in front, and a large inner cusp and hind talon. Second molar as long as the first, hut narrower behind, with tive obtuse cusps. Vertebre: C 7, D 14, L 6,8 3, C 16-20. Body stout. Head broad behind, but with a pointed muzzle. Limbs 1 Storr, Prodromus Meth, Maman, p. 85 (1780). PROCVONID-E 565 plantigrade, but in walking the entire sole is not applied to the ground as it is when the animal is standing. Toes, especially of the fore foot, very free, and capable of being spread wide apart. Claws compressed, curved, pointed, and non-retractile. Tail moder- ately long, cylindrical, thickly covered with hair, annulated, non- prehensile. Fur long, thick, and soft. The well-known Raccoon 4 (Procyon lotor, Fig. 260) of North America is the type of this genus. It is a clumsy thickly-built animal about the size of a Badger, with a coat of long coarse grayish-brown hairs, short ears, and a bushy black and white ringed tail. Its range extends over the whole of Fie. 260.—The Raecoon (Procyon lotor). the United States, and stretches on the west northwards to Alaska and southwards into Central America, where it attains its maximum size. The following notes on the habits of the Raccoon are taken from Dr. C. H. Merriam’s MJummuals of the Adirondack Region : “Raccoons are omnivorous beasts, and feed upon mice, small birds, birds’ eggs, turtles and their eggs, frogs, fish, crayfish, molluscs, insects, nuts, fruits, maize, and sometimes poultry. Ex- cepting the bats and flying squirrels, they are the most strictly nocturnal of all our mammals, and yet I have several times seen them abroad on cloudy days. They haunt the banks of ponds and streams, and find much of their food in these places, such as 6c 1 A corruption of the North American Indian ‘‘arrathkune”’ or ‘‘arathcone.” The French raton or raton laveur, German Waschbér, and other European names are derived from a curious habit the Raccoon has of dipping or washing its food in water before eating it. 566 CARNIVORA crayfish, mussels, and fish, although they are unable to dive and pursue the latter under water, like the otter and mink. They are good swimmers, and do not hesitate to cross rivers that lie in their path. . . . The Raccoon hibernates during the severest part of the winter, retiring to its nest rather early, and appearing again in February or March, according to the earliness or lateness of the season. It makes its home high up in the hollow of some large tree, preferring a dead limb to the trunk itself. It does little in the way of constructing a nest, and from four to six young are commonly born at a time, generally early in April in this region. The young remain with the mother about a year.” The South-American P. cancrivorus, the Crab-eating Raccoon, is very similar to P. lotor, but differs by its much shorter fur, larger size, proportionally more powerful teeth, and other minor characters. It extends over the whole of South America, as far south as the Rio Negro, and is very common in all suitable localities. Its habits are similar to those of the North-American species. Fossil remains of Procyon have been described from the Pleistocene deposits of the United States. Bassaris.\—A. form closely allied to Procyon, but of more slender and elegant proportions, with a sharper nose, longer tail, and more digitigrade feet, and with teeth otherwise like, but smaller, and more sharply denticulated. It was formerly, but erroneously, placed among the Viverride. Two species :—B. astuta, from the southern parts of the United States and Mexico, and B. sumichrasti, from Central America. Bassaricyon.,—This name has been given to a distinct modifica- tion of the Procyonine type of which at present only two examples are known, one from Costa Rica and the other from Ecuador, which, appearing to be different species, have been named B. gabbi and B. allent. They much resemble the Kinkajou (Cercoleptes) in external appearance, but the skull and teeth are more like those of Procyon and Nasua. Nasua.2—Dentition as in Procyon, but the upper canines are larger and more strongly compressed, and the molars smaller. The facial portion of the skull is more elongated and narrow. Verte- bre: C7, D 14, L 6,8 3, C 22-23. Body elongated and rather compressed. Nose prolonged into a somewhat upturned, obliquely truncated, mobile snout. Tail long, non-prehensile, tapering, annu- lated. These animals, commonly called Coatis or Coati-Mundis, live in small troops of eight to twenty, are chiefly arboreal, and feed on fruits, young birds, eggs, insects, etc. Recent researches have reduced the number of supposed species to two, WN. narica of Mexico 1 Lichtenstein, Isis, 1831, p. 512. * Allen, Proc. Ac. Nat. Sei. Philad, 1876, p. 20. ® Storr, Prodromus Meth. Mamm. p. 35 (1780). MUSTELIDA 567 and Central America, and NV. rufa of South America from Surinam to Paraguay. Remains of this genus, mostly referable to the existing species, occur in the cavern-deposits of Brazil. Cercoleptes..—Dentition : i 3, ¢ 4, p 3, m 3; total 36. Molars with low flat crowns, very obscurely tuberculated. Skull short and rounded, with flat upper surface. Vertebre: C 7, D 14, L 6, 8 3, C 26-29. Clavicles present, but in a very rudimentary condition. Head broad and round. Ears short. Body long and musteline. Limbs short. Tail long, tapering, and prehensile. Fur short and soft. Tongue long and very extensile. But one species of this somewhat aberrant genus is known, (. caudivolvulus, the Kinkajou, found in the forests of the warmer parts of South and Central America. It is about the size of a Cat, of a uniform, pale, yellowish- brown colour, nocturnal and arboreal in its habits, feeding on fruit, honey, eggs, and small birds and mammals, and is of a tolerably gentle disposition and easily tamed. Family MUSTELIDA. True molars 4 (or + in Mellivora?). No alisphenoid canal. In the upper molar the inner tubercular portion is always longer in the antero-posterior direction than the secant external portion ; the degree of inflation of the auditory bulla is but slight; and the palate is generally much produced behind the last molars, as is the case with the members of the preceding family. The postglenoid process of the cranium is generally considerably curved over the glenoid fossa, so as to hold very tightly the condyle of the man- dible. The humerus may or may not have an entepicondylar foramen. Except in the Otters, the kidneys resemble those of the Procyonide in being of simple structure. This family is a large and widely distributed one, especially in the northern temperate regions of the earth. The different genera, which are very difficult to arrange in any natural order, are rather artificially divided, chiefly according to the characters of their feet and claws, into the Otter-like (Lutrine), Badger-like (Meline), and Weasel-like (Musteline) forms. Subfamily Lutrinee.—Feet short, rounded (except the hind feet of Latux). Toes webbed. Claws small, curved, blunt. Head broad and much depressed. Upper molar large and quadrate, with its inner tubercular portion much expanded antero-posteriorly (Fig. 261). Kidneys conglomerate. Habits aquatic. Lutra.>— Dentition: i 3, ¢ 4, p 4, m 4; total 36. Upper 1 Tlliger, Prodrumus Syst. Mamm. et Avium, p. 127 (1811). 2 Also in two other species noticed below. One extinct Otter has two upper molars. 3 Erxleben, Syst. Regn. Animal, p. 445 (1777). 568 CARNIVORA carnassial with a trenchant tricuspid blade, and a very large inner lobe, hollowed on the free surface, with a raised sharp edge, and extend- ing along two-thirds or more of the length of the blade. True molar large, with a quadri- cuspidate crown, broader than long. First upper premolar very small, and in some cases absent (Fig. 261). Skull broad and depressed, contracted 1m- mediately behind the orbits. Facial portion very short; brain case large. Vertebree: C 7, D 14-15, L 6-5, 8 3,C Fic. 261.—Palate of Lutra cinerea. (From the 20-26. Body very long. Paleontologia Indica.) Fars: short and. rounded. Limbs short. Feet more or less completely webbed ; claws usually well developed on all the toes, although they may be rudimentary or absent. Trail long, thick at the base and tapering, rather depressed. Fur short and close. The humerus may or may not have an entepicondylar foramen. In conformity with the shape of the skull, the posterior part of the brain is expanded laterally. The Common British Otter (L. vulgaris), as the type of the genus, may be described somewhat fully. It has an elongated, low body, short limbs, short broad feet, with five toes on each, con- nected together by webs, and all with short, moderately strong, compressed, curved, pointed claws. Head rather small, broad, and flat; muzzle very broad ; whiskers thick and strong; eyes small and black ; ears short and rounded. Tail a little more than half the length of the body and head together, very broad and strong at the base, and gradually tapering to the end, somewhat flattened horizontally. The fur is of very fine quality, consisting of a short soft under fur of a whitish-gray colour, brown at the tips, inter- spersed with longer, stiffer, and thicker hairs, very shining, grayish at the base, bright rich brown at the points, especially on the upper parts and outer surface of the legs ; the throat, cheeks, under parts and inner surface of the legs brownish-gray throughout. Individual Otters vary much in size ; but the total length from the nose to the end of the tail averages about 34 feet, of which the tail occupies 1 foot 3 or 4 inches. The weight of a full-sized male is from 18 to 24 lbs., that of a female about 4 Ibs. less. As the Otter lives almost exclusively on fish, it is rarely met with far from water, and usually frequents the shores of brooks, rivers, lakes, and, in some localities, the sea itself. It is a most expert swimmer and diver, easily overtaking and seizing fish in the MUSTELIDA 569 water, but when it has captured its prey it brings it to shore to devour it. When lying upon the bank it holds the fish between its fore-paws, commences at the head, and then eats gradually towards the tail, which it is said always to leave. The female produces three to five young ones at a time, in the month of March or April, and brings them up in a nest formed of grass or other herbage, usually placed in a hollow place in the bank of a river, or under the shelter of the roots of some overhanging tree. The Common Otter is found in localities suitable to its habits throughout Great Britain and Ireland, though far less abundantly than formerly, for, being very destructive to fish, and thus coming into keen competi- tion with those who pursue the occupation of fishing either for sport or for gain, it is rarely allowed to live in peace when once its haunts are discovered. Otter-hunting with packs of hounds of a special breed, and trained for the purpose, was formerly a common pastime in the country. When hunted down and brought to bay by the dogs, the Otter is finally despatched by long spears carried for the purpose by the huntsmen. The Common Otter ranges throughout the greater part of Europe and Asia, the Indian L. nair not being distinct. A closely allied but larger species, L. canadensis, is extensively distributed throughout North America, where it is systematically pursued by professional trappers for the value of its fur. The Common Otter is regularly trained by the natives of some parts of Bengal to assist them in fishing, by driving the fish into the nets. In China Otters are taught to catch fish, being let into the water for the purpose attached to a long cord. Otters are widely distributed over the earth, and, as they are much alike in size and coloration, their specific distinctions are by no means well defined. Besides those mentioned above, the following may be noticed. In the Oriental region there are L. elliott? of India, L. sumatrana of the Malay countries, and L. cinerea ranging over the greater part of the region. The latter species (often known as L. leptonyx) is of small size, with a short head, and rudimentary claws, which may be absent; it was at one time regarded as generically distinct, under the name of donyx. The upper true molar (Fig. 261) is characterised by the great develop- ment of its inner tubercular portion, and the first upper premolar is absent. In the Ethiopian region there are two species, L. capensis and L. maculicollis, Of the Neotropical forms it will suffice to mention the small L. felina and the large L. brasiliensis. The latter is by far the largest of the existing forms, and is characterised by the presence of a prominent flange-like ridge along each lateral 1 See Thomas, Proc. Zool. Soc. 1889, p. 190. » The synonomy of this species is not settled, and the adoption of the name given here only preliminary. 