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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. <A 
median interparietal 
ossification (Fig. 6, 
IP) may divide the 
parietals posteriorly, 
and is itself articu- 
lated with the supra- 
occipital, to the lat- 
eral borders of which 
the parietals are also 
joined. The squam- 


ees a Fic, 7.—Side view of skull of Cape Jumping Hare (Pedetes 
osal (Fig. ‘y Sq) forms caer). xX}. PMx, Premaxilla; Mz, maxilla; Ma, jugal or 
the lateral wall of malar; Fr, frontal; 1, lachrymal; Pa, parietal; Na, nasal; 
Sq, squamosal; 7'y, tympanic ; x0, exoccipital ; AS, alisphen- 
oid; OS, orbitosphenoid ; Per, mastoid bulla. 


the hinder part of 
the brain-case, and 
articulates superiorly with the parietal, and posteriorly with the 
exoccipital. The glenoid cavity (Fig. 8), for the reception of the 
articular condyle of the mandible, is formed by the inferior portion 
of the squamosal, at the point where it gives off the zygomatic 
process to form the hinder portion of the zygomatic arch. The 
middle portion of that arch is formed by the jugal, or malar bone 
(Fig. 7, Ma), which articulates posteriorly with the zygomatic process 
of the squamosal, and anteriorly with the maxilla. The jugal (as 
in Fig. 7) may also articulate with a small bone situated on the 
anterior border of the orbit known as the lachrymal. It is im- 
portant to observe that the zygomatic or temporal arch is a 
squamoso-maxillary one, and that an arcade thus composed is found 
elsewhere only among the extinct Anomodont reptiles, which have 
already been mentioned as showing signs of mammalian aflinity. 
The relative position occupied by the orbito- and alisphenoid is 
sufliciently indicated in Fig. 7 

Wedged in between the squamosal and the bones of the occipital 
and basisphenoidal region are the bones connected with the organ 
of hearing, known as the periotic and tympanic. The position of 
the periotic, which encloses the labyrinth or essential organ of 
hearing, is shown in Fig. 6. The periotic is divided into a very 
dense antero-internal moiety known as the petrosal, and a postero- 
external or mastoid portion (Fig. 8), which appears on the outer wall 
of the brain-case. The tympanic is produced horizontally outwards 
to form the external auditory meatus or tube of the ear, while the 


38 GENERAL ANATOMICAL CHARACTERS 


inner and under surface is frequently dilated into a shell-like 
auditory bulla (Fig. 8). The small bones of the internal ear known 
as the malleus, incus, and stapes are contained in the membranous 
tympanic cavity, 
which is situated in 
a space left among 
this group of bones. 
Further mention of 
these bones is made 
below under the 
head of the sense 
organs. 

In the Carni- 
vora and some other 
groups the foram- 
ina on the base of 
the skull for the 
passage of blood- 
vessels and nerves 
are of considerable 
taxonomic import- 
ance. The position 
of the more im- 
portant of these 
foramina is indi- 
cated in Fig. 8; 
but for details the 
reader may refer to 
the work on the 
Osteology of the Main- 
malia already men- 


tioned. Attention 
Fic. 8.—The right half of the hinder part of the base of the 


cranium of the Wolf (Canis lupus). ¢, Condyloid foramen ; 1, fora- may, “ however, be 
men lacerum posticum ; car, carotid canal; e, eustachian canal; particularly di- 
o, foramen ovale; a, posterior, and a’, anterior aperture of ali- rected to the s0- 
sphenoid canal; P, paroccipital process of exoccipital ; m, mastoid . 
process of periotic ; am, external auditory meatus; g, glenoid for- called alisphenoid 
amen, below which is the glenoid cavity for the condyle of the man- canal, the position 
dible. (Flower, Proc. Zool. Soc., 1869, p. 25.) of which is shown 


in Fig. 8, since this is a feature of some importance in the classifica- 
tion of the Carnivora. This canal is a short channel running hori- 
zontally forward from near the foramen ovale through the alisphenoid, 
and opening anteriorly with the foramen rotundum ; it is traversed 
by the external carotid artery. 

Only in those species, as Man and the smaller kinds of the 
Primates and some other orders, in which the brain holds a large 
relative proportion to the rest of the body, does the external form 


THE SKELETON 39 


of the skull receive much impress from the real shape of the cavity 
containing the brain. The size and form of the mouth, and the 
modifications of the jaws for the support of teeth of various shape 
and number, the ridges and crests on the cranium for the attachment 
of the muscles necessary to put this apparatus in motion, and out- 
growths of bone for the enlargement of the external surface required 
for the support of sense organs or of weapons, such as horns or 
antlers (which outgrowths, to prevent undue increase of weight, are 
filled with cells containing air), cause the principal variations in the 
general configuration of the skull. These variations are, however, 
only characteristically developed in perfectly adult animals, and are 
in many cases more strongly marked in the male than the female 
sex. Throughout all the later stages of growth up to maturity the 
size and form of the brain-case remain comparatively stationary, 
while the accessory parts of the skull rapidly increase and assume 
their distinctive development characteristic of the species. 

The hyoidean apparatus in mammals (Fig. 6) supports the tongue 
and larynx, and consists of an inferior median portion termed the 
basihyal, from which two pairs of half arches, or cornua, extend up- 
wards and outwards. The anterior is the more important, being 
connected with the periotic bone of the cranium. It may be almost 
entirely ligamentous, but more often has several ossifications, the 
largest of which is usually the stylohyal. The posterior cornu 
(thyrohyal) is united at its extremity with the thyroid cartilage of 
the larynx, which it suspends in position. The median portion, 
or basihyal, is sometimes, as in the Howling Monkeys, enormously 
enlarged and hollowed, admitting into its cavity an air-sac connected 
with the organ of voice. 

Vertebral Column.—The vertebral column consists of a series of 
distinct bones called vertebra, arranged in close connection with 
each other along the dorsal side of the neck and trunk, and in the 
median line! It is generally prolonged posteriorly beyond the 
trunk, to form the axial support of the appendage called the tail. 
Anteriorly it is articulated with the occipital region of the skull. 
The number of distinct bones composing the vertebral column 
varies greatly among the Mammalia, the main variation being 
due to the degree of elongation of the tail. Apart from this, in 
most mammals the number is not far from thirty, though it may 
fall as low as twenty-six (as in some Bats), or rise as high as 
forty (Hyrax and Cholepus). The different vertebre, with some 
exceptions, remain through life quite distinct from each other, 
though closely connected by means of fibrous structures which 
allow of a certain, but limited, amount of motion between them. 
The exceptions are the following:—(1) near the posterior part 

1 For the sake of uniformity, in all the following descriptions of the vertebral 
column, the long axis of the body is supposed to be in the horizontal position. 


40 GENERAL ANATOMICAL CHARACTERS 


of the trunk, in nearly all mammals which possess completely 
developed hinder limbs, two or more vertebree become ankylosed 
together to form the “sacrum,” or portion of the vertebral column 
to which the pelvic girdle is attached; (2) in some species of 
Whales and Armadillos there are constant ossific unions of certain 
vertebrae of the cervical region. 

Although the vertebre of different regions of the column of the 
same animal or of different animals present great diversities of 
form, yet there is a certain general resemblance among them, or a 
common plan on which they are constructed, which is more or less 
modified by alteration of form or proportions, or by the addition or 
suppression of parts to fit them to fulfil their special purpose in the 
economy. An ordinary or typical vertebra consists, in the first 
place, of a solid piece of bone, termed the body or centrum (Fig. 
9, ¢), of the form of a disk or short cylinder. The bodies of con- 


Fia. 9.—Anterior surface of Human 
thoracic vertebra (fourth). c, Body or 
centrum; nc, neural canal; , pedicle, 
and J, lamina of the arch; ¢, transverse 
process ; az, anterior zygapophysis. 


Fic. 10.—Side view of the first lum- 
bar vertebra of a Dog (Canis familiaris). 
s, Spinous process ; az, anterior zygapo 
physis ; pz, posterior zygapophysis ; m, 
metapophysis ; a, anapophysis; ¢, trans- 


verse process. 


tiguous vertebre are connected together by a very dense, tough, and 
elastic material called the “intervertebral substance,” of peculiar and 
complex arrangement. This substance forms the main, and in some 
cases the only, union between the vertebrae. Its elasticity provides 
for the vertebree always returning to their normal relation to each 
other and to the column generally, when they have been disturbed 
therefrom by muscular action. A process (») arises on each side 
from the dorsal surface of the body. These processes, meeting in 
the middle line above, form an arch, surmounting a space or short 
canal (nc). Since it contains the posterior prolongation of the 
great cerebro-spinal nervous axis, or spinal cord, this space is called 
the neural canal, and the arch the neural arch, in contradistinction 
to another arch on the ventral surface of the body of the verte- 
bree, called the hemal arch. The latter is, however, never formed 


THE SKELETON 41 


in mammals by any part of the vertebra itself, but by certain 
distinct bones placed more or less in apposition to it, namely the 
ribs in the thoracic, and the “chevron bones” in the caudal region. 
In most cases the arch of one vertebra is articulated with that of 
the next by distinct surfaces with synovial joints, placed one on 
each side, called ‘“‘zygapophyses ” (u:, 2), but these are often entirely 
wanting when flexibility is more needed than strength, as in the 
greater part of the caudal region of long-tailed animals. In addition 
to the body and the arch, there are certain projecting parts called 
processes, chiefly serving for the attachment of the numerous 
muscles which move the vertebral column. Of these two are single 
and median, viz. the spinous process, neural spine, or neurapophysis 
(s), arising from the middle of the upper part of the arch, and the 
hypapophysis from the under surface of the body. The latter, how- 
ever, is as frequently absent as the former is constant. The other 
processes are paired and lateral. They are the transverse processes 
(t), of which there may be two, an upper and a lower, in which case 
the former ‘is called, in the language of Owen (to whom we are 
indebted for the terminology of the parts of vertebrae in common 
use), “ diapophysis,” and the latter “ parapophysis.” Other processes 
less constantly present are called respectively “ metapophyses ” (1) 
and “anapophyses ” (a). 

The vertebral column is divided for convenience of description 
into five regions—the cervical, thoracic or dorsal, lumbar, sacral, and 
caudal. This division is useful, especially as it is not entirely 
arbitrary, and in most cases is capable of ready definition ; but at 
the contiguous extremities of the regions the characters of the 
vertebrae of one are apt to blend into 
those of the next region, either normally 
or as peculiarities of individual skeletons. 