570 CARNIVORA margin of the tail, on which account it was referred by Dr. Gray to a distinct genus, with the name of Pteronura sambachi. It should be observed that all Otters have a very distinct inner cusp to the blade of the lower carnassial, but that the relative size of this cusp varies in the different species. Extinct Otters. —Several species of fossil Otters have been described. Thus in the Indian Siwaliks we have L. paleindica, which is closely allied to L. swmatrana, and a larger form described as L. bathygnathus. The Pliocene of Hessen-Darmstadt yields L. hessica ; while L. dubia, of the Middle Miocene of France, is a species characterised by the small size of the inner cusp of the lower carnassial—a character in which it resembles those Tertiary forms deseribed as Trochictis, which are believed to connect Lutra with the Musteline. Two very large Otters, respectively from the Indian Siwaliks and the Italian Miocene, named L. sivalensis and L. campanii, may be regarded either as representing a very distinct Enhydriodont group of Lutra or as referable to a separate genus Enhydriodon. They are characterised by certain peculiarities in the structure of the teeth, and the second upper premolar may be absent in the Indian form. Lastly, the genus Potamotherium con- tains a small Otter (P. valetont) from the Lower Miocene of the Continent, which differs from all other known MMustelide in having a minute second upper true molar. This species is evidently a very generalised form approximating to the Viverride in its dental formula, and also in the characters of the teeth themselves. The brain, as recently described by Dr. Filhol, differs from that of Lutri and other Mustelines in the great relative width of the anterior extremity of the hemispheres and olfactory lobes, and also in the disposition of the sulci, in both of which respects it more nearly resembles the Viverride. Latux.$—Dentition: i 3, ¢ 4, p 3, mi; total 32. Differs from all other existing Carnivora in having but two incisors on each side of the lower jaw, the one corresponding to the first (very small in the true Otters) being constantly absent. Though the molar teeth generally resemble those of Lutra in their proportions, they differ very much in the exceeding roundness and massiveness of their crowns and bluntness of their cusps. Feet webbed. Fore feet small, with five subequal toes, furnished with short compressed claws; palms naked. Hind feet very large, depressed, and fin- like. The phalanges flattened as in the Seals. The fifth toe the longest and stoutest, the rest gradually diminishing in size to the first, all with moderate claws. Tail moderate, cylindrical, and obtuse ; about one-fourth the length of the head and body. 1 Gloger, Nova Acta Ac. Coes. Leop.-Car. vol. xiii. pt. 2, p. 511 (1827): Syn. Enhydra; Fleming, Philosophy of Zoology, vol. ii. p. 187 (1822). Preoceupied by Finhydris, Merrem, Tent. Syst, Amphib. p. 140 (1820). MUSTELIDA: 571 The Sea-Otter (L. lutris, Fig. 262) is the sole representative of this genus. The entire length of the animal from nose to end of tail is about 4 feet, so that the body is considerably larger and more massive than that of the English Otter. The skin is peculiarly loose, and stretches when removed from the animal so as to give the idea of a still larger creature than it really is. The pellage is remarkable for the preponderance of the beautifully soft woolly under fur, the longer stiffer hairs being very scanty. The general colour is a deep liver brown, everywhere silvered or frosted with the hoary tips of the longer hairs. These are, however, removed when the skin is dressed for commercial purposes. Fic, 262.—The Sea-Otter (Latax lutris). From Wolf, Proc. Zool. Soc. 1865, pl. vii. Sea-Otters are only found upon the rocky shores of certain parts of the North Pacific Ocean, especially the Aleutian Islands and Alaska, extending as far south on the American coast as Oregon; but, owing to the unremitting persecution to which they are sub- jected for the sake of their skins, which rank among the most valuable known to the furrier, their numbers are greatly diminish- ing, and, unless some restriction can be placed upon their destruc- tion, such as that which protects the Fur-Seals of the Pribyloff Islands, the species is threatened with extermination, or, at all events, excessive scarcity. When this occurs, the occupation of five thousand of the half-civilised natives of Alaska, who are dependent upon Sea-Otter hunting as a means for obtaining their living, will be gone. The principal hunting grounds at present are the little rocky islets and reefs around the island of Saanach and 572 CARNIVORA the Chernobours, where they are captured by spearing, clubbing, or nets, and recently by the more destructive rifle bullet. They do not feed on fish, like the true Otters, but on clams, mussels, sea- urchins, and crabs, for the mastication of which the blunt cusps of their teeth are admirably suited. The female brings forth but a single young one at a time, apparently at any season of the year. They are excessively shy and wary, and all attempts to rear the young ones in captivity have hitherto failed. Subfamily Meline.—Feet elongated. Toes straight. Claws non-retractile, slightly curved, subcompressed, blunt; those of the fore foot especially large. Upper molar variable. Kidneys simple. Habits mostly terrestrial and fossorial. Mephitis.\A—Dentition: + 3, ¢ 4, p 3, m 4; total 34. Upper molar larger than the carnassial, subquadrate, rather broader than long. Lower carnassial with talon less than half the length of the whole tooth. Bony palate terminating posteriorly opposite the hinder border of the last molar tooth. Facial portion of skull short and somewhat truncated in front. Vertebre: C 7, D 16, L 6,8 2, C 21. Head small. Body elongated. Limbs moderate, subplantigrade. Ears short and rounded. Tail long, abundantly clothed with very long fine hair. Anal glands largely developed. The secretion of these glands, which can be discharged at the will of the animal, has an intolerably offensive odour, which circumstance has rendered the Skunks, as they are commonly called, proverbial. They are strictly nocturnal animals, terrestrial and burrowing, feed- ing chiefly on small mammals, birds, reptiles, insects, worms, roots, and berries. All the known species have a prevalent black colour, varied by white strips or spots on the upper part (Fig. 263). They generally carry the body much arched, and the tail erect, the long loose hair of which waves like a plume over the back. There are three species, all inhabitants of the American continent, over which they have an extensive range. The Common Skunk (J mephitica, Fig. 263) is an animal of about the size of a small Cat, ranging from Hudson’s Bay to Guatemala. The following account of its habits is given by Dr. C. H. Merriam in his Mammals of the Adirondack Region :-— “The skunk preys upon mice, salamanders, frogs, and the eggs of birds that nest on or within reach from the ground. At times he eats carrion, and if he chances to stumble upon a hen’s nest the eggs are liable to suffer; and once in a while he acquires the evil habit of robbing the hen-roost, but as a rule skunks are not addicted to this vice. Of all our native mammals perhaps no one is so universally abused and has so many unpleasant things said about it as the innocent subject of the present biography ; and yet no other species is so valuable to the farmer. Pre-eminently an insect-eater, Cuvier, ‘‘Tabl. de Classif.” in Legons d’ Anat. Compar. vol. i. (1800). MUSTELIDE 573 he destroys more beetles, grasshoppers, and the Tike than all ow: other mammals together, and in addition to these he devours vast numbers of mice. He does not evinee that dread of man that is se manifest in the great majovity of our mammals, and when met during any of his cireumambulations rarely thinks of running away. He is slow in movement and deliberate in action, and does not often huvry himself in whatever he does. His ordinary gait is a measured walk, but when pressed for time he breaks into a low shutting eullop. It is hard to intimidate a skunk, but when onee really frightened he manages to get over the ground at a very fair pace. Pia. 268. —The Common Skunk (Mephitis meprition, Skunks remain active throughout the greater part of the year in this region, and hibernate only during the severest portion of the winter. ‘They differ from most of our hibernating mammals in that the inactive period is apparently dependent solely on the tempera- ture, while the mere amount of snow has no intluence whatever upen their movements. Skunks, particularly when young, make very pretty pets, being attractive in) appearance, gentle in disposition, interesting in manners, and cleanly in halits—rare qualities indeed! They are playful, sometimes mischievous, and manifest considerable affection for these who have the care of them. Their tlesh is white, tender, and sweet, and is delicious eating. Skunks have large families, from six to ten young being commonly 574 CARNIVORA raised each season; and as a rule they all live in the same hole until the following spring.” The two ducts leading from the anal glands open at the tips of two small conical papille placed in such a position that the animal can protrude them externally, and can thus guide the direction of the jet of nauseous fluid, which can be propelled by the powerful muscles surrounding the glands to a distance of from 8 to 12 feet. The Long-tailed Skunk (JZ. macrura), from Central and Southern Mexico, has two lateral stripes, and a longer and more bushy tail than the common species. Jf putorius, of the Southern United States and thence southwards to Yucatan and Guatemala, is of a much smaller size, with four interrupted white lateral stripes, and a skull differing considerably in form from that of the type species. It is regarded by some writers as representing a distinct genus, Spilogale ; and has been recently divided by Dr. C. H. Merriam into several nominal species. Conepatus.\—The Skunk of tropical America (C. mapacito), ranging from Texas to Chili and Patagonia, differs considerably from the true Skunks, although in colour it is almost precisely similar to the common species, with which it also agrees in the variation of the relative development of the black and white. Its build is heavier than that of Mephitis ; the snout and head are more Pig-like ; and the nostrils open downwards and forwards instead of laterally on the sides of the muzzle. The skull also has many special characters, and the teeth are different in shape and, as a rule, in number also, the first minute premolar of J/ephitis being almost invariably absent, so that the dental formula is i 3, ¢ 4, p 2, m 4; total 32. Remains of Conepatus, which have been referred to three species, are found in the cavern-deposits of Brazil. Arctonyx.>—Dentition: i 3, ¢ 4, p 4,m4; total 38. Incisor Ine curved, the outer teeth being placed posteriorly to the others. Lower incisors proclivous. First premolars often rudimentary or absent. Upper molar much larger than the carnassial, longer in the antero-posterior direction than broad; lower carnassial with a very large, low, tuberculated talon. Cranium elongated and depressed ; face long, narrow, and concave above. Bony palate extending as far backwards as the level of the glenoid fossa ; palatal bones dilated ; suborbital foramina very large. Vertebre: C7, D16, L 4,84, C20. Snout long, naked, mobile, and truncated, with large terminal nostrils, much like that of a Pig. Eyes small. Kars very small and rounded. Body compressed rather than depressed. Limbs of moderate length and digitigrade in walking. Gray, dan. Mag. Nat. Hist. sev. 2, vol. i. p. 581 (1837). ? F. Cuvier, Hist. Nat. des Mammiferes (1825). AWUSTELID AL 575 Tail moderate, tapering. A full soft under fur, with longer, bristly hairs interspersed. The best-known species is 4. collavis, the Sand- Badger, or Bhdlu-soor} (i.e. Bear-pig) of the natives, found in the mountains of the north-east of India and Assam. It is rather larger than the English Badger, higher on its legs, and very Pig-like in general aspect, of a light gray colour, with flesh-coloured snout and feet ; and is nocturnal and omnivorous in habits. The imper- fectly known 4. tavoides from Assam and Arakan, and perhaps China, is a much smaller species. A third form probably exists in Eastern Tibet. Professor Mivart remarks that the brain-case of -lrctonyx is narrower than in any other Arctoid; while the palate is relatively longer than in any other Carnivore except Procyon ; and the metatarsus is relatively shorter than in any other member of the order. AMydaus.-—Dentition as in the last genus, but the cusps of the teeth more acutely pointed. Cranium elongated, face narrow and produced. Suborbital foramen small, and the palate, as in all the succeeding genera of this group, produced backwards about midway between the last molar tooth and the glenoid fossa. Vertebre: C7, D 14-15, L6-5, 8 3, C12. Head pointed in front; snout produced, mobile, obliquely truncated, the nostrils being inferior. Limbs rather short and stout. Tail extremely short, but clothed with rather long bushy hair. Anal glands largely developed, and emitting an odour like that of the American Skunks. One species, J. meliceps, the Teledu, a small burrowing Badger, found in the mountains of Java at an elevation of 7000 or more feet above sea-level. Aeles.