Cervical Vertebre.—The cervical region 
constitutes the most anterior portion of 
the column, or that which joins the 
cranium. The vertebrae which belong to 
it are either entirely destitute of movable 
ribs, or if they have any these are small, 
and do not join the sternum. As a general 
rule they have a considerable perforation 
through the base of the transverse process pitta ate 
(the vertebrarterial canal, Fig. 11, 7); or, Pera ae ser eine Dos, 
as it is sometimes described, they have 5, spinous process; az, anterior 
two transverse processes, superior and zysapophysis; «, vertebrarterial 
: + + : oye canal; t, transverse process ; ¢’, its 
inferior, which meet at their extremities j\pridr jamelia. 
to enclose a canal. This, however, rarely 
applies to the last vertebra of the region, in which only the upper 
transverse process is usually developed. The transverse process, 


42 GENERAL ANATOMICAL CHARACTERS 


moreover, very often sends down near its extremity a more or 
less compressed plate (inferior lamella), which, being considered 
serially homologous with the ribs of the thoracic vertebre (though 
not developed autogenously), is often called the “costal” or 
“pleurapophysial” plate. This is usually largest on the sixth, and 
altogether wanting on the seventh vertebra. The first and second 
cervical vertebrae, called respectively “atlas” and “axis,” are 
specially modified for the function of supporting and permitting 
the free movements of the head. They are not united together 
by the intervertebral substance, but connected only by ordinary 
ligaments and synovial joints. 

The cervical region in mammals presents the remarkable 
peculiarity that, whatever the length or flexibility of the neck, the 
number of vertebre is the same, viz. seven, with the exception of 
the Manatee and Hoffman’s Two-toed Sloth (Cholepus hoffmannt), 
which both have but six, and the Three-toed Sloth (Bradypus 
tridactylus), which has nine, though in this case the last two usually 
support movable ribs, which are not sufficiently developed to reach 
the sternum. 

According to Parker there may occasionally be eight cervicals 
in the Pangolins (Manis). 

Dorsal Vertebree.—The dorsal (or, as it would be more correctly 
termed, thoracic) region consists of the vertebra succeeding those 
of the neck, which have ribs movably articulated to them. These 
ribs arch round the thorax—the anterior one, and usually the 
greater number of those that follow, being attached below to the 
sternum. 

Lumbar Vertebree.—The lumbar region consists of those vertebree 
of the trunk in front of the sacrum which bear no movable ribs. 
It may happen that, as the ribs decrease in size posteriorly (the 
last being sometimes more or less rudimentary), the step from the 
thoracic to the lumbar region may be gradual and rather undeter- 
mined in a given species; but most commonly this is not the 
case, and the distinction is as well defined here as in any other 
region. Asa general rule there is a certain relation between the 
number of the thoracic and lumbar vertebrae, the whole number 
being tolerably constant in a given group of animals, and any 
increase of the one being at the expense of the other. Thus in all 
known Artiodactyle Ungulata there are 19 dorso-lumbar vertebree ; 
but these may consist of 12 dorsal and 7 lumbar vertebra, or 13 
dorsal and 6 lumbar, or 14 dorsal and 5 lumbar. The smallest 
number of dorso-lumbar vertebre in mammals occurs in some 
Armadillos, which have but 14. The number found in Man, 
the higher Apes, and most Bats, viz. 17, is exceptionally low; 
19 prevails in the Artiodactyla, nearly all Marsupials, and very 
many Rodents; 20 or 21 in Carnivora and most Insectivora ; 


THE SKELETON 43 


and 23 in Perissodactyla. The highest and quite exceptional 
numbers are in the Two-toed Sloth (Cholepus) 27, and the Hyrax 
30. The prevailing number of rib-bearing vertebre is 12 or 13, 
any variation being generally in excess of these numbers. 

Sacral Vertebre.—The sacral region offers more difficulties 0. 
definition. Taking the human “os sacrum” as a guide for 
comparison, it is generally defined as consisting of those vertebra 
between the lumbar and caudal regions which are ankylosed 
together to form a single bone. It happens, however, that the 
number of such vertebrae varies in different individuals of the 
same or nearly allied species, especially as age advances, when a 
certain number of the tail vertebre generally become incorporated 
with the true sacrum. Other suggested tests—as those vertebrae 
which have a distinct additional (pleurapophysial) centre of ossifica- 
tion between the body and the ilium, those to which the ilium is 
directly articulated, or those in front of the insertion of the ischio- 
sacral ligaments—being equally unsatisfactory or unpractical, the 
old one of ankylosis, as it is found to prevail in the average 
condition of adults in each species, is used in the enumeration of 
the vertebre in the following pages. The Cetacea, having no iliac 
bones, have no part of the vertebral column modified into a 
sacrum. 

Caudal Vertebre.—The caudal vertebre are those placed behind 
the sacrum, and terminating the vertebral column. They vary 
in number greatly—being reduced to 5, 4, or even 3, in a most 
rudimentary condition, in Man 
and in some Apes and Bats, and 
being numerous and powerfully 
developed, with strong and com- 
plex processes, in many mammals, 
especially among the Edentata, 
Cetacea, and Marsupialia. The 
highest known number, 46, is 
possessed by the African Long- 
tailed Pangolin. Connected with 
the under surface of the caudal 
vertebree of many mammals which 
have the tail well developed are 
certain bones formed more or less 
like an inverted arch, called chev- 
ron bones, or by the French os en — Fra. 12.— Anterior surface of fourth 
a These are always situated caudal vertebra of Porpeise (Phoceenc. cate: 

. munis). 8s, Spinous process ; m, metapophy- 
nearly opposite to an interverte- sis; t, transverse process ; h, chevron bone. 
bral space, and are generally artic- 
ulated both to the vertebra in front and the vertebra behind, but 
sometimes chiefly or entirely either to one or the other. 


44 GENERAL ANATOMICAL CHARACTERS 


In some of the Anomodont Reptiles and Labyrinthodont 
Amphibians these chevrons are attached to the intercentra—or 
imperfect disks alternating with the true centra—which suggests 
that they are primarily intercentral elements which have been trans- 
ferred to the edges of the centra by the disappearance of the inter- 
centra. 

Sternum.—The sternum of mammals is a bone, or generally a 
series of bones, placed longitudinally in the mesial line, on the 
inferior or ventral aspect of the thorax, and connected on each side 
with the vertebral column by a series 
of more or less ossified bars called 
“ribs.” It is present in all mammals, 
but varies much in character in the 
different groups. It usually consists 
of a series of distinct segments placed 
one before the other, the anterior 
being called the presternum or “ manu- 
brium sterni” of human anatomy, and 
the posterior the xiphisternum, or 
xiphoid or ensiform process, while the 
intermediate segments, whatever their 
number, constitute the mesosternum 
or “body.” In the Whalebone Whales 
the presternum alone is developed, and 
but a single pair of ribs is attached 
to it. 

Libs—The ribs form a series of 
long, narrow, and more or less flattened 
bones, extending laterally from the 

ie as ena ide’ Caleta aes sides of the vertebral column, curving 
sternal ribs. ps, Presternam; ms, G@Oownwards towards the median line 
mesos ternunn: rs, xiphisternum ; cy of the body below, and mostly joining 
fie ee Le the sides of the sternum. The posterior 
ribs, however, do not directly articulate 
with that bone, but are either attached by their extremities to 
the edges of each rib in front of them, and thus only indirectly 
join the sternum, or else they are quite free below, mecting no part 
of the skeleton. These differences have given rise to the division 
into “true” and “false” ribs (by no means good expressions), signi- 
fying those that join the sternum directly and those that do not : 
and of the latter, those that are free below are ealled “ floating ” 
ribs. The portion of each rib nearest the vertebral column and 
that nearest the sternum differ in their charactors, the latter being 
usually but imperfectly ossified, or remaining permanently eartila- 
ginous. These are called “costal cartilages,” or when ossified 
“sternal ribs.” 


ms 


THE SKELETON 45 


In the anterior part of the thorax the vertebral extremity of 
each rib is divided into two parts, “head” or “capitulum,” and 
“tubercle”; the former is attached to the side of the body of the 
vertebra, the latter to its 
transverse process; the 
former attachment corre- 
sponds to the interspace 
between the vertebra, the 
head of the rib commonly 
articulating partly with 
the hinder edge of the 
body of the vertebra ante- 
cedent to that which bears 
its tubercle. Hence the 
body of the last cervical 
vertebra usually supports 
part of the head of the first 
rib. In the posterior part 
of the series the capitular 
and tubercular attach- 


ments commonly coalesce, Fria. 14.—Sternum and strongly ossified sternal ribs 


and the rib is attached of Great Armadillo (Priodon gigas). ps, Presternum ; 
xs, Xiphisternum. 


solely to its corresponding 
vertebra. The number of pairs of ribs is of course the same as that 
of the thoracic vertebree. 

The circumstance that in some of the Anomodont reptiles and 


Fic. 15.—Skeleton of Lion (Felis leo). cd, Caudal vertebre ; cp, carpus; cr, coracoid process 
of scapula ; cv, cervical vertebre ; d, dorsal vertebrae ; fb, fibula; fm, femur; h, humerus; il, 
ilium ; isch, ischium; 2, lumbar vertebre ; m, metatarsus ; mc, metacarpus ; 7p, patella ; pb, pubis ; 
ph, phalanges ; pv, pelvis; r, radius; s, sacral vertebre; sc, scapula; sk, skull; th, tibia; ts, 
tarsus ; u, ulna; zy, zygomatic arch. 


Labyrinthodonts the capitula of the ribs articulate with the inter- 
central elements of the vertebral column has suggested, as in the 


46 GENERAL ANATOMICAL CHARACTERS 


instance of the chevron bones, that the intercentral capitular articu- 
lation of the ribs of mammals is a feature directly inherited 
from those extinct types by the gradual disappearance of the 
intercentra. 

Appendicular Skeleton.—The appendicular portion of the frame- 
work consists, when completely developed, of two pairs of limbs, 
anterior and posterior (Fig. 15). 

Anterior Limb.—The anterior limb is present and fully developed 
in all mammals, being composed of a shoulder girdle and three seg- 
ments belonging to the limb proper, viz. the upper arm or brachium, 
the fore-arm or antebrachium, and the hand or manus. 