>—Dentition : 1 3, ¢ 4, p 4, m 4; total 38. The first premolar in both jaws extremely minute and often deciduous. Upper molar very much larger than the carnassial, subquadrate, as broad as long. Lower carnassial with a broad, low, tuberculated talon, more than half the length of the whole tooth. The postglenoid processes of the skull are so strongiy developed, and the glenoid fossa is so deep, that the condyle of the lower jaw is firmly held in its place even after all the surrounding soft parts are removed. Vertebre: C7, D15, L5,83,C 18. Muzzle pointed. Ears very short. Body stout, broad. Limbs short, strong, subplantigrade. Tail short. The best-known species is the common Badger (Jf. turus) of Europe and Northern Asia, still found in many parts of England, where it lives in woods, is nocturnal, burrowing, and very omni- vorous, feeding on mice, reptiles, insects, fruit, acorns, and roots. Other nearly allied species, AL. leucurus and Jf. chinensis, are found in continental Asia, J. canescens in Persia, and 1. anakwme in Japan. The appearance of the common Badger is too well known to 1 Possibly the name should be Balu-soor (Sand-pig). ° F. Cuvier, Hist. Nat. des Mammiferes (1825). 3 Storr, Prodromus Meth. Mamm. p. 84 (1780). 576 CARNIVORA need description, but}it may be mentioned that a full-grown individual stands about a foot in height at the shoulder, and measures from 24 to 3 feet in length. The young are born in a naked and blind condition, usually in litters of three or four. It appears that the usual period of gestation is about eleven and a half months, but instances are recorded where the period has been protracted to upwards of fifteen months. Fossil remains of the common Badger are found in the Pleistocene deposits of Europe, while extinct species have been described from the Lower Pliocene beds of Maragha, in Persia. Taxidea..—Dental formula as in JA/eles, except that the rudi- mentary anterior premolar appears to be always wanting in the upper jaw. The upper carnassial much larger in proportion to the other teeth. Upper molar about the same size as the carnassial, triangular, with the apex turned backwards. Talon of lower car- nassial less than half the length of the tooth. Skull very wide in the occipital region ; the lambdoidal crest very greatly developed, and the sagittal but slightly, contrary to what obtains in Jfeles. Vertebre: C7, D 15, L 5,8 3, C 16. Body very stoutly built and depressed. Tail short. The animals of this genus are peculiar to North America, where they represent the Badgers of the Old World, resembling them much in appearance and _ habits. T. americana is the common American Badger of the United States ; Tf. berlandieri, the Mexican Badger, is perhaps only a local variety. Meltivora.,—Dentition : ¢ 3, ¢ 4, p 3, m 1; total 32. Upper carnassial large, with its inner tubercle quite at the anterior end of the blade, as in the following genera; molar much smaller and transversely extended, having a very small outer and a larger rounded inner lobe. Talon of lower carnassial very small, scarcely one-fourth of the whole length of the tooth, and with but one cusp ; lower tubercular molar absent. Vertebre: C7,D14,L4,8 4, C15. Body stout, depressed. Limbs short, strong. Head depressed, nose rather pointed. External ears rudimentary. Tail short. The animals of this genus are commonly called Ratels. JZ. indica from India, and Jf. ratel (Fig. 264) from South and West Africa, have nearly the same general appearance and size, being rather larger than a common Badger. Their coloration is peculiar, all the upper surface of the body, head, and tail being ashy gray, while the lower parts, separated by a distinct longitudinal boundary line, are black. The two species may be distinguished by the circumstance that the African one has a distinct white line round the body at the junction of the gray of the upper side with the black of the lower, while in the Indian form this line is absent; the teeth also of the former are, on the whole, larger, rounder, and heavier than those of 1 Waterhouse, Prov. Zool. Soc. 1838, p. 154. * Storr, Prodromus Meth, Mamm. p. 34 (1780). MUSTELIDAZ 577 the latter. In spite of these differences the two are, however, so nearly allied that they might almost be considered as local races of a single widely spread species. The following account of the Indian species is extracted from Dr. Jerdon’s Mammals of India: “The Indian badger is found throughout the whole of India, from the extreme south to the foot of the Himalayas, chiefly in hilly districts, where it has greater facilities for constructing the holes and dens in which it lives ; but also in the north of India in alluvial plains, where the banks of Fia, 264.—The African Ratel (Mellivora ratel). large rivers afford equally suitable localities wherein to make its lair. It is stated to live usually in pairs, and to eat rats, birds, frogs, white ants, and various insects, and in the north of India it is accused of digging out dead bodies, and is popularly known as the grave-digger. It doubtless also, like its Cape congener, occasionally partakes of honey. It is often very destructive to poultry, and I have known of several having been trapped and killed whilst committing such depredations in Central India and in the northern Cirecars. In confinement the Indian badger is quiet and will partake of vegetable food, fruits, rice, etc.” A fossil species of Mellivora, apparently closely allied to the existing forms, occurs in the Pliocene Siwaliks of India. The same deposits have also yielded remains of an extinct genus described as Mellivorodon. 37 578 CARNIVORA Helictis\—Dentition : i 3, ¢ 3, p 4, m 4; total 38. Upper carnassial with a large bicuspid inner tubercle ; upper molar smaller, wider transversely than in the antero-posterior direction. Lower carnassial with talon about one-third the length of the tooth. Skull elongated, rather narrow and depressed. Facial portion especially nar- row. Infra- orbital foramen very large. Head rather small and pro- duced in front, with an elon- gated, obliquely truncated,naked Fic. 265.—Helictis personata. (From Blanford, Mammalia of British gnout. Ears India, p. 175.) small. Body elongated. Limbs short. Tail short or moderate, bushy. Several species are described (ZH. orientalis, personata [Fig. 265], moschata, subawrantiaca), all from Eastern Asia; they are all small animals compared with the other members of the subfamily, climbing trees with agility and living much on fruit and berries as well as on small mammals and birds. The two first named species occur in British India, H. orientalis also ranging into Java; the Chinese fZ, subawrantiaca is brilliantly coloured in the region of the throat.? Fic, 266.—Left lateral and superior aspect of the brain of Helictis sabaurantiaca. (From Garrod, Proc. Zool. Soc. 1879, p. 807.) The brain of Helictis, represented in the accompanying figure, shows the general type of cerebral structure characteristic of the Mustelide. The brain of this genus differs, however, from that of every other Carnivore in that the hippocampal gyrus rises to the surface on either side of the great longitudinal fissure, in 1 Gray, Proc. Zool. Soc. 1831, p. 94. ° Garrod, ibid. 1879, pl. xxix. MOUSTELIDA : 579 consequence of which there is no crucial fissure, and the so-called “Ursine lozenge,” so characteristic of the Arctoidea, is incomplete behind. The superior gyrus, as in Ictonyx and Afustela, ceases at the superior posterior angle of the hemisphere. Lctonyx.A—Dentition : i 3, ¢4, p 3, m4; total 34. In general characters the teeth much resemble those of the Polecats (JMustela), being more delicately cut and sharply cusped than in most of the foregoing. Upper molar smaller than the carnassial, narrow from before backwards. Lower carnassial with a small narrow talon and distinct inner cusp. General form of body Musteline. Limbs short. Fore feet large and broad, with five stout, nearly straight, blunt, and non-retractile claws, of which the first and fifth are considerably shorter than the others. Tail moderate, with longer hairs towards the end, giving it a bushy appearance. Hairs generally long and loose. The best-known species of this genus, J. zorilla, the Cape Polecat, was placed by Cuvier in the genus Jfustela, and by Lichtenstein in Mephitis; and in many characters it forms a transition between these genera. It is about the size of an English Polecat, but conspicuous by its coloration, having broad, longitudinal bands of dark brown, alternating with white. Its odour is said to be as offensive as that of the American Skunks. From the Cape of Good Hope it ranges as far north as Senegal. Another species, I. frenata, from Sennaar and Egypt, has been described. Subfamily Mustelinze.—Toes short, partially webbed; claws short, compressed, acute, curved, often semiretractile. Upper molar of moderate size, wide transversely. Kidneys simple. Terrestrial and arboreal in habits. Galictis."—Dentition: 7 3,¢ 4, p #,m4; total 34. Molars small but stout. Upper carnassial with the inner tubercle near the middle of the inner border of the tooth. Lower carnassial with talon small, and inner cusp small or absent. Body long. Limbs short ; claws non-retractile. Palms and soles naked. Head broad and depressed. Tail of moderate length. The best-known species are G. vittata, the ‘vison (genus Grisonia, Gray), and G. barbara, the Tayra (genus Galera, Gray), both South American; G. allamandi is an inter- mediate form. Remains of (alictis occur in the Pleistocene cavern-deposits of Brazil, and also in the Pleistocene of North America. Mustela.2—Dentition: i 8, ¢ 4, p aaa m4; total 34 or 38. Upper carnassial with inner tubercle close to the anterior edge of the tooth. Molar nearly as large as carnassial. Lower carnassial with small or no inner cusp. Vertebre: C7, D14, L6, 8 3, C 18-23. Body long and slender. Limbs short, digitigrade. Feet 1 Kaup, Thierreich, vol. i. p. 352 (1835). 2 Bell, Proc. Zool. Soc. 1837, p. 45. 3 Linn. Syst. Nat. 12th ed. vol. i. p. 66 (1766). 580 CARNIVORA rounded ; toes short, with compressed, acute, semiretractile claws. Tail moderate or long, more or less bushy. The genus Alustel, as restricted by Cuvier (Jteyne Animal, 1817), contains a very natural assemblage of animals commonly called Martens, Sables, Polecats, Stoats, Ermines, and Weasels, all closely allied in structure and habits. A structural division, however, occurs between the two first-named and all the others, especially shown in the presence of an additional small premolar tooth on each side of the jaw; and, availing himself of this and some other minor characters, Cuvier divided the genus into two subgenera, for the first of which he retained the name of J/ustela, and to the second assigned that of Puforius. Three years later Nilsson (Skand. Fauna, 1820) definitely constituted the two groups into genera, applying to the first the name of Jartes, by which the animals composing it had been generally designated by the Latin-writing zoologists of the preceding century, and keeping A/ustela for the more typical Weasels and their immediate allies. Later zoologists have been divided between the nomenclature of Cuvier, which has the priority, and that of Nilsson, which on other grounds is pre- ferable. Those who adopt the latter affirm that Cuvier’s names, being only used by him in a subgeneric sense, and not binominally, need not be applied generically, but this is contrary to the practice usually followed in such cases ; and therefore, if the original genus be divided, the name J/ustela should be retained for the Martens, and Putorius for the Polecats and Weasels. Here, however, the genus will be employed in its wider sense, and divided into two groups. The typical group of the Martens! presents the following distinctive features. Body long, slender, and very flexible, though less so than in the true Weasels. Head somewhat triangular; muzzle pointed, the nose extending a little beyond the lips; eyes large and prominent; ears conspicuous, broad, somewhat triangular, rounded at the ends, furred outside and in. Limbs short; feet rounded ; toes short, five on each foot, all with short, compressed, curved, sharp-pointed claws; soles densely furred between the naked pads. Tail moderately long, more or less bushy. Outer fur long, strong, and glossy; a very abundant soft under fur. Skull elongated and depressed. Facial portion moderate and rather compressed. Zygomata arched and wide, but slender. Postorbital processes small. Auditory bull large, but not very globose. Mandible with a strong triangular vertical coronoid process and a well-developed angular process. Premolars 4 Upper incisors in a straight transverse line, rather long and ? By all old authors of authority,"as Ray, Pennant, Shaw, and Fleming, the word is written ‘‘ Martin,” but this form of spelling is now generally reserved by way of distinction for the bird. The term ‘Marten-Cat,” often used, is a misnomer, MUSTELIDE 581 compressed ; first and second subequal, third considerably larger. Lower incisors very small, especially the first, and crowded together, the second placed rather behind the others. Canines long and sharp-pointed. Upper premolars: first very small, with simple crown and one root; second and