Shoulder-girdle.—The shoulder or pectoral girdlein the large majority 
of mammals is in a rudimentary or rather modified condition, com- 
pared with that in which it exists in other vertebrates. In the Mono- 
tremata (Ornithorhynchus and Echidna) alone is the ventral portion, or 
coracoid, complete and articulated with the sternum below, as in the 
Sauropsida ; and in this group alone do we find an anterior ventral 
element, apparently corresponding with the precoracoid of the Anom- 
odont reptiles, although generally known as the epicoracoid. In all 
other mammals the coracoid, though ossified from a distinct centre, 
forms only a process, sometimes a scarcely distinct tubercle, projecting 
from the anterior border of the glenoid cavity of the scapula. The 
last-named cavity, which in the Monotremes is formed jointly by 
the scapula and coracoid, receives the head of the humerus, or 
arm-bone. The scapula is always well developed, and generally 
broad and flat (whence its vernacular name “blade bone”), with a 
ridge called the “spine” on its outer surface, which usually ends in 
a free curved process, the “acromion.” As the scapula affords 
attachment to many of the muscles which act upon the anterior 
limb, its form and the development of its processes are greatly 
modified according to the uses to which the member is put. Thus it 
is most reduced and simple in character in those animals whose limbs 
are mere organs of support, as the Ungulates ; and most complex 
when the limbs are also used for grasping, climbing, or digging. 
The development or absence of the clavicle or “collar-bone,” an 
accessory bar which connects the sternum with the scapula and 
steadies the shoulder-joint, has a somewhat similar relation, though 
its complete absence in the Bears shows that this is not an invariable 
rule. A complete clavicle is found in Man and all the Primates, in 
Chiroptera, all Insectivora (except Potamogale), in many Rodents, in 
most Edentates, and in all Marsupials, except Perameles. More or 
less rudimentary clavicles (generally suspended freely in the muscles) 
are found in the Cat, Dog, and most Carnivora, Myrmecophaga, and 
some Rodents. Clavicles are altogether absent in most of the Urside, 
all the Pinnipedia, Manis among Edentates, the Cetacea, Sirenia, 
Ungulates, and some Rodents. 


THE SKELETON 47 


The Monotremes are peculiar in possessing a T-shaped 
interclavicle like that of many reptiles, lying upon the sternum, 
and articulating superiorly with the clavicles. 

Brachium and Antebrachium.—The proximal segment of the 
anterior or pectoral limb proper contains a single bone, the humerus, 
and the second segment two bones, the radius and the ulna, placed 
side by side, and articulating with the humerus at their proximal, 
and with the carpus at their distal extremity (Fig. 15). In their 
primitive and unmodified condition these bones may be considered as 
placed one on each border of the limb, the radius being preaxial or 
anterior, and the ulna postaxial or posterior, when the distal or free 
end of the limb is directed outwards, or away from the trunk. This 
is their position in the earliest embryonic condition, and is best 
illustrated among adult mammals in the Cetacea, where the two 
bones are fixed side by side and parallel to each other. In the 
greater number of mammals the bones assume a very modified and 
adaptive position, usually crossing each other in the forearm, the 
radius in front of the ulna, so that the preaxial bone (radius), 
though external (in the ordinary position of the limb) at the upper 
end, is internal at the lower end; and the hand, being mainly fixed 
to the radius, also has its preaxial border internal. In the large 
majority of mammals the bones are fixed in this position, but in 
some few, as in Man, a free movement of crossing and uncrossing— 
or pronation and supination, as it is termed—is allowed between 
them, so that they can be placed in their primitive parallel condition, 
when the hand (which moves with the radius) 
is said to be supine, or they may be crossed, 
when the hand is said to be prone. 

The humerus frequently has a foramen 
piercing the inner border of the distal 
extremity, known as the entepicondylar 2 
foramen, which corresponds with a similar 
one found in the Anomodont reptiles. The 
hollow in the head of the ulna for the recep- ~~" i 
tion of the head of the humerus is known 
as the greater sigmoid cavity, and that for Ai 
the head of the radius as the lesser sigmoid aa Wei 
cavity (Fig. 16). The term olecranon is | iii) i i 
applied to that process of the ulna which Pena 3 
forms the prominence of the elbow. PFW Sed pes ck Wied 

In most mammals walking on four limbs, ane aie Pha Heap (eaus). 
in which the hand is permanently prone, the «, Anterior tubercle; ol, 
ulna is much reduced in size, and the radius pon serene fs een 
increased, especially at the upper end; so ve 
that the articular surface of the latter, instead of being confined to 
the external side of the trochlea of the humerus, extends all across 


{ 


48 GENERAL ANATOMICAL CHARACTERS 


its anterior surface, and the two bones, instead of being external 
and internal, are anterior and posterior. In many hoofed or Ungu- 
late mammals, and in Bats, the ulna is reduced to little more than 
its upper articular extremity, and firmly ankylosed to the radius 
—-stability of these parts being more essential than mobility. 
Alanus.—The terminal segment of the anterior limb is the hand 
or manus. Its skeleton consists of three divisions: (1) the 
“carpus,” a group of small, more or less rounded or angular bones 
with flattened surfaces applied to one another, and, though arti- 
culating by synovial joints, having scarcely any motion between 
them ; (2) the “metacarpus,” a series of elongated bones placed side 
by side, with their proximal ends articulating by almost immovable 
joints with the carpus; (3) the “phalanges” or bones of the digits, 
usually three in number to each, articulating to one another by freely 
movable hinge-joints, the first being connected in like manner to 
the distal end of the metacarpal bone to which it corresponds. 
Carpus.—To understand thoroughly the arrangement of the 
bones of the carpus in mammals, it is necessary to study their 
condition in some of the lower vertebrates. Fig. 17 represents 
the manus in one of its fullest and at the same time most 
generalised forms, as seen in one of the 
Water Tortoises (Chelydra serpentina). The 
carpus consists of two principal rows of 
bones. The upper or proximal row con- 
tains three bones, to which Gegenbaur 
has applied the terms radiale (r), inter- 
medium (i), and ulnare (uw), the first being 
on the radial or preaxial side of the limb. 
The lower or distal row contains five 
bones, called carpale 1, 2, 3, 4, and 5 
respectively, commencing on the radial 
side. Between these two rows, in the 
middle of the carpus, is a single bone, 
the centrale (c). In this very symmetrical 
carpus it will be observed that the rudiale 
supports on its distal side two bones, 
Set eR ee ee carpale land 2; the zntermedium is in a 
right manus of a Water Tortoise line with the centrale and carpale 3 which 
e se th ee tne ~ together form a median axis of the hand, 
ulnare ; i, intermedium ; 7, radiale ; while the oe has also two bones articu- 
entrale : 1-5 Ave bi s of ¢ o rit, s ste as aaa 
Me one Ge ee 
m5, the five metacarpals. carpals o the distal 
row supports a metacarpal. 


fing ae caaee Sees ap 

The opinion has recently been expressed by Baur that the bone termed 
radiale in Fig. 17 is really a second centrale, and that the radiale is represented 
by a minute bone generally known as the radial sesamoid. The mammalian 


THE SKELETON 49 


In the carpus of the Mammalia there are usually two additional 
bones developed in the tendons of the flexor muscles, one on each 
side of the carpus, which may be called the radial and ulnar 
sesamoid bones ; the latter, which is the more constant and generally 
larger, is commonly known as the pisiform bone. The fourth and 
fifth carpals of the distal row are always united into a single bone, 
and the centrale is very often absent. As a general rule all the 
other bones are present and distinct, though it not unfrequently 
happens that two may have coalesced to form a single bone, or 
one or more may be altogether suppressed. 

The following table shows the principal names in use for the 
various carpal bones,—those in the second column being the terms 
generally employed by English anatomists :— 


Radiale =Scaphoid = Naviculare. 

Intermedium = Lunar = Semilunare, Lunatum. 
Ulnare = Cuneiform = Triquetrum, Pyramidale. 
Centrale =Central = Intermediwm (Cuvier). 
Carpale1 =Trapezium =Multangulum majus. 
Carpale2 =Trapezoid = Multangulum minus. 
Carpale3 =Magnum = Capitatwm. 

od ani \ =Unciform = Hamatum, Uncinatum. 
Carpale 5 q 


The radial and ulnar sesamoids are regarded by Bardeleben? as 
the rudiments of a prepollex and a postminimus digit ; the primitive 
number of digits being thus supposed to have been seven. These 
bones have been observed in all orders of mammals having five 
complete digits. Occasionally, as in Pedetes caffer, the so-called 
prepollex consists of two bones, of which the distal one bears a 
distinct nail-like horny covering. In Bathyergus maritimus the 
pisiform, or postminimus, is likewise double; the two elements 
being regarded by their describer as representing the carpal and 
metacarpal of the presumed seventh digit. 

Similarly in the posterior limb the tibial sesamoid, and a fibular 
ossification corresponding to the pisiform, are regarded as represent- 
ing a prehallux and a postminimus. 

Metacarpus and Phalanges——The metacarpal bones, with the 
digits which they support, are never more than five in number, and 
are described numerically —first, second, etc., counting from the 
radial towards the ulnar side. The digits are also sometimes named 
(1) the pollex, (2) index, (3) medius, (4) annularis, (5) minimus. 


scaphoid is accordingly also regarded as a second centrale. In the same com- 
munication, Dr. Baur expresses his disbelief in the existence of remnants of a 
prepollex and of a seventh digit in mammals and other vertebrates. (See Anat. 
Anzeiger, vol. iv. pp. 49-52, 1889.) 
1 On the Prepollex and Prehallux, ete., Proc. Zool. Soc. 1889, pp. 259-262. 
4 


50 GENERAL ANATOMICAL CHARACTERS 


One or more may be in a rudimentary condition, or altogether 
suppressed. If one is absent, it is most commonly the first. 
Excepting the Cetacea, no mammals have more than three phalanges 
to each digit, but they may occasionally have fewer by suppression 
or ankylosis. The first or radial digit is an exception to the usual 
rule, one of its parts being constantly absent, since, while each of the 
other digits has commonly a metacarpal and three phalanges, it has 
only three bones altogether ; whether the missing one is a meta- 
carpal or one of the phalanges is a subject which has occasioned 
much discussion, and has not yet been satisfactorily decided. The 
terminal phalanges of the digits are usually specially modified to 
support the nail, claw, or hoof, and are called “ungual phalanges.” 
In walking, some mammals (as the Bears) apply the whole of the 
lower surface of the carpus, metacarpus, and phalanges to the 
ground ; to these the term “ plantigrade” is applied. Many others 
(as nearly all the existing Ungulata) only rest on the last one or two 
phalanges of the toes, the first phalanx and the metacarpals being 
vertical and in a line with the fore-arm. These are called “digiti- 
grade.” Intermediate conditions exist between these two forms, to 
which the terms “phalangigrade” (as the Camel) and “subplanti- 
grade” (as in most Carnivora), are applied. When the weight is 
borne entirely on the distal surface of the ungual phalanx, and the 
horny structures growing around it, as in the Horse, the mode of 
progression is called “ unguligrade.” 