third nearly equal in size and two-rooted, with simple compressed sharp-pointed crowns, with very slightly developed accessory cusps; fourth (the carnassial) with blade consisting chiefly of the central and posterior lobes, the anterior being rudimentary, inner tubercle small and confined to the anterior part of the tooth. True molar tubercular, about twice as wide transversely as in the antero-posterior direction, having an outer, more elevated, but smaller portion, bearing three blunt tubercles ; to the inner side of this the crown is contracted, and its surface deeply hollowed; it then expands again into a broad low lobe, with the central part elevated, and a raised, even, semicircular, slightly crenated internal border. Lower premolars : first very small, simple, and one-rooted ; second, third, and fourth increasing slightly in size, with high compressed pointed crowns and posterior accessory cusps, best marked in the third. First molar (carnassial) with well-marked bilobed blade, talon scarcely more than one-third of the length of the tooth, and a very small inner cusp. Second molar small, single-rooted, with a low, flattened, subcircular or oval tubercular crown. In geographical distribution the Martens are limited to the northern hemisphere, ranging throughout the greater part of the temperate regions of both Old and New Worlds, as far north as conditions of existence suited to their habits are met with, and southwards in America to 35° N. lat., while in Asia one species is met with as far in this direction as the island of Java. The various species appear to be very similar in their habits. They live in woods and rocky places, and are thoroughly arboreal, spending most of their time in trees, although descending to the ground in quest of prey. They climb with great facility, and are agile and graceful in their movements. Some species are said occasionally to resort to berries and other fruit for food, but as a rule they are strictly carnivorous, feeding chiefly on birds and their eges, small mammals, as squirrels, hares, rabbits, and moles, but chiefly mice of various kinds, of which they destroy great numbers, and occasionally snakes, lizards, and frogs. In proportion to their size they are among the most bloodthirsty of animals, though less so than the true Weasels. The female usually makes her nest of moss, dried leaves, and grass in the hollow of a tree, but sometimes in a hole among rocks or ruined buildings, and produces several young at a birth, usually from four to six. Though wild and untameable to a great degree if captured when fully grown, when taken young they are very docile, and have frequently been made \ 582 CARNIVORA pets of, not having the strong unpleasant odour of the smaller Mustelide. The common European Marten appears to have been partially domesticated by the Greeks and Romans, and to have been used to keep houses clear from rats and mice before cats were introduced.!_ In the same way, according to Hodgson, the Yellow- bellied Weasel (JZ. cathia) “is exceedingly prized by the Nipalese for its service in ridding houses of rats. It is easily tamed; and such is the dread of it common to all Murine animals that not one will approach a house where it is domiciled.” It is, however, to the great value attached to the pelts of these animals that their importance to man is chiefly due. Though all yield fur of serviceable quality, the commercial value varies immensely, not only according to the particular species from which it is obtained, but according to individual variation, depending upon age, sex, season, and other trifling circumstances. The skins from northern regions are more full and of a finer colour and gloss than those from more temperate climates, as are those of animals killed in winter compared with the same individuals in the summer season. The caprices of fashion have, moreover, set wholly factitious values upon slight shades of colour, recognised and named by experienced furriers, but not: indicating any specific or other distinctions of which zoologists have any cognisance. Enormous numbers of animals are annually caught, chiefly in traps, to supply the demand of the fur trade, Siberia and North America being the principal localities from which they are obtained. With the exception of the Pekan (JZ. pennanti) all the Martens are so much alike in size, general colouring, and cranial and dental characters that the discrimination of the species, and assignment of the proper geographical distribution to each, has been a subject which has sorely perplexed the ingenuity and patience of zoologists. The following description by Dr. Elliott Coues of the external characters of the American Pine Marten (J/. americana) will apply almost equally well to most of the others: “It is almost impossible to describe the colour of the Pine Marten, except in general terms, without going into the details of the endless diversities occasioned by age, sex, season, or other incidents.. The animal is ‘brown,’ of various shades from orange or tawny to quite blackish ; the tail and feet are ordinarily the darkest, the head lightest, often quite whitish ; the ears are usually rimmed with whitish; on the throat there is usually a large tawny-yellowish or orange-brown patch, from the chin to the fore legs, sometimes entire, sometimes broken into a number of smaller, irregular blotches, sometimes wanting, some- times prolonged on the whole under surface, when the animal is ? See Rolleston, ‘‘On the Domestic Cats, Felis domesticus and Mustela foina, of Ancient and Modern Times,” Jowrnal of Anatomy and Physiology, vol. ii. p. 47, 1868. MUSTELIDE 583 bicolor like a Stoat in summer. The general ‘ brown’ has a grayish cast, as far as the under fur is concerned, and is overlaid with rich lustrous blackish-brown in places where the long bristly hairs prevail. The claws are whitish ; the naked nose pad and whiskers are black. The tail occasionally shows interspersed white hairs, or a white tip.” The species generally recognised as distinct are the following, the first five belonging to the Old and the last two to the New World :— AL foiue, the Beech Marten, Stone Marten, or White-breasted Marten.—Distinguished from the following by the greater breadth of the skull, and some minute but constant dental characters, by Fic. 207,—The Pine Marten (Jlustela martes). the dull grayish-brown colour of the fur of the upper parts, and the pure white of the throat and breast. It inhabits the greater part of the continent of Europe, but is more southern than the next in its distribution, not being found in Sweden or Norway, nor, according to the investigations of Mr. Alston, in the British Isles, although included in their fauna by all earlier writers; to the eastward it ranges into Afghanistan and the Himalaya. AE. martes, the Pine Marten (Fig. 267).—Outer fur rich dark brown ; under fur reddish-gray, with clear yellow tips; breast spot usually yellow, varying from bright orange to pale cream-colour or yellowish-white. Length of head and body 16 to 18 inches; of tail (including the hair) 9 to 12 inches. This species is extensively distributed throughout northern Europe and Asia, and was formerly 584 CARNITVORA common in most parts of Great Britain and Treland, Though commonly called “ Pine Marten,” it does not appear to have any special preference for coniferous trees, except that, inasmuch as they constitute the greater proportion of the forests of the countries which it inhabits, it is more often met with in them than in any other. With regd to its recent oceurrence in the British Isles, Mr. Alston writes in the Prac, Zool, Soe, 1879 :— “Although greatly reduced in numbers by perseettion, it still maintains its ground in the wilder districts of Scothud, the north of England, Wales, and Ireland; and occasionally specimens are killed in countios where the species was thought to lave heen long oxtinet. In Seotland it is still found, though comparatively rarely, in the Lows and in most. of the Highland mainland counties, being perhaps most abundant in Sutherland and Ross shire, especially in the deer forests. In the Lowlands a Marten is now a very great rarity ; but a fino example was killed in Ayrshire in the winter of 1875-76. In the north of England Mr. W. A. Durnford says. the species is still plentiful in the wilder parts of Cumberland, West- moreland, and Laneashire, and in’ Lineolushire several have been recorded, the latest killed in 1865, by Mr. Cordeaux. In Norfolk one was shot last year; and [ have myself examined a fine oxumple which was shot in’ Elertfordshire, within 20 miles of London, in Decomber E872. In Dorsetshire the lust is said) to have been killed in 1804; but a specimen occurred in Lhanpshire about forty years ago, and another in Survey in VS47. 0 In Preland the following counties were enumerated by Thompson as habitats of thix species: Donogal, Londonderry, Antrim, Down, Armagh, Fermanagh, Longford, Galway, Tipperary, Cork, and Kerry. The Cal-crann is probably now a rarer animal in Ireland than it was when Thompson wrote; but it still exists ino various districts, especially in County Kerry, whenee the society has received several living examples ; and Professor A. Leith Adams states that it has been seen of late years even in county Dublin.” AL cibetling, the Sable (Geman, Zobel and Zebel Swedish, sibel. Tuassian, sobel, a word probably of Puranian origin).— Closely resembling the last, if indeed differing from it except in the quality of the fur, which is the most highly valued of that of all the group. Found chilly in Mastern Siberia. A flaviquia, the Indian Marten.—Inhabits the southern slopes of ,the Himalaya, the Nilgiri Hills, the interior of Ceylon, the Malay Peninsula, aud Java, ‘The coloration of this species is very striking, the upper parts being blackish-brown, and the throat: and breast yellow or orange, tn the bright coloured variety. 1b differs from the other species in having the soles of the feet more ov legs naked, AP. ielenpas.-— Japan. MUSTELIDA 585 AM. amerivanu, the North-American Sable or Marten.—A species so closely allied to the European Pine Marten and Asiatic Sable that it is very difficult to assign constant distinguishing characters between them. The importance of the fur of this animal as an article of commerce may be judged of from the fact that 15,000 skins were sold in one year by the Hudson's Bay Company as long ago as 1743, and the more recent annual imports into Great Britain have exceeded 100,000. It is ordinarily caught in wooden traps of very simple construction, being little enclosures of stakes or brush in which the bait is placed upon a trigger, with a short upright stick supporting a log of wood, which falls upon its victim on the slightest disturbance. A line of such traps, several to a mile, often extends many miles. The bait is any kind of meat, a mouse, squirrel, piece of fish, or bird’s head. It is principally trapped during the colder months, fron: October to April, when the fur is in good condition, us it is nearly valueless during the shedding in summer. Dr. Coues tells us that, notwithstanding the persistent and uninterrupted destruction to which the American Sable is subjected, it docs not appear to diminish materially in numbers in unsettled parts of the country. It holds its own partly in conse- quence of its shyness, which keeps it away from the abodes of men, and partly becanse it is so prolific, bringing forth six to eight young ata litter, Its home is sometimes a den under ground or beneath rocks, but oftencr the hollow of « tree, and it is said frequently to take forcible possession of a squirrel’s nest, driving off or devouring the rightful proprictor, A. pennanti, the Pekan ov Pennant’s Marten, also called Fisher Marten, though there appears to be nothing in its habits to justify the appellation.—This is the largest species of the group, the head and body measuring from 24 to 30 inches, and the tail 14 to 18 inches, It is also more robust in form than the others, its general aspect being more that of a Pox thin a Weasel ; in fact, its usual name among the American hunters is “ Black Fox.” Its general colour is blackish, lighter by mixture of brown or gray on the head and upper fore part of the body, with no light patch on the throat, and unlike the other Martens generally darker below than above. Tt was generally distributed in wooded districts throughout the greater part of North America, as far north as Great Slave Lake, G3° N. lat. and Alaska, and extending south to the parallel of 35°; Int at the present time it is almost exterminated in the settled parts of the United States cast of the Mississippi. Fossil remains of «a Marten from the Pliocene Siwaliks of India indicate w species which cannot be distinguished from those now inhabiting the same region; while remains of JZ. martes occur in European cavern-deposits, ind in the fens of Cambridgeshire. With the Puforiine group (genus Pulorins) we come to those 386 CARNIVORA smaller forms distinguished by having only three premolars in each jaw, by the absence of an inner cusp to the blade of the lower carnassial, as well as by certain external characters. This group contains a few species known as Minks, differing from the rest by slight structural modifications, and especially by their semiaquatic habits. They are distinguished from the Polecats, Stoats, and Weasels, which constitute the remainder of the group, by the facial part of the skull being narrower and more approaching in form that of the Martens, by the premolar teeth (especially the anterior one in the upper jaw) being larger, by the toes being partially webbed, and by the absence of hair in the intervals between the naked pads of the soles of the feet. The two best-known species, so much alike in size, form, colour, and habits that although they are widely separated geographically some zoologists question their specific distinction, are M. lutreola, the Nérz or Sumpfotter (Marsh- Otter) of Eastern Europe, and M. vison, the Mink of North America. The former inhabits Finland, Poland, and the greater part of Russia, though not found east of the Ural Mountains. Formerly it extended westward into Central Germany, but it is now very rare, if not extinct, in that country. The latter is found in places which suit its habits throughout the whole of North America. Another form, Jf. sibirica, from Eastern Asia, of which much less is known, appears to connect the true Minks with the Polecats. For the following description, chiefly taken from the American form (though almost equally applicable to that of Europe), we are mainly indebted to Dr. Coues’s Fur-bearing Animals of North America. In size it much resembles the English Polecat,—the length of the head and body being usually from 15 to 18 inches, that of the tail to the end of the hair about 9 inches. The female is consider- ably smaller than the male. The tail is bushy, but tapering at the end. The ears are small, low, rounded, and scarcely project beyond the adjacent fur. The pellage consists of a dense, soft, matted under fur, mixed with long, stiff, lustrous hairs on all parts of the body and tail. The gloss is greatest on the upper parts; on the tail the bristly hairs predominate. Northern specimens have the finest and most glistening pellage ; in those from southern regions there is less difference between the under and over fur, and the whole pellage is coarser and harsher. In colour different specimens present a - considerable range of variation, but the animal is ordinarily of a rich dark brown, scarcely or not paler below than on the general upper parts ; but the back is usually the darkest, and the tail is nearly black. The under jaw, from the chin about as far back as the angle of the mouth, is generally white. In the European Mink the upper lip is also white, but as this occasionally occurs in American speci- mens it fails as an absolutely distinguishing character. Besides the white on the chin, there are often other irregular white patches MUSTELIN.# 587 on the under parts of the body. In very rare instances the tail is tipped with white. The fur, like that of most of the animals of the group to which it belongs, is an important article of commerce. The principal characteristic of the Mink in comparison with its congeners is its amphibious mode of life. It is to the water what the other Weasels are to the land, or Martens to the trees, being as essentially aquatic in its habits as the Otter, Beaver, or Musk-Rat, and spending perhaps more of its time in the water than it does on land. It swims with most of the body submerged, and dives with perfect ease, remaining long without coming to the surface to breathe. It makes its nest in burrows in the banks of streams, breeding once a year about the month of April, and producing five or six young at a birth. Its food consists of frogs, fish, freshwater Fig. 268.—The Common Poleeat (Mustela putorins). molluses and crustaceans, as well as mice, rats, musk-rats, rabbits, and small birds. In common with the other animals of the genus, it has a very peculiar and disagreeable effluvium, which, according to Cones, is more powerful, penetrating, and lasting than that of any animal of the country except the Skunk. It also possesses the courage, ferocity, and tenacity of life of its allies. When taken young, however, it ean be readily tamed, and lately Minks have heen extensively bred in captivity in America, both for the sake of their fur and for the purpose of using them in like manner as Ferrets in England, to clear buildings of rats. The Polecats include four species confined to the northern hemisphere, the best known of which is the Common Polecat (IL putorius, Fig. 268). The Ferret is a domesticated variety of this species, generally of a yellowish-white colour ; whereas the Wild 588 CARNIVORA Polecat is dark brown above and black beneath, the face being variegated with dark brown and white markings. The skull is rough, strongly ridged, and of a far more powerful type than that of the Stoats, Weasels, or Martens; being in the female much smaller and lighter than in the male. The fur, which is long, coarse, and of compiwatively small value, changes its colour very little, if at all, at the different seasons of the year. The distribution and habits of this species have been described by Blasins, the following being an abstract of his account. The Polecat ranges over the greater part of Europe, reaching northwards into Southern Sweden, and in Russia to the region of the White Sea. It does not occur in the extreme South, but is common eyery- where throughout Central Europe. In the Alps it ranges far above the tree-line during the summer, but retreats in winter to lower ground. In fine weather it lives cither in the open air, in holes, fox-earths, rabbit-warrens, under rocks, ov in wood-stacks, while in winter it seeks the protection of deserted buildings. During the day it sleeps in its hiding-place, sallying forth at night to plunder dovecots and hen-houses. It climbs but little, and shows far less activity than the Marten. It feeds ordinarily on small mammals, such as rabbits, hamsters, rats, and mice, on such hirds as it can catch, especially poultry and pigeons, and also on snakes, lizards, frogs, fish, and eggs. Its prey is devoured only in its lair, but, even though it can carry away but a single victim, it commonly kills everything that comes in its way, often destroying all the inhabitants of a hen-house in order to gratify its passion for slaughter. The pairing time is towards the end of the winter, and the young, from three to eight in number, are born in April or May, after a period of gestation of about two months. The young, if taken carly, may be easily trained, like Ferrets, for rabbit-catehing. The Polecat is very tenacious of life, and will hear many severe wounds before succumbing ; it is also said to receive with impunity the bite of the adder. Its fetid smell has hecome proverbial. Four other species of Polecats are known, viz.—The Siberian Polecat (AL erersimennt) of Western and Northern Asia is nearly allied to the European species, but the head and back are almost white, and the skull is stouter and more constricted behind the orbits) The Tibetan Jf larrata is distinguished from the last by the presence of a process connecting the pterygoid with the auditory bulla, and hy a difference in the shape of the upper molar. The American Polecat (MZ. vigripes), inhabiting the central plateau of the United States, and extending southwards into ‘Texas, is another closely allied species, although some zoolovists have made it the type of the genus Cynomyouar, Finally, the Mottled Poleeat (AL. surmatica) is a species sparsely distributed in’ Eastern Europe and parts of Western Asia, but common in Southern Afghanistan. WUSTELIDE 589 Its skull, although smaller, resembles that of the common species : but the coloration is very different, all the upper part: baie mottled with large irregular reddish spots on a white ground, and the under side, limbs, and tail deep shining black. The tail is long. The Common Polecat occurs in a fossil condition in the cave- deposits of Europe. The remaining members of the genus comprise the true Weasels and Stoats, which are of almost cosmopolitan distribution. In the Common Weasel (Jf rulgaris, Fig. 269) the upper parts, outside of limbs and tail, are a uniform reddish-brown, the under parts pure Fic, 209.—The Common Weasel (Mustela ruljuris). white. In very cold regions, both in Europe and America, it turn: completely white in winter, but less regularly and at a lower temperature than the Stoat, from which it is easily distinguished br its smaller size, and by its wanting the black end of the tail. The length of the head and body of the male is usually about 3 inches, that of the tail 2! inches: the female is smaller. This species is pretty generally distributed throughout Europe, Northern and Central Asia, British North America, and the northern portions of the United States. It possesses in a full degree all the active, courageous, and bloodthirsty disposition ot the rest of the genus, but its diminutive size prevents it attacking and destroying any but the smaller mammals and birds. Mice, rats, voles. moles, and frogs constitute its principal food It is generally found on or 590 CARNIVORA near the surface of the ground, but it can not only pursue its prey through very small holes and crevices of rocks and under dense tangled herbage, but follow it up the stems and branches of trees, or even into the water, swimming with perfect ease. It constructs a nest of dried leaves and herbage, placed in a hole in the ground or a bank or hollow tree, in which it brings up its litter of four to six (usually five) young ones. The mother will defend her young with the utmost desperation against any assailant, having been often known to sacrifice her own life rather than desert them. The Stoat or Ermine (J/. erminea) has nearly the same distribu- tion as the Weasel, but in Asia it is said to extend into parts of the Kashmir Himalaya. Its size, as already mentioned, consider- ably exceeds that of the Weasel ; and its most distinctive feature is the black tip at the end of the tail, which remains when the rest of the pellage turns white. The white winter skins from the northern regions of its habitat, where the fur is thick and close, form the well-known and valuable ermine of commerce. Remains of the Stoat are found in the Pleistocene cavern-deposits of Europe. The other species of Weasels are very numerous and widely distributed. Extinct Mustelines—A number of European Miocene Carnivores may be referred to the genus Jfustela in its wider sense, and serve to confirm the propriety of this use of the term. Thus JL. sectoria is a species of somewhat larger size than the Stoat, with p +, while in AL. angustifrons the number of premolars is 2, and in J. mustelina 4; the latter species agreeing very closely in size with the Stoat. The extinct Plesictis, in which there are p + and the lower car- nassial has a large inner cusp, is distinguished from J/ustela by the circumstance that the temporal ridges of the skull never unite to form a sagittal crest. Moreover, the inner tubercular portion of the upper molar (as in some of the Miocene species of J/ustelw) is shorter in an antero-posterior direction than the secant outer moiety; and the auditory bulla is more inflated than in A/ustela, although it has no septum. Both these features indicate a decided approximation to the Viverroid genus Stenoplesiotis (p. 539); and since there are no well-marked characters of family value by which these two genera can be distinguished the available evidence points to a transition from the Viverroid to the Musteloid type. Austela lurteti, of the Middle Miocene of France, should perhaps be referred to Letonye. Peecilogale.A—This genus has been made for the reception of the South African Mustela albinucha, in which the coloration is similar to that of Ictonyz, but the number of cheek-teeth is usually reduced to pg, m4, although there may be a second lower molar. The auditory bulla is quite flat. Lyncodon.?—This name has been proposed for a small Musteline 1 0. Thomas, Ann. Mag. Nat. Hist. ser. 5, vol. xi. p. 870 (1883). > Gervais, Dict. Univ. d’ Hist. Nat. t. iv. p, 685 (1849). AMLUSTELIDA 591 from Patagonia, with p 2, m +, which Mr. O. Thomas suggests may be nothing more than an aberrant southern form of Mustela (Putorius) brasiliensis. The auditory bulla is more inflated than in the typical Weasels. This animal is somewhat larger than the Stoat. Gulo.\—Dentition: 1 3,¢ +, p 4, m 4; total 38. Crowns of the teeth very stout. Upper molar very much smaller than the car- nassial. Lower carnassial large, with very small talon and no inner cusp. Third upper incisor unusually large, almost like a canine. The dentition, though really but a modification of that of the Weasels, presents a great general resemblance to that of the Hyena. Palate prolonged somewhat behind the last molar. Humerus with an ente- picondylar foramen. Vertebre: C 7,D 15, L 5,8 3,C15. Body Fic. 270.