In the Chiroptera the digits are enormously elongated, and 
support a cutaneous expansion constituting the organ of flight. In 
the Cetacea the manus is formed into a paddle, being covered by 
continuous integument, which conceals all trace of division into 
separate digits, and shows no sign of nails or claws. In the Sloths 
the manus is long and very narrow, habitually curved, and terminat- 
ing in two or three pointed curved claws in close apposition with 
each other, and incapable, in fact, of being divaricated ; so that it is 
reduced to the condition of a hook, by which the animal suspends 
itself to the boughs of the trees among which it lives. These are 
only examples of the endless modifications to which the distal 
extremity of the limb is subjected in adaptation to the various 
purposes to which it is applied. 

Posterior Limb.—The posterior limb is constructed upon a plan 
very similar to that of the anterior extremity. It consists of a 
pelvic girdle and three segments belonging to the limb proper, viz. 
the thigh, the leg, and the foot or pes (Fig. 15). 

Pelvic Girdle.—The pelvic girdle is present in some form in all 
mammals, though in the Cetacea and the Sirenia it is in an exceed. 
ingly rudimentary condition. In all mammals except those be- 
longing to the two orders just named, each lateral half of the pelvic 
girdle consists essentially, like the corresponding part of the anterior 


THE SKELETON 51 


limb, of a flattened rod of bone crossing the long axis of the trunk, 
having an upper or dorsal and a lower or ventral end. The upper 
end diverges from that of the opposite side, but the lower end 
approaches, and, in most cases, meets it, forming a symphysis, 
without the intervention of any bone corresponding to the sternum. 
The pelvic girdle differs from the shoulder girdle in being firmly 
articulated to the vertebral column, thus giving greater power to 
the hinder limb in its function of supporting and propelling the 
body. Like the shoulder girdle, it bears on its outer side, near 
the middle, a cup-shaped articular cavity (“acetabulum”), into 
which the proximal end of the first bone of the limb proper is 
received. Each lateral half of the girdle is called the “os 
innominatum,” or innominate bone, and consists originally of three 
bones which unite at the acetabulum. The “ilium” or upper bone 
is that which articulates with the sacral vertebre. Of the two 
lower bones the anterior or “pubis” unites with its fellow of 
the other side at the symphysis; the posterior is the “ischium.” 
These lower elements form two bars of bone, united above and 
below, but leaving a space between them in the middle, filled only 
by membrane, and called the “thyroid” or “obturator” foramen. 
The whole circle of bone formed by the two innominate bones 
and the sacrum is called the pelvis. In the Monotremata 
and Marsupialia, a pair of thin, flat, elongated ossifications 
called epipubic or marsupial bones are attached to the fore part 
of the pubis, and project forward into the muscular wall of the 
abdomen. 

Thigh and Leg.—The first segment of the limb proper has one 
bone, the femur, corresponding with the humerus of the anterior 
limb. The second segment has two bones, the tibia and fibula, corre- 
sponding with the radius and ulna. These bones always lie in their 
primitive unmodified position, parallel to each other, the tibia on 
the preaxial and the fibula on the postaxial side, and are never 
either permanently crossed or capable of any considerable amount 
of rotation, as in the corresponding bones of the fore limb. In the 
ordinary walking position the tibia is internal, and the fibula ex- 
ternal. In many mammals the fibula is in a more or less rudi- 
mentary condition, and it often ankyloses with the tibia at one or 
both extremities. The patella or “knee-cap,” which is found in an 
ossified condition in all mammals, with the exception of some of 
the Marsupialia, is a large sesamoid bone developed in the tendon 
of the extensor muscles of the thigh, where the tendon passes over 
the front of the knee-joint, to which it serves as a protection. 
There are frequently smaller ossicles, one or two in number, situated 
behind the femoral condyles, called “fabellz.” The processes for 
the attachment of muscles near the upper end of the femur are 
termed trochanters; and the third trochanter, found on the hinder 


52 GENERAL ANATOMICAL CHARACTERS 


aspect of the shaft of this bone in many forms is of considerable 
taxonomic importance. 

Pes,—The terminal segment of the hind limb is the foot or pes. 
Its skeleton presents in many particulars a close resemblance to that 
of the manus, being divisible into three parts: (1) a group of 
short, more or less rounded or square bones, constituting the 
tarsus ; (2) a series of long bones placed side by side, forming the 
metatarsus ; and (3) the phalanges of the digits or toes. 

The bones of the tarsus of many of the lower Vertebrata closely 
resemble both in number and arrangement those of the carpus, as 
shown in Fig. 17. They have been described in their most general- 
ised condition by Gegenbaur under the names expressed in the first 
column of the following table. The names in the second column are 
those by which they are generally known to English anatomists, 
while in the third column some synonyms occasionally employed 
are added. 


ene) = \ = Astragalus 1 = Talus. 

Fibulare = Calcaneum = 0s calcis. 
Centrale = Navicular = Scaphoideum. 
Tarsale 1 = Internal cuneiform = Entocunetforme. 
Tarsale 2 = Middle cuneiform = Mesocunetforme. 
Tarsale 3 = External cuneiform = Ectocunetforme. 
Tarsale 4 ! 

Tarsale 5 \ = Cubeid. 


The bones of the tarsus of mammals present fewer diversities of 
number and arrangement than those of the carpus. The proximal 
row (see Fig. 18) always consists of two bones, namely the astra- 
galus (a), which probably represents the coalesced scaphoid and lunar 
of the hand, and the caleaneum (c). The former is placed more to 
the dorsal side of the foot than the latter, and almost exclusively 
furnishes the tarsal part of the tibio-tarsal or ankle-joint. The cal- 
caneum, placed more to the ventral or “plantar” side of the foot, is 
elongated backwards to form a more or less prominent tuberosity, 
the “tuber calcis,” to which the tendon of the great extensor muscles 
of the foot is attached. The navicular bone (n) is interposed between 
the proximal and distal row on the inner or tibial side of the foot, 
but on the outer side the bones of the two rows come into contact. 
The distal row, when complete, consists of four bones, which, be- 
ginning on the inner side, are the three cuneiform bones, internal 
(c), middle (¢?), and external (c*), articulated to the distal surface 
of the navicular, and the cuboid (cb), articulated with the calcaneum. 
Of these the middle cuneiform is usually the smallest in animals 


1 Cope and Baur consider that the astragalus corresponds only with the inter- 
medium, and that the tibiale may exist as a distinct element. 


THE DIGESTIVE SYSTEM 53 


in which all five digits are developed; but when the hallux is 
wanting the internal cuneiform may be rudimentary or altogether 
absent. The three cuneiform bones sup- 
port respectively the first, second, and third 
metatarsals, and the cuboid supports the 
fourth and fifth ; they thus exactly corre- 
spond with the four bones of the distal row 
of the carpus. 

In addition to these constant tarsal 
bones, there may be supplemental or 
sesamoid bones: one situated near the 
middle of the tibial side of the tarsus, 
largely developed in many Carnivora and 
Rodentia ; another, less frequent, on the 
fibular side; and a third, often developed 
in the tendons of the plantar surface of 
the tarsus, is especially large in Armadillos. 
There is also usually a pair of sesamoid 
bones on the plantar aspect of each meta- 
tarso-phalangeal articulation. In the young 
of the carnivorous genus C'ryptoprocta there 
may be a second centrale, which usually 
coalesces with the ectocuneiform. 

; The metatarsal bones never exceed five a rae ua ste ght 
in number, and the phalanges follow the smetatarsus; Ph, phalanges; c, 
same numerical rule as in the manus, never caleaneum; a, astragalus; cb, 
exceeding three in each digit. Moreover, ei Ee a 
the first digit, counting from the tibial side, form ; c3, external cuneiform. The 
or hallux, resembles the pollex of the hand ‘its are indicated by Roman 
. « numerals, counting from the 
in always having one segment less than tiyiai to the tibular side. 

the other digits. As the function of the 

hind foot is more restricted than that of the hand the modifica- 
tions of its structure are less striking. In the Cetacea and the 
Sirenia it is entirely wanting, though in some existing members of 
the first-named order rudiments of the bones of both the first and 
second segments of the limb have been detected, and a femur is 
present in the Miocene Sirenian Hulitherium. 


IV, THE DIGESTIVE SYSTEM. 


General Considerations. —The search after the purpose which 
every modification of structure subserves in the economy is always 
full of interest, and, if conducted with due caution and sutticient 
knowledge of all the attendant circumstances, may lead to important 
generalisations. It must always be borne in mind, however, that 


54 GENERAL ANATOMICAL CHARACTERS 


adaptation to its special function is not the only cause of the 
particular form or structure of an organ, but that this form, having 
in all probability been arrived at by the successive and gradual 
modification of some other different form from which it is now to a 
greater or less degree removed, has other factors besides use to be 
taken into account. In no case is this principle so well seen as in 
that of the organs of digestion. These may be considered as 
machines which have to operate upon alimentary substances in very 
different conditions of mechanical and chemical combination, and to 
reduce them in every case to the same or precisely similar 
materials ; and we might well imagine that the apparatus required 
to produce flesh and blood out of coarse fibrous vegetable substances 
would be different from that which had to produce exactly the 
same results out of ready-made flesh or blood ; and in a very broad 
sense we find that this is so. Thus, if we take a large number of 
carnivorous animals, belonging to different fundamental types, and 
a large number of herbivorous animals, and strike a kind of average 
of each, we shall find that there is, pervading the first group, a 
general style, if we may use the expression, of the alimentary organs, 
different from that of the others. That is to say, there is a specially 
carnivorous and a specially herbivorous modification of these parts. 
But, if function were the only element which has guided such 
modification, it might be inferred that, as one form must be supposed 
to be best adapted in its relation to a particular kind of diet, that 
form would be found in all the animals consuming such diet. But 
this is far from being the case. Thus the Horse and the Ox, for 
instance—two animals whose food in the natural state is precisely 
similar—are most different as regards the structure of their ali- 
mentary canal, and the processes involved in the preparation of that 
food. Again, the Seal and the Porpoise, both purely fish-eaters, 
which seize, swallow, and digest precisely the same kind of prey, in 
precisely the same manner, have a totally different arrangement of the 
alimentary canal. If the Seal’s stomach is adapted in the best conceiv- 
able manner for the purpose it has to fulfil, why is not the Porpoise’s 
stomach an exact facsimile of it, and vice versd ? We can only answer 
that the Seal and Porpoise belong to different natural groups of 
animals, formed either on different primitive types, or descended 
from differently constructed ancestors. On this principle only can 
we account for the fact that, whereas, owing to the comparatively 
small variety of the different alimentary substances met with in 
nature, few modifications would appear necessary in the organs of 
digestion, there is really endless variety in the parts devoted to 
this purpose. 