—The Wolverene (Gulo luscus). and limbs stoutly made. Feet large and powerful, subplantigrade, with large, compressed, much curved, and sharp-pointed claws. Soles of the feet (except the pads of the toes) covered with thick bristly hairs. Ears very small, nearly concealed by the fur. Eyes small. Tail short, thick, and bushy. Fur full, long, and rather coarse. The one species, the Wolverene or Glutton (G. luscus, Fig. 270), an inhabitant of the forest regions of Northern Europe, Asia, and America, much resembles a small Bear in appearance. It is a very powerful animal for its size, climbs trees, and lives on grouse, squirrels, hares, foxes, beavers, reindeer, and is said to attack even horses and cows. The Wolverene has a curious habit of stealing and secreting articles of which it can make no possible use, as is exemplified in the following instance related by Dr. Coues: 1 Storr, Prodromus Meth. Mamm. p. 34 (1780). 592 CARNIVORA “A hunter and his family, having left their lodge unguarded during their absence, on their return found it completely gutted— the walls were there, but nothing else. Blankets, guns, kettles, axes, cans, knives, and all the other paraphernalia of a trapper’s tent had vanished, and the tracks left by the beast showed who had been the thief. The family set to work, and, by carefully following up all his paths, recovered, with some trifling exceptions, the whole of the lost property.” The pairing season occurs in March, and the female, secure in her burrow, produces her young, four or five at a birth, in June or July. In defence of these she is exceedingly bold, and the Indians, according to Coues, “have been heard to say that they would sooner encounter a she-bear with her cubs than a carcajou (the Indian name of the glutton) under the same circumstances.” Fossil remains of the Wolverene are found in cavern and other Pleistocene deposits in various parts of Europe. Suborder PINNIPEDIA. The Eared-Seals, Walruses, and Seals differ from the rest of the Carnivora mainly in the structure of their limbs, which are modified for aquatic progression,—the two proximal segments being very short and partially enveloped in the general integument of the body; while the third segment, especially in the hinder extremities, is elongated, expanded, and webbed. There are always five well- developed digits on each limb. In the hind limb the two marginal digits (first and fifth) are stouter and generally longer than the others. The teeth also differ from those of the more typical Carnivora. The incisors are always fewer than 3. The cheek series consists generally of four premolars and one molar of very uniform characters, with never more than two roots, and with conical, more or less compressed, pointed crowns, which may have accessory cusps, placed before or behind the principal one, but are never broad and tuberculated ; and there is no differentiated carnassial tooth. The milk-teeth are very small and simple, and are shed or absorbed at a very early age, usually either before or within a few days after birth. The brain is relatively large; the cerebral hemispheres being broad in proportion to their length, with numerous and complex convolutions. There is a very short cecum. The kidneys are divided into numerous distinct lobules. There are no Cowper’s glands. The mammz are either two or four, and abdominal in position. No clavicles. Tail always very short. Eyes very large and exposed, with flat cornea. The animals of this group are all aquatic in their mode of life, spending the greater part of their time in the water, swimming and diving with great facility, feeding mainly on fish, crustaceans, and other marine animals, and progressing on land with difficulty. OTARIID.LE 593 They always come on shore, however, for the purpose of bringing forth their young. They are generally marine, but they occasion- ally ascend large rivers, and some inhabit inland seas and lakes, as the Caspian and Baikal. Though not numerous in species, they are widely distributed over the world, but occur most abundantly on the coasts of lands situated in cold and temperate zones. The suborder is divisible into three well-marked families: the Ofuriidi, Fur-Seals or Sea-Bears, which form a transition from the Fissiped Carnivora to the Seals ; the Zrichechidw, containing the Walrus ; and the Phocide or typical Seals. The resemblances between the skull and other parts of the body of the Fur-Seals and the Ursoid true Carnivora is suggestive of some genetic relationship between the two groups, and Pro- fessor Mivart! expresses the opinion that the one group is the direct descendant of the other. The same writer goes on to suggest that if the Eared-Seals have been derived from Bear-like Carnivores this need not necessarily hold good with the true Seals, which may have had another, and possibly Lutrine, origin. The presence of an alisphenoid canal in Ursus and the Ofariide, and its absence in Lutru and the Phocidw, together with other osteological features, are cited in support of this view; but although these resemblances and differences are certainly remarkable, yet much more evidence is required before the hypothesis can be accepted as even a probable one. It must, moreover, be borne in mind that the true Bears are a very modern group; and there is a possibility that the Pinnipeds may prove to have been independently derived from the extinct Carnivora noticed below under the name of Creodonta. Family OTARUD. When on land the hind feet are turned forwards under the body, and aid in supporting and moving the trunk as in ordinary mammals. A small external ear. Testes suspended in a distinct external scrotum. Skull with postorbital processes, and an alisphenoid canal. Angle of mandible inflected. Palms and soles of feet naked. Otaria.2—Dentition: i 3, ¢ 1, p 4, m cele total 34 or 36. First and second upper incisors small, with the summits of the crowns divided by a deep transverse groove into an anterior and a posterior cusp of nearly equal height; the third large and canine-like. Canines large, conical, pointed, recurved. Molars and premolars usually 4, of which the second, third, and fourth are preceded by milk-teeth shed a few days after birth: sometimes (as in Fig. 271) a sixth upper molar (occasionally developed on one l Proce. Zool. Sov. 1885, p. 497. > Péron, Mouage aux Terres Australes, vol. ii. y. 37 note (1816). 38 594 CARNIVORS! side and not the other); all with similar characters, venerally uniradicular ; crown moderate, compressed, pointed, with a single principal cusp, and sometimes a cingulum, and more or less de- velopedanterior and — posterior ACCESSOTY CUSPS. Vertebra: C7, 1+15, b 5.84, C 9-14. Head rounded. Eyes large, Pinna of cur small, narrow, and pointed. Neck : 5 lone. Skin of Mia, 271.—Skull of Otaria forsteri. (From Gray, Pree. Zool. Soc. & 1872, p. 660.) all the feet ex- tended far be- yond the nails and ends of the digits, with a deeply-lobed margin. The nails small and often quite rudimentary, especially those of the first and fifth toes of both feet, the best-developed and most constant being the three middle claws of the hind foot, which are clongated, compressed, and curved. The Eared-Seals, commonly called Sea-Bears or Nea-Lions, are widely distributed, especially in the temperate regions of both hemispheres, though absent from the coasts of the North Atlantic. As might be inferred from their power of walking on all fours, they spend more of their time on shore, and range inland to greater distances, than the true Seals, especially at the breeding time, though they are obliged always to return to the water to seck their food. They are gregarious and polygamous, and the males are usually much larger than the females, a circumstance which has given rise to some of the confusion existing in the specific deter- mination of the various members of the genus. Some of the species possess, in addition to the stiff, close, hairy covering common to all the group, an exceedingly fine, dense, woolly under fur. The skins of these, when dressed and deprived of the longer harsh outer hairs, constitute the “sealskin” of commerce, so much valued for wearing apparel, which is not the product of any of the true Neals. The best-known species are (@. sfe/lcri, the Northern Sex Lion, the largest of the genus, from the North Pacilic, about 10 fect in length ; 0. jubata, the Patagonian or Southern Nea Lion (Fig. 272), from the Falkland Islands and Patagonia; 0. culiforniana, from California, frequently exhibited alive in menagerics in Europe ; O. wesind, the common Sea-Bear or Fur-Neal of the North Pacifie, the skins of which are imported in immense numbers from the Prybiloff Islands ; 0. pusilla, from the Cape of Good Hope; 0. forsteri and OTARUDAE 595 others, from the coasts of Australia and various islands scattered over the southern hemisphere. These have been grouped by some voologists into many genera, founded upon very trivial modifica- tions of teeth and skull In a recent memoir Mr. Beddard ! con- cludes that if the genus be split wp at all, it should be divided into Otaria, containing only OQ, jubufa (vith its numerous synonyms), and -trelocephalus, comprising all the other species. ‘The latter group is distinguished by the more narrow and pointed nose, the longer ears, the palate not excavated nor truncated posteriorly, the presence of Mia. 272.0 The Patagonian Sea-Lion (frie pubata). From Selater, Prec. Zool, Suc, W860, p. SO. a hook-like process to the pterygoids, and by the posterior border of the nasals extending behind the zyvgoma, Tho following account of GO. wsdee in the Peybilott [slands is taken, with slight verbal alteration, from Nordenskiold’s /oyuge of the Peyas * The Sea-Bears are found year after year during summer at certain parts of the coast, known as *rookeries,” where, collected in hundreds of thousands, they pass several months without the least food. ‘The males or * bulls’ come tirst to the place, most of them in the month of May or in the beginning of June. The most violent conthets, often with a deadly issue for one of the parties, now atise regarding the space of about a hundred square fect whieh each bull-seal considers necessary for his home. The hssOn the structure of Hooker's Sea-Lion (lrefocephalus hookerd.” Trans. Zool, Soe Vol. xt, p. 809 (1890). 396 CARNIVORA strongest and most successful in fight retain the best places near the shore: the weaker have to crawl] farther up on land, where the chances of getting a sufficient number of spouses are not particularly great. The fighting goes on with many feigned attacks and parades. At first the contest concerns only the proprietorship of the soil. The attacked, therefore, never follows his opponent beyond the area he has once taken up, but haughtily lays itself down, when the enemy has retired, in order to collect strength for a new combat. The animal in such a case grunts with satisfaction, throws himself on his back, scratches himself with his fore feet, attends to his toilet, or cools himself by slowly fanning with one of his hind feet : but he is always on the alert and ready for a new fight, until he is tired out and meets his match and is driven farther up from the beach. In the middle of June the females come up from the sea. At the waters edge they are received in a very gallant way by some strong bulls that have succeeded in securing for themselves places next the shore, and now are bent by fair means or foul on annexing the females for their harem. But searcely is the female that has come up out of the water established with male No. 1 than he rushes towards a new female on the surface of the water. Male No. 2 now stretches out his neck and without ceremony lays hold of the female of No. 1, to be afterwards exposed to a similar trick by No. 3. In such cases the females are quite passive, never fall out with each other, and bear with patience the severe wounds they often get when they are pulled about by the combatants, now in one direction, now in another. All the females are finally dis tributed in this way after furious combats among the males, those of the latter who are nearest the beach getting from 12 to 15 consorts to their share. Soon after landing the females bring forth their young, which are treated with great indifference, and are protected by their adopted father only within the limits of the harem. Next comes the pairing season, and when it has passed there is an end to the arrangement and distribution into families at first so strictly maintained. The males, rendered lean by three months’ absolute fasting, by degrees leave the rookery, which is left in possession of the Walruses and the young Sea Bears, including a number of young males that have not ventured to the place before. In the middle of September, when the young have learned to swim, the place is quite abandoned, with the exception of single animals that have for some reason remained behind.” Family TRICHECHID-. In many characters the single genus representing this family is intermediate between the Otariide and Phocide, but it has a completely aberrant dentition. It has no external ears, as in the TRICHECHID A: 597 Phocide ; but when on land the hind feet are turned forwards and used in progression, though less completely than in the Otartidd. The upper canines are developed into immense tusks, which descend wu long distance below the lower jaw. All the other teeth (Fig. 273), including the lower canines, are much alike, small, simple, and one-rooted, the molars with flat crowns. The skull is without postorbital process, but has an alisphenoid canal. Trichechus.A—Dentition of young: i 3, ¢+, p andm 4. Many of these teeth are, however, lost early or remain through life in a rudimentary state concealed by the gums. The teeth which are usually developed functionally are i 3, ¢ 1, p 3, m %; total 18. Vertebrw: C7, D 14,L 6,8 4,0 12. Head round, Eyes rather small. Muzzle short and broad, with on each side a group of long, Via, 278,—Diagram of the dentition of the Walrus (Trickcehus rosmarus), The denticles placed apart from the others are milk-teeth, and disappear soon after birth. The small teeth in connection with the jaws frequently persist throughout life. very stiff, bristly whiskers. The remainder of the hair-covering very short and adpressed. Tail very rudimentary. Fore feet with subequal toes, carrying five minute flattened nails. Hind feet with subequal toes, the fifth slightly the largest, having cutaneous lobes projecting beyond the ends as in Oturiv , first and fifth with minute flattened nails; second, third, and fourth with large, elongated, subcompressed pointed nails. Trichechus is the almost universally accepted generic name by which the Walrus or Morse? is known to zoologists, but some con- fusion has been introduced into literature by the revival of the nearly obsolete terms /tosmarns by some authors and Odelenus ly others. 7. rosmearus is the name of the species met with in the Arctic seas ; that of the North Pacific, if distinct, is 7. obesus. The preceding and following descriptions will apply equally to both. 1 Linn, Sys, Nat, 12th ed, vol. i. p. 49 (1766). * The former word is a modification of the Scandinavian vallross or hvalros (‘whale-horse’’), the latter an adaptation of the Russian name for the animal. 598 CARNIVORA A full-grown male Walrus measures from 10 to 11 feet from the nose to the end of the very short tail, and is a heavy, bulky animal, especially thick about the shoulders. The soles of both fore and hind feet are bare, rough, and warty. The surface of the skin generally is covered with short, adpressed hair of a light, yellowish- brown colour, which, on the under parts of the body and base of the flippers, passes into dark reddish-brown or chestnut. In old animals the hair becomes more scanty, sometimes almost entirely disappearing, and the skin shows ample evidence of the rough life and pugnacious habits of the animal in the innumerable scars with which it is usually covered. Itis everywhere more or less wrinkled, Fic. 274.—The Walrus (Lrichechus rosmarus). but especially over the shoulders, where it is thrown into deep and heavy folds. The tusks are formidable weapons of defence, but their principal use seems to be scraping and digging among the sand and shingle for the molluscs and crustaceans on which the Walrus feeds. They are said also to aid in climbing up the slippery rocks and ledges of ice on which so much of the animal's life is passed. Although this function of the tusks is affirmed by numerous authors, some of whom appear to have had opportunities of actual observation, it is explicitly denied by Malmeren. Walruses are more or less gregarious in their habits, being met with generally in companies or herds of various sizes. They are only found near the coast or on large masses of floating ice, and rarely far out in the open sea ; and, though often moving from one part of their feeding ground to another, they have no regular seasonal migrations. Their young are born between the months of TRICHECHIDA 599 April and June, usually but one at a time, never more than two. Their strong affection for their young, and their sympathy for each other in times of danger, have been particularly noticed by all who have had the opportunity of observing them in their native haunts. When one of their number is wounded, the whole herd usually join in a concerted and intelligent defence. Although harmless and inoffensive when not molested, they exhibit considerable fierceness when attacked, using their great tusks with tremendous effect either on human enemies who come into too close quarters or on Polar Bears, the only other adversaries they can meet with in their own natural territory. Their voice is a loud roaring, and can be heard at a great distance; it is described by Dr. Kane as “some- thing between the mooing of a cow and the deepest baying of a mastiff, very round and full, with its bark or detached notes repeated rather quickly seven or nine times in succession.” The principal food of the Walrus consists of bivalved molluscs, especially Mya truncata and Sawxicava rugosa, two species very abundant in the Arctic regions, which it digs up from the mud and sand in which they lie buried at the bottom of the sea by means of its tusks. It crushes and removes the shells by the aid of its grinding teeth and tongue, swallowing only the soft part of the animal. It also feeds on other molluscs, sand-worms, star-fishes, and shrimps. Portions of various kinds of alge or sea-weeds have been found in its stomach, but whether swallowed intentionally or not is still doubtful. The commercial products of the Walrus are its oil, hide (used to manufacture harness and sole-leather and twisted into tiller ropes), and tusks. The ivory of the latter is, however, inferior in quality to that of the Elephant. Its flesh forms an important article of food to the Eskimo and Tchuktchis. Of the coast tribes of the last-named people the Walrus forms the chief means of support. “The flesh supplies them with food, the ivory tusks are made into implements used in the chase and for other domestic purposes, as well as affording a valuable article of barter, and the skin furnishes the material for covering their summer habitations, harness for their dog-teams, and lines for their fishing gear ” (Scammon). Geographically the Walrus is confined to the northern circum- polar regions of the globe, extending apparently as far north as explorers have penetrated, but its southern range has been much restricted of late in consequence of the persecutions of man. On the Atlantic coast of America it was met with in the sixteenth century as low as the southern coast of Nova Scotia, and in the last century it was common in the Gulf of St. Lawrence and on the shores of Labrador. It still inhabits the coast round Hudson’s Bay, Davis Straits, and Greenland, where, however, its numbers are daily decreasing. It is not found on the Arctic coast of America between the 97th and 600 CARNIVORA 158th meridians. In Europe occasional stragglers have reached the British Isles, and it was formerly abundant on the coasts of Finmark. It is rare in Iceland, but Spitzbergen, Nova Zembla, and the western part of the north coast of Siberia are still constant places of resort, in all of which a regular war of extermination 1s carried on. The North Pacific, including both sides of Behring’s Strait, northern Kamschatka, Alaska, and the Pribyloff Islands, are also the haunts of numerous Walruses, which are isolated from those of the North Atlantic by the long stretches of coast, both of Siberia and North America, where they do not occur. The Pacific Walrus appears to be as large as, if not larger than, that of the Atlantic; its tusks are longer and more slender, and curved inwards ; the whiskers are smaller, and the muzzle (of the skull) relatively deeper and broader. These and certain other minor differences have induced some naturalists to consider it specifically distinct under the name of Trichechus obesus. Its habits appear to be quite similar to those of the Atlantic form. Though formerly found in immense herds, it is rapidly becoming scarce, as the methods of destruction used by the American whalers, who have systematically entered upon its pursuit, are far more certain and deadly than those of the native Tchuktchis, to whom, as mentioned before, the Walrus long afforded the principal means of subsistence. Fossil remains of Walruses and closely allied animals have been found in the United States, and in England, Belgium, and France, in deposits of Pliocene age. Family PHocrp-®. The true Seals are the most completely adapted for aquatic life of all. the Pinnipeds, When on land the hind limbs are extended behind them and take no part in progression, which is effected by a series of jumping movements produced by the muscles of the trunk, in some species aided by the fore limbs only. The palms and soles of the feet are hairy. There is no pinna to the ear, and no scrotum, the testes being abdominal. The upper incisors have simple, pointed crowns, and vary in number in the different groups. All the forms have well-developed canines and 2 teeth of the cheek- series. In those species of which the milk-dentition is known, there are three milk molars (Fig. 275), which precede the second, third, and fourth permanent molars ; the dentition is therefore pi, m +, the first premolar having as usual no milk-predecessor. ‘The skull has no postorbital process and no alisphenoid canal; and the angle of the mandible is not inflected. The fur is stiff and adpressed, without woolly under fur. Subfamily Phoeinze.—Incisors 3. All the feet with five well- developed claws. The toes on the hind feet subequal, the first and PHOCIDE 601 fifth not greatly exceeding the others in length, and with the interdigital membrane not extending beyond the toes. Halicheerus.\—Dentition : i 3,¢4, p 4,m 4; total 34. Crowns of molars large, simple, conical, recurved, slightly compressed, OO Vy Fic. 275.—Upper permanent and deciduous dentition of the Greenland Seal (Phoce, granlandica). The first and second deciduous incisors are already absorbed. with sharp anterior and posterior edges, but without accessory cusps, except sometimes in the two hinder ones of the lower jaw. With the exception of the last one or two in the upper jaw and the last in the lower jaw they are all uniradicular. Vertebre: C 7,D15,L5,8 4, C 14. One species, H. grypus, the Gray Seal of the coasts of Scandinavia and the British Isles (see page 604.) Phoca.2—Dental formula as the last. Teeth smaller and more pointed. Molars (Figs. 275 and 276) with two roots (except the first in each jaw); and their crowns with accessory cusps. Vertebre: C 7, D 15, L 5, 8 4, C 12-15. Head round and_ short. Fore feet short, with five very strong, subcom- pressed, slightly curved, rather sharp claws, sub- Fig. 276.—Skull of Common Seal, showing form of teeth. equal in length. On the hind feet the claws much narrower and less curved. The species of this genus are widely distributed throughout the northern hemisphere, and include P. barbata, the Bearded Seal; P. gran- landica, the Greenland Seal; P. vitulina, the Common Seal (Fig. 277); and P. hispida, the Ringed Seal of the North Atlantic ; P. caspica, from the Caspian and Aral Seas; and P. sibirica, from Lake Baikal. 1 Nilsson, Faun. Scandinav. vol. i. p. 377 (1820). * Linn. Syst. Nat. 12th ed. vol. i. p. 55 (1766). 602 CARNITORA Although the members of this subfamily swim and dive with the greatest ease, often remaining as much as a quarter of an hour or more below the surface, and are dependent for their sustenance entirely on living prey captured in the water. vet they frequently resort to sandy beaches, rocks, or ice-floes, either to sleep or to bask in the sun, and especially for the purpose of bringing forth their young. The latter appears to be the universal habit, and, strange as it may seem, the young seals—of some species at least—take to the water at first very reluctantly, and have actually to be taught to swim by their parents. The number of young produced is usually one annually, though occasionally two. They are at first covered with a coat of very thick, soft, nearly white fur, and until it falls off they do not usually enter the water. This occurs in the Greenland and Gray Seal when from two to three weeks old, but in the Common Seal apparently much earlier. One of this species born in the London Zoological Gardens had shed its infantile woolly coat and was swimming and diving about in its pond within three hours after its birth. The movements of the true Seals upon the ground or ice are very different from those of the Eared-Seals. Thus the hinder limbs (by which mainly they propel themselves through the water) are on land always perfectly passive, stretched backwards, with the soles of the feet applied to each other, and often raised to avoid contact with the ground. Sometimes the fore limbs are equally passive, being placed close to the sides of the body, and motion is then effected by a shuffling or wriggling action produced by the muscles of the trunk. When, however, there is any necessity for a more rapid mode of progression the animals use the fore paws. either alternately or simultaneously, pressing the palmar surface on the ground and lifting and dragging the body forwards in a succession of short jumps. In this way they manage to move so fast that a man has to step out beyond a walk to keep up with them; but such rapid