- Mouth.—The digestive apparatus of mammals, as in other ver- 
tebrates, consists mainly of a tube with an aperture placed at or 
near either extremity of the body,—the oral and the anal orifice, — 


THE DIGESTIVE SYSTEM 55 


and furnished with muscular walls, the fibres of which are so 
arranged as by their regular alternate contraction and relaxation to 
drive onwards the contents of the tube from the first to the second 
of these apertures. The anterior or commencing portion of this 
tube and the parts around it are greatly and variously modified in 
relation to the functions assigned to them of selecting and seizing 
the food, and preparing it by various mechanical and chemical 
processes for the true digestion which it has afterwards to undergo 
before it can be assimilated into the system. For this end the tube 
is dilated into a chamber or cavity called the mouth, bordered 
externally by the lips, which are usually muscular and prehensile, 
and supported by a movable framework carrying the teeth; the 
structure and modifications of which have been already described. 
The roof of the mouth is formed by the palate, terminating behind 
by a muscular, contractile arch, having in Man and some few other 
species a median projection called the uvula, beneath which the 
mouth communicates with the pharynx. The anterior part of the 
palate is composed of mucous membrane tightly stretched over the 
flat or slightly concave bony lamina separating the mouth from 
the nasal passages, and is generally raised into a series of trans- 
verse ridges, which sometimes, as in Ruminants, attain a con- 
siderable development. In the floor of the mouth, between the 
rami of the mandible, and supported behind by the hyoidean 
apparatus, lies the tongue; an organ the free surface of which, 
especially in its posterior part, is devoted to the sense of taste, but 
which also, by its great mobility (being composed almost entirely 
of muscular fibres), performs important mechanical functions 
connected with masticating and procuring food. Its modifications 
of form in different mammals are very numerous. Between the 
long, extensile, vermiform tongue of the Anteaters, which is 
essential to the peculiar mode of feeding of those animals, and the 
short, sessile, and almost functionless tongue of the Porpoise, every 
intermediate condition is found. Whatever the form, the upper 
surface is always. covered with numerous fine papille, in which 
the terminal filaments of the gustatory nerve are distributed. 

Salivary Glands.—The fluid known as the saliva is secreted by 
an extensive and complex system of glands discharging into the 
cavity of the mouth (buccal cavity), the position and relation of 
some of which are exhibited in the woodcut on the next page 
(Fig. 19). 

This apparatus consists of small glands embedded in the mucous 
membrane or submucous tissue lining the cavity of the mouth, 
which are of two kinds (the follicular and the racemose), and of 
others in which the secreting structure is aggregated in distinct 
masses removed some distance from the cavity; other tissues besides 
the lining membrane being usually interposed, and pouring their 


56 GENERAL ANATOMICAL CHARACTERS 


secretion into the cavity by a distinct tube or duct, which traverses 
the mucous membrane. To the latter alone the name of “salivary 
glands” is ordinarily appropriated, although the distinction 
between them and the smaller racemose glands is only one of 
convenience for descriptive purposes, their structure being more or 
less nearly identical ; and, since the fluids secreted by all become 
mixed in the mouth, their functions are, at all events in great part, 
common. Under the name of salivary glands are commonly 


Fic, 19.—Salivary Glands of the Genet. A, Right side of the head dissected ; p, parotid 
gland ; d, Steno’s duct; sm, submaxillary gland, traversed by the jugular veins (jv); 0, aperture 
of Steno’s duct. B, Part of the head with the lip drawn up to show (st.d) aperture of 
Steno’s duct; z.gl, zygomatic gland; 0, aperture of do.; z, zygomatic arch. (Mivart, Proe. 
Zool. Soc. 1882, p. 504.) 


included—(1) the “parotid” (p), situated very superficially on the 
side of the head, below or around the cartilaginous external 
auditory meatus, and the secretion of which enters the mouth by 
a duet (often called Steno’s or Stenson’s) which crosses the masseter 
muscle and opens into the upper and back part of the cheek 
(Fig. 19); and (2) the “submaxillary” (sm), situated in the neck 
near or below the angle of the mandible, and sending a long duet 


THE DIGESTIVE SYSTEM 57 


(Wharton’s) forwards to open on the fore-part of the floor of the 
cavity of the mouth, below the apex of the tongue. These are the 
most largely developed and constant of the salivary glands, being 
met with in various degrees of development in almost all animals 
of the class. Next in constancy are (3) “the sublingual,” closely 
associated with the last-named, at all events in the locality in which 
the secretion is poured out; and (4) the “zygomatic” (z.gl), found 
only in some animals in the cheek, just under cover of the anterior 
part of the zygomatic arch, its duct entering the buccal cavity near 
that of the parotid. 

The most obvious function common to the secretion of these 
various glands, and to that of the smaller ones placed in the mucous 
membrane of the lips, the cheeks, the tongue, the palate, and fauces, 
is the mechanical one of moistening and softening the food, to 
enable it the more readily to be tasted, masticated, and swallowed, 
though each kind of gland may contribute in different manner 
and different degree to perform this function. The saliva is, 
moreover, of the greatest importance in the first stage or introduc- 
tion to the digestive process, as it dissolves or makes a watery 
extract of all soluble substances in the food, and so prepares them 
to be further acted on by the more potent digestive fluids met with 
subsequently in their progress through the alimentary canal. In 
addition to these functions it seems now well established by experi- 
ment that saliva serves in Man and many animals to aid directly 
in the digestive process, particularly by its power of inducing the 
saccharine transformation of amylaceous substances. As a general 
rule, in mammals the parotid saliva is more watery in its 
composition, while that of the submavxillaries, and still more the 
sublingual, contains more solid elements and is more viscid ;—so 
much so that some anatomists consider the latter, together with the 
small racemose glands of the cheeks, lips, and tongue, as mucous 
glands, retaining the name of salivary only for the parotid. These 
peculiar properties are sometimes illustrated in a remarkable 
degree, as, for example, the great secretion of excessively viscid 
saliva which lubricates the tongue of the Anteaters and Armadillos, 
associated with enormously developed submaxillary glands ; while, 
on the other hand, the parotids are of great size in those animals 
which habitually masticate dry and fibrous food. 

Stomach.—After the preparation which the aliment has under- 
gone in the mouth,—the extent of which varies immensely in 
different forms, being reduced almost to nothing in such animals as 
the Seals and Cetaceans, which, to use the familiar expression, 
“bolt” their food entire, and most fully carried out in the Rumin- 
ants, which “chew the cud,”—it is swallowed, and carried along 
the cesophagus by the action of its muscular coats into the stomach. 
In the greater number of mammals this organ is a simple saccular 


58 GENERAL ANATOMICAL CHARACTERS 


dilatation of the alimentary canal, as in Figs. 20, 21, but in others 
it undergoes remarkable modifications and complexities. The lining 
of the stomach is thickly beset with tubular glands, which are 
generally considered to belong to two different forms, recognisable 
by their structure, and different in their function—the most 
numerous and important secreting the gastric juice (the active 
agent in stomachic digestion), and hence called “peptic” glands, 
while the others are concerned only in the elaboration of mucus. 
The relative distribution of these glands in different regions of the 
walls of the stomach varies greatly in different animals, and in 
many species there are large tracts of the mucous membrane which 
do not secrete a fluid having the properties of gastric juice, but 
often constitute more or less distinct cavities devoted to storing 


py 


Fia. 20.—Stomach and pancreas of the Genet. Posterior or dorsal surface. w, Esophagus ; 
8, pancreas ; pd, pancreatic duct; bd, biliary duct from the liver. (From Mivart, Proc. Zool. 
Soc. 1882, p. 305.) 
and perhaps softening or otherwise preparing the food for digestion. 
Sometimes there is a great ageregation of glands forming distinet 
thickened patches of the stomach wall, as in the Beaver and Koala, 
or even collected in pyriform pouches with a common narrow 
opening into the cavity, as in the Manatee and the curious African 
Rodent Lophiomys. The action of the gastric fluid is mainly 
exerted upon the nitrogenous elements of the food, which it 
dissolves and modifies so as to render them capable of undergoing 
absorption, effected partly by the blood-vessels of the stomach, 
although the greater part passes through the pylorus, an aperture 
surrounded by a circular muscular valve, into the intestinal canal. 
Here it comes in contact with the secretion of a vast number of 
small glands called the crypts of Lieberkuhn, somewhat similar 
to those of the stomach; and also of several special glands of a 
different character, namely, the small racemose, duodenal, or 


THE DIGESTIVE SYSTEM 59 


Brunner’s glands, the pancreas, and the liver; the position of the 
ducts of the two latter organs being indicated in Fig. 20. 