action costs considerable effort, and they very soon become heated and exhausted. These various modes of progression appear to be common to all species so far as has been observed. Most kinds of Seals are gregarious and congregate, especially at the breeding season, in immense herds. Such is the habit of the Greenland Seal (Phocu grwnlundica), which resorts in the spring to the ice-floes of the North Sea, around Jan Mayen Island, where about 200,000 are killed annually by the crews of the Seoteh, Dutch, and Norwegian sealing vessels. Others, like the Common Seal of the British islands (P. vitulina), though having a wide geographical range, are never met with in such large numbers or far away from land. This species is stationary all the vear round, but some have a regular season of migration, moving south in PHOCIDAY 603 winter and north in summer. They are usually harmless, timid, inoffensive animals, though, being polygamous, the old males often fight desperately with each other, their skins being frequently found covered with wounds and scars. They are greatly attached to their young, and remarkably docile and easily trained when in captivity ; indeed, although there would seem little in the structure or habits of the Seal to fit it by nature to be a companion of man, yet there is perhaps no wild animal which attaches itself so readily to the person who takes care of and feeds it. Seals appear to have much curiosity, and it is a very old and apparently well- Pia. 277,—The Common Seal (Phoee vituline). attested observation that they are strongly attracted by musical sounds. Their sense of smell is very acute, and their voice varies from a harsh bark or grunt to a plaintive bleat. Seals feed chiefly on fish, of which they consume enormous quantities ; some, how- ever, subsist largely on crustaceans, especially species of Giannis, which swarm in the northern seas, also on molluscs, echinoderms, and even occasionally sea-birds, which they seize when swimming or floating on the water. Although the true Seals do not possess the beautiful under fur (‘‘seal-skin” of the furriers) which makes the skin of the Nea-Bears so precious, yet their hides are still sufficiently valuable as articles of commerce, together with the oil yielded by their fat, to subject them to a devastating persecution, by which their numbers are being continually diminished. 604 CARNIVORA Two species of seals only are met with regularly on the British coasts, the Common Seal and the Gray Seal. The former (Fig. 277) is a constant resident in all suitable localities round the Scottish, Irish, and English coasts, from which it has not been driven away by the molestations of man. Although, naturally, the most secluded and out-of-the-way spots are selected as their habitual dwelling-places, there are few localities where they may not be occasionally met with. Within the writers’ knowledge one was seen not many years ago lying on the shingly beach at so populous a place as Brighton, and another was caught in the river Welland, near Stamford, 30 miles from the sea. They frequent bays, inlets, and estuaries, and are often seen on sandbanks or mudflats left dry at low tide, and, unlike some of their congeners, are not found on the ice-floes of the open sea, nor, though gregarious, are very large numbers ever seen in one spot. The young are produced at the end of May or beginning of June. They feed chiefly on fish, and the destruction they occasion among salmon is well known to Scottish fishermen. The Common Seal is widely distributed, being found not only on the European and American coasts bordering the Atlantic Ocean, but also in the North Pacific. It is from 4 to 5 feet in length, and variable in colour, though usually yellowish-gray, with irregular spots of dark brown or black above and yellowish-white beneath. The Gray Seal (Halichwrus grypus) is of considerably larger size, the males attaining when fully adult a length of 8 feet from nose to end of hind feet. It is of a yellowish-gray colour, lighter beneath, and with dark gray spots or blotches, but, like most other Seals, is liable to great variations of colour according to age. This species appears to be restricted to the North Atlantic, having been rarely seen on the American coasts, but not farther south than Nova Scotia ; it is chiefly met with on the coasts of Ireland, England, Scotland, Norway, and Sweden, including the Baltic and Gulf of Bothnia, and Iceland, though it does not appear to range farther north. It is apparently not migratory, and its favourite breeding places are rocky islands; the young being born in the end of September or beginning of October. Subfamily Monaehinze.—Incisors 2. Cheek-teeth two-rooted, except the first. On the hind feet the first and fifth toes greatly exceeding the others in length, with nails rudimentary or absent. AMonachus.\—Dentition : i 2,¢4, p 4, m 14; total 32. Crowns of molars strong, conical, compressed, hollowed on the inner side, with a strongly marked lobed cingulum, especially on the inner side, and slightly developed accessory cusps before and behind. The first and last upper and the first lower molar considerably smaller than the others. Vertebre: C 7,D15,L5,8 2,011. All the Fleming, Philosophy of Zoology, vol. ii. p. 187 (1822). PHOCIDE 605 nails of both fore and hind feet very small and rudimentary. One species, M. albiventer, the Monk-Seal of the Mediterranean and adjacent parts of the Atlantic. The other genera! of this section have the same dental formula, but are distinguished by the characters of the cheek-teeth and the feet. They are all inhabitants of the shores of the southern hemisphere. Ogmorhinus.2—All the teeth of the cheek-series with three distinct pointed cusps, deeply separated from each other; of these the middle or principal cusp is largest and slightly recurved ; the other two (anterior and posterior) are nearly equal in size, and have their apices directed towards the middle one. Skull much elongated. One species, 0. leptonyz, the Sea-Leopard, widely distrib- uted in the Antarctic and southern temperate seas. Lobodon.2—Cheek-teeth with much-compressed elongated crowns and a principal recurved cusp, rounded and somewhat bulbous at the apex, and one anterior, and one, two, or three posterior, very distinct accessory cusps. One species, L. carcinophaga. Pwecilophoca.*—Cheek-teeth small, with simple, subcompressed, conical crowns, having a broad cingulum, but no distinct accessory cusps. One species, P. weddelli. Ommatophoca.>—All the teeth very small; those of the cheek- series with pointed recurved crowns, and small posterior and still less developed anterior accessory cusps. Orbits very large. Nails quite rudimentary on front, and absent on hind feet. The skull bears a considerable resemblance to that of the members of the next subfamily, towards which it may form a transition. There is one species, 0. rossi, of which very little is known. Subfamily Cystophorinz.—Incisors 2. Teeth of cheek-series generally one-rooted. Nose of males with an appendage capable of being inflated. First and fifth toes of hind feet greatly exceeding the others in length, with prolonged cutaneous lobes, and rudi- mentary or no nails. Cystophora.-—Dentition: 7 2, ¢ +, p 4, m 4; total 30. The last molar has generally two distinct roots. Beneath the skin over the face of the adult male, and connected with the nostrils, is a sac which, when inflated, forms a kind of hood covering the 1 For details of these and the other genera see Mivart, Proc. Zool. Soc. 1885, p. 486, et seq. 2 Peters, Monatsd. K. P. Akad. Wissensch. zu Berlin, p. 393 (1875), substituted for Stenorhynchus, F, Cuvier ; preoccupied for a genus of Crustacea. 3 Gray, Zoology of Erebus and Terror, vol. i. p. 5 (1844). + New name, Syn. Leptonyx, Gray, Charlesworth’s Mag. Nat. Hist. vol. i. p. 582 (1837) ; preoccupied by Swainson, 1821. 5 Gray, Zoology of Erebus and Terror, vol. i. p. 7 (1844). 8 Nilsson, Faun. Scandinav, vol. i. p. 882 (1820). 606 CARNIVORA upper part of the head. Nails present, though small, on the hind feet. One species, (. cristutu, the Hooded or Bladder-Nose Seal of the Polar Seas. Macrorhinus.A—Dentition as the last, but cheek-teeth of simpler character, and all onerooted. All the teeth, except the canines, very small relatively to the size of the animal. Hind feet without nails.. Vertebre: C 7, D 15, L 5,8 3, C 11. Nose of adult male produced into a short tubular proboscis, ordinarily flaccid, but capable of dilatation and elongation under excitement. One species, J. Iconinus, the Elephant Seal, or Sea-Elephant of the whalers, the largest of the whole family, attaining the length of nearly 20 feet. Formerly abundant in the Antarctic Seas, and also found on the coast of California. Extinct Seals—Remains of animals of this group have been found in late Miocene and Pliocene strata in Europe and America, the most abundant and best-preserved being those of the Pliocene Antwerp Crag, the subject of an illustrated monograph by Van Beneden. Nothing has, however, yet been discovered which throws any light upon the origin of the group, since all the extinct forms at present known come within the definition of the existing families; and, though annectant forms between these occur, there are as yet no transitions to a more generalised type of mammal. Indeed, all those of which the characters are best known belong to the completely developed Phocine or Trichechine, and not to the Otariine, type. The typical genus Phocu occurs in the Antwerp Crag, while remains of Seals provisionally referred to this genus are found in the Pliocene of the Crimea and the Miocene of Malta and Virginia. Of the other Antwerp forms Callophocu is said to be allied to Phoca grenlundica, Platyphoca to Phoca barbata, Phocanella to Phoca foetida, Gryphoca to Halicherus, Palwophoca and Aonatherium to Aonachus, and Afesotaria to Cystophora ; while Prophoca does not appear to come very close to any existing form. It should be observed that it is extremely doubtful whether all these fossil Seals are really entitled to generic distinction. Bibliography of Pinnipedia.—J. A. Allen, History of North American Pinnipeds, 1880; St. George Mivart, ‘‘ Notes on the Pinnipedia,” Proc. Zool. Soc. 1885, p. 484; P. J. Van Beneden, Ossements fossiles d’ Anvers, in the Jkin. Acad. Roy. d. Belgique. Suborder CREODONTA. The discovery of fossil remains in Eocene and early Miocene formations both in Europe and North America shows that numerous species of terrestrial carnivorous animals existed upon the earth during those periods which cannot be referred to either of the 1 F. Cuvier, Afém. du Muséum, vol. xi. p. 200 (1824), ‘‘ Macrorhine.” CREODONTA 607 sections into which the order has now become broken up. By some zoologists these have been supposed to be Marsupials, or at least to show transitional characters between the Metatherian and Eutherian subclasses. By others they are looked upon as belonging altogether to the latter group, and as the common ancestors of existing Carnivores and Insectivores, or perhaps rather as descendants or relatives of such common ancestors, retaining more of the generalised characters than any of the existing species. They shade off almost insensibly into numerous other forms less distinctly carnivorous, to the whole of which, including the modern Insectivora, Cope (to whom we are indebted for much of our knowledge of the American extinct species) gives the name of BUNOTHERIA, those more specially related to the existing Carnivora forming the suborder Fic. 278.—Anterior portion of the skull of Hyenodon leptorhynchus. (After Filhol.) Creodonta. These are instances, however, in which the application of the principles of classification adopted in the case of existing species, of which the entire structure is known, and which have become divided into isolated groups by the extinction of inter- mediate forms, is almost impossible. If the generally accepted view of evolution is true, and the extreme modifications pass insensibly into each other by minute gradations (a view the paleontological proof of which becomes strengthened by every fresh discovery), there must be many of these extinct forms which cannot be assigned to definitely characterised groups. There are, however, some which stand out prominently from the others as formed on distinct types, having no exact representatives at present living on the earth. The more typical Creodonts appear, however, to be so closely related to the true Carnivora through the extinet Ifiacidw (p. 539), 608 CARNIVORA that it is on the whole advisable to regard them as representing a distinet suborder of Carnivora. In the strong development of the canines ‘Fig. 273: they are distinguished irom the modern Insect- ivora: and chey also differ from the latter and resemble the true Carniveres in the form of the incisors. the second one in the lower jaw (when three are present) being thrust up above the level of the other two in the manner obtaiming in mest of the modern Carni- vora. Some of the most generasised forms included in the present group approximate sc closely to the Condylarthrous Ungulates as to indicate that both c