Intestinal Canal.—The intestinal canal varies greatly in relative 
length and capacity in different animals, and it also offers manifold 
peculiarities of form, being sometimes a simple cylindrical tube of 
nearly uniform calibre throughout, but more often subject to altera- 
tions of form and capacity in different portions of its course,—the 
most characteristic and constant being the division into an upper 
and narrower, and lower and wider portion, called respectively the 
small and the large intestine, the former being divided quite arbi- 
trarily and artificially into duodenum, jejun- 
um, and ileum, and the latter into colon and 
rectum. One of the most striking peculiari- 
ties of this part of the alimentary canal is 
the frequent presence of a diverticulum or 
blind pouch, the caput cecwm coli, as it was 
first called, a name generally abbreviated into 
“cecum,” situated at the junction of the 
large and the small intestine, a structure pre- 
senting an immense variety of development, 
from the smallest bulging of a portion of the 
side wall of the tube to a huge and complex 
sac, greatly exceeding in capacity the whole 
of the remainder of the alimentary canal. It 
is only in herbivorous animals that the caecum 
is developed to this great extent, and among 
these there is a curious complementary re- ; ' 

< a . : Fia, 21. — Diagrammatic 
lationship between the size and complexity pian of the general arrange- 
of this organ and that of the stomach. ment of the alimentary canal 
Where the latter is simple the cacum is i ® typical Mammal 
a sophagus; st, stomach; p, 
generally the largest, and vice versd. Both the pylorus; s,s, small intestine 
cecum and colon are often sacculated, a dis- (abbreviated); ¢, cwcum ; J, J, 
position caused by the arrangement of the ae oe 
longitudinal bands of muscular tissue in their ~ 
walls; but the small intestine is always smooth and simple-walled 
externally, though its lining membrane often exhibits various 
contrivances for increasing the absorbing surface without adding to 
the general bulk of the organ, such as the numerous small villi by 
which it’ is everywhere beset, and the more obvious transverse, 
longitudinal, or reticulating folds projecting into the interior, met 
with in many animals, of which the “ valvul conniventes” of Man 
form well-known examples. 

Besides the crypts of Lieberkuhn found throughout the in- 
testinal canal, and the glands of Brunner confined to the duodenun, 
there are other structures in the mucous membrane, about the 
nature of which there is still much uncertainty, called “solitary ” and 


60 GENERAL ANATOMICAL CHARACTER 


“agminated” glands ; the latter being more commonly known by the 
name of “Peyer’s patches.” These were formerly supposed to be 
secretory organs, which discharged some kind of fluid into the 
intestine, but are now more generally considered to belong to that 
group of structures of somewhat mysterious function of which the 
lymphatic and lacteal glands are members. The solitary glands are 
found scattered irregularly throughout the whole intestinal tract ; 
the agminated, on the other hand, are always confined to the small 
intestine, and are most abundant in its lower part. They are 
subject to great variation in number and in size, and even 
in different individuals of the same species, and also differ in 
character at different periods of life, becoming atrophied in old 
age. 

i Liver.—The distinct glands situated outside the walls of the 
intestinal canal, but which pour their secretion into it, are the 
pancreas and the liver. The latter is the more important on 
account of its size, if not on account of the direct action of its 
secretion in the digestive process. This large gland, so complex in 
structure and function, is well developed in all mammals, and its 
secreting tube, the bile-duct, always opens into the duodenum, or 
that portion of the canal which immediately succeeds the stomach. 
It is situated on the right side of the abdomen in contact with the 
diaphragm and the stomach, but varies greatly in relative size, and 
also in form, in different groups of mammals. In most mammals a 
gall-bladder, consisting of a pyriform diverticulum from the bile- 
duct, is present, but in many this appendage is wanting, and it is 
difficult to find the rationale of its presence or absence in relation 
to use or any other circumstance in the animal economy. 

The descriptions of the livers of various animals to be met 
with in treatises or memoirs on comparative anatomy are very 
difficult to understand for want of a uniform system of nomencla- 
ture. The difficulty usually met with arises from the circumstance 
that this organ is divided sometimes, as in Man, Ruminants, and 
the Cetacea, into two main lobes, which have been always called 
respectively right and left, and in other cases, as in the lower 
Monkeys, Carnivora, Insectivora, and several other orders, into a 
larger number of lobes. Among the latter the primary division usu- 
ally appears at first sight tripartite, the whole organ consisting of a 
middle, called “ cystic” or “suspensory ” lobe, and two lateral lobes, 
called respectively right and left lobes. This introduces confusion 
in describing livers by the same terms throughout the whole series 
of mammals, since the right and left lobes of the Monkey or Dog, 
for instance, do not correspond with parts designated by the same 
names in Man and the Sheep. There are, moreover, conditions 
where neither the bipartite nor the tripartite system of nomencla- 
ture will answer, so that we should have considerable difficulty in 


THE DIGESTIVE SYSTEM 61 


describing them without some more general system. In order to 
arrive at such a system it appears desirable to consider the liver in 
all cases as primarily divided by the umbilical vein (see Fig. 22, w) 
into two segments, right and left. This corresponds with its 
development and with the condition characteristic of the organ in 
the inferior classes of vertebrates. The situation of this division 
can almost always be recognised in adult animals by the persistence 
of some traces of the umbilical vein in the form of the round 
ligament, and by the position of the suspensory ligament. 

When the two main parts into which the liver is thus divided 
are entire, as in Man, the Ruminants, and Cetacea, they may be 
spoken of as the right and left lobes; when fissured, as the right 
and left segments of the liver, reserving the term lobe for the sub- 


Fia, 22,—Diagrammatic plan of the inferior surface of a multilobed liver of a Mammal. 
The posterior or attached border is uppermost. 2, Umbilical vein of the foetus, represented by 
the round ligament in the adult, lying in the umbilical fissure ; dv, the ductus venosus ; ve, 
the inferior vena cava ; p, the vena porte entering the transverse fissure ; Uf, the left lateral 
fissure ; rlf, the right lateral fissure ; cf, the cystic fissure ; 11, the left lateral lobe ; Ic, the left 
central lobe ; re, the right central lobe ; 71, the right lateral lobe ; s, the Spigelian lobe ; c, the 
caudate lobe ; g, the gall-bladder. 


divisions. This will involve no ambiguity, for the terms right and 
left lobe will no longer be used for divisions of the more complex 
form of liver. In the large majority of mammals each segment is 
further divided by a fissure, more or less deep, extending from 
the free towards the attached border, which are called right and 
left lateral fissures (Fig. 22, rif and Jif). When these are more 
deeply cut than the umbilical fissure (uv), the organ has that 
tripartite or trefoil-like form just spoken of, but it is easily seen 
that it is really divided into four regions or lobes, those included 
between the lateral fissures being the right and left central (rc and 
le) separated by the umbilical fissure, and those beyond the lateral 
fissures on each side being the right and left lateral lobes (71 and J/). 


62 GENERAL ANATOMICAL CHARACTERS 


The essentially bipartite character of the organ and its uniformity 
of construction throughout the class are thus not lost sight of, even 
in the most complex forms. The left segment of the liver is rarely 
complicated to any further extent, except in some cases by minor 
or secondary fissures marking off small lobules, generally inconstant 
and irregular, and never worthy of any special designation. On 
the other hand, the right segment is usually more complex. The 
gall-bladder, when present, is always attached to the under surface 
of the right central lobe, sometimes merely applied to it, in other 
cases deeply embedded in its substance. In many instances the 
fossa in which it is sunk is continued to the free margin of the 
liver as an indent, or even a tolerably deep fissure (cf). The 
portal fissure (p), through which the portal vein and hepatic artery 
enter and the bile-duct emerges from the liver, crosses the right 
central lobe transversely, near the attached border of the liver. 
The right lateral lobe always has the great vena cava (vc) either 
grooving its surface or tunnelling through its substance near the 
inner or left end of its attached border ; and a prolongation of this 
lobe to the left, between the vein and the portal fissure, sometimes 
forming a mere flat track of hepatic substance, but more often 
a prominent tongue-shaped process, is the so-called “Spigelian lobe” 
(s). From the under surface of the right lateral lobe a portion is 
generally partially detached by a fissure, and called the “caudate 
lobe” (c). In Man this lobe is almost obsolete, but in most 
mammals it is of considerable magnitude, and has very constant 
and characteristic relations. It is connected by an isthmus at the 
left (narrowest or attached) end to the Spigelian lobe, behind which 
isthmus the vena cava is always in relation to it, channelling 
through or grooving its surface. It generally has a pointed apex, 
and is deeply hollowed to receive the right kidney, to the upper 
and inner side of which it is applied. 

Considerations derived from the comparatively small and simple 
condition of the liver of the Ungulata, compared with its large 
size and complex form in the Carnivora, have led to the perhaps 
too hasty generalisation that the first type is related to a herbivorous 
and the latter to a carnivorous diet. The exceptions to such a 
proposition are very numerous. The fact of the great difference 
between the liver of the Cetacea and that of the Seals cannot 
be accounted for by difference of habits of life, though it perhaps 
may be by difference of origin. 


? For further details of these modifications, see Flower’s “Lectures on the 
Comparative Anatomy of the Organs of Digestion of the Mammalia,” Medical 
Times and Gazette, Feb.-Dec. 1872. 


CIRCULATORY AND RESPIRATORY SYSTEMS 63 


V. CIRCULATORY, ABSORBENT, RESPIRATORY, AND URINARY 
SYSTEMS. 


Blood.—The blood of mammals is always red, and during the 
life of the animal hot, having a nearly uniform temperature, 
varying within a few degrees on each side of 100° Fahr. The 
corpuscles are, as usual in the vertebrates, of two kinds: (1) 
colourless, spheroidal, nucleated, and exhibiting amceboid move- 
ments; while (2) the more numerous, on which depends the 
characteristic hue of the fluid in which they are suspended, are 
coloured, non-nucleated, flattened, slightly biconcave discs, with 
circular outline in all known species except the Camels and Llamas, 
where they have the elliptical form characteristic of the red 
corpuscles of nearly all the other vertebrates, though adhering to 
the mammalian type in the absence of nucleus and relatively small 
size. As arule they are smaller as well as more numerous than in 
other classes, but vary considerably in size in different species, and 
not always in relation to the magnitude of the animal; a Mouse, 
for instance, having as large corpuscles as a Horse. Within the 
limits of any natural group there is, however, very often some such 
relation, the largest corpuscles being found among the large species 
and the smallest corpuscles among the small species of the group, 
but even to this generalisation there are many exceptions. The 
transverse diameter of the red corpuscles in Man averages zy5q of 
an inch, which is exceptionally large, and only exceeded by the 
Elephant (z7;5), and by some Cetacea and Edentata. They are 
also generally large in Apes, Rodents, and the Monotremata, and 
small in the Artiodactyles, least of all in the Chevrotains (Tragulus), 
being in 7. javanicus and meminna not more than >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 <Allodon of the 
Upper Jurassic of the United States. 

The first discovery of the remains of Mesozoic mammals was 
made in the Keuper or Upper Trias of the Rhetian Alps in 
Bavaria. In 1847 Professor Pleininger of Stuttgart, while sifting 
some sand from the Keuper of Diegerloch and Steinenbronn, 
found, among an immense mass of teeth, scales, and unrecog- 
nisable fragments of skeletons of fish and saurians, two minute 
teeth, each with well-defined, enamelled, tuberculated crowns 
and distinct roots, plainly showing their mammalian character. 
These were considered by their discoverer to indicate a predaceous 
and carnivorous animal of very small size, to which he gave the name 
of Jicrolestes antiquus. Subsequently Mr. C. Moore discovered in a 
bone bed of Rhetie (topmost Trias) age, filling a fissure in the 
Mountain Limestone at Holwell, near Frome in Somersetshire, 
various isolated teeth with their crowns much worn, but apparently 
including both upper and lower molars and a canine, which are 
assigned by Sir R. Owen to Pleininger’s genus JJicrolestes, and 
described specifically as Jf mocrei. Under the name of Aypsi- 
prymnopsis rheticus, Professor Boyd Dawkins described a single tooth 
with two roots discovered in the Rhetic Marlstone at Watchet in 
Somersetshire. Sir R. Owen referred the latter tooth to Vicrolestes, 
and if its describer is right in regarding it as a much worn premolar 
of the type of those of Plagiaulax (Fig. 25) there would be evidence 
that Microlestes was closely allied to the latter, from the molars 
of which those of J/icrolestes are scarcely distinguishable. 

Plagiaulax, of the Dorsetshire Purbeck (Figs. 24, 25), is at once 
distinguished from Tritylodon by its secant premolars, which, as already 
mentioned, recall those of some of the J/wcrepodide, although readily 


112 GEOLOGICAL DISTRIBUTION 


distinguished by the convexity of the cutting edge and their oblique 
grooving. This remarkable and highly specialised type has been the 
occasion of one of the most interesting discussions on the inferences 
which may be drawn as to the affinities and habits of an otherwise 
unknown animal from the structure of a small portion of its organisa- 
tion which occurs in the annals of natural history—a discussion 
carried on with great ability, ingenuity, and wealth of illustration 
on both sides. Dr. Falconer maintained that it was more nearly 
allied to the Rat-Kangaroo (Potorous or Hypsiprymnus) than to any 
other existing form, and that, as it is known that these animals 
feed upon grass and roots, “it may be inferred of Plagiaulax that 
the species were herbivorous or frugivorous. I can see nothing in 
the character of their teeth,” he adds, “to indicate that they were 
either insectivorous or omnivorous.” Sir R. Owen, on the other 
hand, from the same materials came to the conclusion that ‘the 
physiological deductions from the above-described characteristics of 
the lower jaw and teeth of Plagiaulaz are that it was a carnivorous 
Marsupial. It probably found its prey in the contemporary small 
insectivorous mammals and Lizards, supposing no herbivorous form 
like Stereognathus to have co-existed during the Upper Oolitic 
period.” 

It is impossible here to give at any length the arguments by 
which these opposing views are respectively supported, but it may 
be indicated that the first-mentioned is strongly countenanced by 
the consideration of the following facts: (1) all existing Marsupials 
may be divided, so far as their dentition is concerned, into two 
groups—(a) those which have a pair of large more or less procumbent 
incisors close to the symphysis of the lower jaw, and rudimentary 
or no canines (diprotodont dentition), and (#) those which have 
numerous small incisors and large pointed canines (polyprotodont 
dentition) ; (2) the vast majority of the former group are purely 
vegetable feeders, and almost all of the latter are carnivorous or 
insectivorous ; and (3) Plagiaulaa, so far as its structure is known, 
shows an analogy with the former group; and, as we have no sure 
basis for inferences as to the habits of an unknown animal, but the 
knowledge of the habits of such as are known, we have no grounds 
for supposing that its habits differed from those forms having an 
analogous type of dental structure.1 

Allied types, such as Ctenacodon, are also met with in the Upper 


1 The whole discussion is contained in the following memoirs: (1) H. 
Falconer, ‘‘ Description of Two Species of the Fossil Mammalian genus 
Plagiaulax, from Purbeck,” Quart. Journ. Geol. Soc. vol. xiv. 1857 ; (2) B. Owen, 
art. ‘‘ Paleontology,” Encyclopedia Britannica, 8th ed., 1859; (3) H. Falconer, 
“On the Disputed affinity of the Mammalian genus Plagiaulax,” Quart. Journ. 
Geol. Soc. vol. xviii. 1862; (4) R. Owen, ‘‘ Monograph of the Fossil Mammalia 
of the Mesozoic Formation,” Paleontographical Society, 1871. 


MESOZOIC MAMMALS 113 


Jurassic of North America; and the Plagiaulucide also persisted 
into the lower part of the Eocene division of the Tertiary period ; 
Neoplagiaulax being a Tertiary form common to Europe and the 
United States, while Liotomus and Ptilodus are at present known 
only from the latter country. 

The present group is also represented in the upper Cretaceous 
of the United States by Selenacodon (Meniscoéssus in part), Cimoliomys, 
etc. Polymastodon, of the Lowest or Puerco Eocene of New Mexico 
is the largest known form, and is characterised by the presence 
of only one premolar and the elongated molars. The angle of 
the mandible is inflected after the Marsupial fashion. 

Polyprotodont Types.—The second type of mammalian dentition 
found in the Mesozoic period resembles that occurring among 
recent Polyprotodont Marsupials—that is to say there are at 
least three lower incisors, the canines are well developed, and the 
premolars and molars are cuspidate, the number of the latter reach- 
ing in some cases to seven or eight. There has been much dis- 
cussion as to the taxonomic position of these forms, and while the 
majority of writers admit the Marsupial affinities of at least a 
moiety, it has been contended that others indicate distinct ordinal 
groups more or less closely allied to the Insectivora. At present, 
however, there is no decisive evidence to support such a view. 
Important proof of the Marsupial affinity of one of these forms is 
afforded by the replacement of the teeth, which appears to be of the 
same nature as in the existing Marsupials, that is to say, the last 
premolar alone is preceded by a milk-tooth. 

The most generalised forms appear to be Dromatherium and 
Microconodon, from Lower Mesozoic beds in the United States, of 
which enlarged views of the teeth are given in Fig. 4 (1, 2), p. 
31. Professor Osborn points out the extremely simple character of 
these teeth, and it is quite possible that these forms may prove 
to be Prototheria. There are three premolars and seven molars in 
the lower jaw of Dromatherium. 

A common form in the Purbeck of Dorsetshire is Triconodon 
(Triacanthodon), in which the formula of the lower teeth is 23, ¢ 1, 
p 4, m 3-4. A lower jaw is shown in 
Fig. 28, and an enlarged view of a molar 
tooth in Fig. 4 (5). The molar teeth con- 
sist of three flattened cones placed in the 
same antero-posterior line, those of the ; 

. . . . Fic. 28.—Reversed view of the 
upper and lower jaw being alike. Pria- left ramus of the mandible of 
codon, of the Jurassic of the United States, Triconodon mordax ; from the 
is probably inseparable from Triconodon. ac ieee Oa Aauatal 
In the genus Phascolotherium (Fig. 29) of ~~ ‘ 
the Lower Jurassic Stonesfield Slate, the lower teeth may be 
classified as 7 4,¢ 1, p 3, m 4, the premolars and molars being 


1i4 GEOLOGICAL DISTRIBUTION 


much alike. The molars approximate to the type of those of 
Triconodon, but the anterior and posterior cones are relatively 
smaller. Like that of the last-named genus, the mandible of 


Fria. 29.—Inner view of the right ramus of the mandible of Phascolotherium buchklandi ; 
from the Stonesfield Slate. The outline shows the natural size. i, Incisors (one missing) ; ¢, 


canine; p, premolars; m, molars. The mylohyoid groove is seen near the lower border. (After 
Owen.) 


Phascolotherium is remarkable for the extremely low position of 
its articular condyle. In Amphilestes (Fig. 30) of the Stonesfield 
Slate the molars appear to be of the same general type as those 
of Phascolotherium, but are more numerous, although their exact 
number cannot be determined. A somewhat different type 
of lower molar is displayed by the genus Aimblotherium, of the 
Dorsetshire Purbeck, to which dmphitherium of the Stonesfield Slate 
was probably allied. This type of tooth is shown in Fig. 4 (8, 9, 
12) p. 31, and, as there stated, represents that modification of the 
tritubercular type known as the tubercular sectorial. The three 
primitive tritubercular cusps form what is known as the blade of 
the tooth, behind which 
there is the talon or 


Pp hypocone. A similar 
\o\1234 5 Bi 2 
1\ wets form of molar oceurs 


3 


Na 
eh in the existing Opos- 
sums and Bandicoots. 


Fia. 30.—Reversed inner view of the left ramus of the 
mandible of Amphilestes broderipi; from the Stonesfield 
Slate. Twice natural size. The restoration of the anterior 
teeth is conjectural, and the condyle is placed too high. 


The number of lower 
teeth in .fmblotherium 
ist 4, ec 1, p 4, m 


7-8. Numerous allied 
types, such as fchyro- 
don and Dryolestes occur in the Upper Jurassic of Europe or the 
United States, while from only one side of the jaw being exposed 
in each ease so-called genera like Séylodon and Stylacodon have been 
formed upon specimens showing the opposite side to that which 
is exposed in the types of Amblotherium and Amphitherium. The 


(After Owen.) 


TERTIARY MAMMALS 115 


only parallel among existing forms to the excessive number of 
molar teeth found in these Mesozoic genera occurs in the Mar- 
supial genus Jfyrmecobius, of which a description is given in a 
succeeding chapter. Jaws more or less closely resembling those 
described under the names mentioned above are also found in 
the uppermost Cretaceous of the United States. A feature com- 
mon to these Mesozoic mammals and J/yrmecobius and some other 
existing forms is the presence of a narrow channel on the inner 
side of the mandibular ramus known as the mylohyoid groove 
(Fig. 29). 

The last type of molar dentition occurring among the Mesozoic 
Mammalia is that found in the 
lower jaws (Fig. 31), upon which 
the genus Spalacotherium was 
established, the upper jaws, 
described as Peralestes, being 


Fic, 31.—Part of the left ramus of the man 
apparently referable to the same dible, viewed from the outer side, of Spala- 


animal. Upper and lower teeth cotherium tricuspidens; from the Purbeck of 


: . Swanage, Twice natural size. (After Owen. 
of this form are represented in Giramnge. Thane namineL Sze. CREM OWE) 


Fig. 4 (6, 7), p. 31, where they are described as typical examples 
of the tritubercular type of molars, the upper teeth having one 
inner and two outer cusps, and the reverse condition obtaining in 
the lower ones. This type of molar presents a marked resemblance 
to that found in the existing Insectivorous genus Chrysochloris ; the 
number of lower teeth in Spalacotherium is, however, 1 3, ¢ 1, 
p+m 10, by which it is widely distinguished from all the Insect- 
ivora. j/enacodon, of the Upper Jurassic of the United States, 
appears to be allied to Spalacotherium. 

Tertiary Mammals.—The more important types of Tertiary 
mammals will, as already mentioned, be noticed under the heads 
of the groups to which they are severally allied; but a few general 
remarks on this subject may be advantageously recorded in this chap- 
ter. In the first place, it may be observed that the comparatively 
scanty evidence of mammalian life hitherto yielded by the Cretaceous, 
coupled with the number and variety of forms approximating to 
the existing groups found even in the lowest Tertiary, indicates a 
great imperfection of the geological record. At present, indeed, 
we have no decisive evidence of the existence of any members of 
the Eutherian subclass previously to the Tertiary; but it can hardly 
be doubted that in some part of the world they had made their 
appearance before that epoch. The Eutherian mammals of the 
lowest Eocene, both in Europe and the United States, are of an 
extremely generalised type; and although many of them approximate 
to existing groups, they show such a combination of characters, now 
restricted to individual groups, as to indicate that several of the 
various orders into which the subclass is now divided were at that 


116 GEOGRAPHICAL AND GEOLOGICAL DISTRIBUTION 


period very intimately connected. A marked feature of these 
early Eutherians is the prevalency of trituberculism in the dentition, 
not less noteworthy being the frequent occurrence of pentadactylism 
- in the feet, while many of the individual bones were devoid of the 
grooves and ridges found in those of later types. By the time 
that we reach the upper division of the Eocene period, such as the 
horizon of the well-known gypsum of the Paris basin, nearly all the 
chief groups of mammals had become clearly differentiated from 
one another, although their representatives were usually more 
generalised than their existing allies. From this date to the later 
geological periods there is a gradual approximation to the types of 
mammalian life existing at the present day. 

In addition to the features of trituberculism and pentadactyl- 
ism so characteristic of the oldest known Eutherians, we may notice 
some other points in connection with the earlier types. Thus the 
older Tertiary mammals, as we have already stated, had relatively 
smaller and simpler brains than the later types, so that a gradual 
evolution in this respect may be traced from the Eocene to the 
Pleistocene. Again, there is a great tendency among the Eocene 
forms to a retention of the typical Eutherian dental formula noticed 
on page 25, and also to the absence of an interval, or diastema, in 
the dental series. Concomitantly with this feature we may notice 
the short crowns and simpler structure of the molar teeth of the 
earlier Ungulates as compared with those of to-day, of which details 
will be given in a later chapter. Another instance of the more 
generalised characters of the earlier mammals is afforded by the 
absence or slight development of horns, antlers, and tusks among 
the Ungulata. Thus the earlier Rhinoceroses were hornless, and 
the Deer either without antlers or with antlers of a very simple 
kind, while the male Swine were not furnished with the formidable 
tusks of the existing Wild Boars. Finally, all, or nearly all of the 
mammals, from the lowest Eocene of Rheims present the pecu- 
larity of having a vertical perforation in the astragalus. 

The intimate connection existing during the Middle Tertiary 
between many families of mammals now widely distinguished from 
one another may be more conveniently noted when we come to the 
consideration of the families in question. 


CHAPTER V 
THE SUBCLASS PROTOTHERIA OR ORNITHODELPHIA 


General Characters The characters of the Prototheria can at 
present only be deduced from the two existing families, since 
hitherto no extinct animals which can be referred with certainty 
to other divisions of this remarkable and well-characterised group 
have been discovered. These two isolated forms, in many respects 
widely dissimilar, yet having numerous common characters which 
unite them together and distinguish them from the rest of the 
Mammalia, are the Ornithorhynchide and the Echidnide, both re- 
stricted in their geographical range to the Australian region of the 
globe. Taken altogether they represent the lowest type of evolution 
of the mammalian class, and most of the characters in which they 
differ from the other two subclasses tend to connect them with the 
inferior, vertebrates, the Sauropsida and Amphibia; for, though 
the name Ornithodelphia owes its origin to the resemblance of the 
structure of the female reproductive organs to those of birds, there 
is nothing especially bird-like about them. 

Their principal distinctive characters are these. The brain has 
a very large anterior commissure, and a very small corpus callosum, 
agreeing exactly in this respect with the Marsupials. The cerebral 
hemispheres, in Echidna at least, are well developed and convoluted 
on the surface. The auditory ossicles present a low grade of de- 
velopment, the malleus being very large, the incus small, and the 
stapes columelliform. The coracoid bone is complete, and articu- 
lates with the sternum, and there is a precoracoid (epicoracoid) in 
advance of the coracoid, while there is also a large “ interclavicle” 
or episternum in front of the sternum, and connecting it with the 
clavicles. There are also “epipubic” bones. The oviducts (not 
differentiated into uterine and Fallopian portions) are completely 
distinct, and open, as in oviparous vertebrates, separately into a 
cloacal chamber, and there is no distinct vagina. The testes of 
the male are abdominal in position throughout life, and the vasa 


118 MONOTREMATA 


deferentia open into the cloaca, not into a distinct urethral passage. 
The penis, attached to the ventral wall of the cloaca, is perforated 
by a canal in the greater part of its length, and not merely grooved, 
as in reptiles and those birds which have such an organ. The 
canal is open at the base and brought only temporarily in contact 
with the termination of the vasa deferentia, so as to form a seminal 
urethra when required ; but it never transmits the urinary secretion. 
This condition is a distinct advance on that of the Sauropsida in 
the direction of the more complex development of these parts in 
most of the other Mammalia. The ureters do not open into the 
bladder, but behind it into the dorsal wall of the genito-urinary 
passage. The mammary glands have no distinct nipple, but pour 
out their secretion through numerous apertures situated in a cup- 
shaped depression of the abdominal skin, forming a mammary 
marsupium, especially developed in the females during lactation. 
It should be mentioned that, according to the observations of Pro- 
fessor Gegenbaur, the mammary glands of the Monotremes are the 
simplest found in the entire class. The region of the glands is, 
indeed, distinguished from the rest of the abdomen merely by its 
thicker layers of muscles. The glands themselves are closely con- 
nected with the hair-follicles, and belong to the sudoriparous type, 
whereas the glands of all other mammals are of sebaceous origin. 

The young are produced from eggs laid by the female parent, 
which are meroblastic, like those of birds; that is to say only a 
portion of the yolk segments and forms the embryo, the remainder 
serving for the nourishment of the latter. 

The above are the principal distinguishing characters of the 
group, and apply not only to the subclass, but of course equally to 
the one order Monotremata, in which the two existing genera are 
included. In addition to these more important characters, the 
following minor features may also be mentioned. 

The scapula differs from that of all other mammals in that the 
ridge corresponding to the spine of other forms is situated on the 
anterior border instead of in the middle of the outer or dorsal surface. 
The humerus is much expanded at its two extremities, and has a very 
prominent deltoid crest, and a well-marked entepicondylar foramen. 

The dorso-thoracic vertebree are nineteen in number, and have 
no terminal epiphyses to their bodies. The tranverse processes of 
the cervical vertebre are of autogenous formation, and remain 
suturally connected with the remainder of the vertebra until the 
animal is full-grown. ‘Though in this respect they present an 
approximation to the Sauropsida (Reptiles and Birds), they differ 
from these classes, inasmuch as there is not a gradual transition from 
these autogenous transverse processes of the neck (or cervical ribs, 
as they may be considered) into the thoracic ribs, for in the seventh 
vertebra the costal element is much smaller than in the others, 


ORNITHORHYNCHIDAZ 119 


indicative of a very marked separation of neck from thorax, not 
seen in the existing Sauropsida. The upper ends of the ribs 
are attached to the sides of the bodies of the dorsal vertebre 
only, and not to the transverse processes. The sternal ribs are 
well ossified, and there are distinct partly ossified intermediate ribs. 
The cerebral cavity, unlike that of the lower Marsupials or the 
Reptiles, is large and hemispherical, flattened below and arched 
above, and about as broad as long. The cribriform plate of the 
ethmoid is nearly horizontal. The cranial walls are very thin, and 
smoothly rounded externally, and the sutures become completely 
obliterated in adult skulls, as in Birds. The broad occipital region 
slopes upwards and forwards, and the face is produced into a long 
and depressed rostrum. The bony palate is prolonged backwards, 
so that the posterior nares are nearly on a level with the glenoid 
fosse. The mandible is without distinct ascending ramus; the 
coronoid process and angle are rudimentary, and the two halves are 
loosely connected at the symphysis. The fibula has a_ broad, 
flattened process, projecting upwards from its upper extremity 
above the articulation, like an olecranon. In the male there is an 
additional, flat, curved ossicle on the hinder and tibial side of the 
plantar aspect of the tarsus, articulating chiefly to the tibia, which 
supportsin theadult a sharp-pointed perforated horny spur, with which 
is connected the duct of a gland situated beneath the skin of the back 
of the thigh, the function of which is not yet clearly understood. (A 
rudimentary spur is found in the young female Ornithorhynchus, but 
this disappears when the animal becomes adult.) The stomach is 
sub-globular and simple ; the alimentary canal has no ileo-czecal valve, 
or marked distinction between large and small intestine, but has a 
small, slender vermiform cecum with glandular walls. The liver 
is divided into the usual number of lobes characteristic of the 
Mammalia, and is provided with a gall-bladder. 

In the presence of three distinct bones developed from cartilage 
in the shoulder-girdle (viz. scapula, coracoid, and pre- or epi-coracoid) 
the Monotremes agree with the Anomodont reptiles (see p. 83), 
and with no other représentatives of that class. The precoracoid 
of the Anomodonts is, however, distinguished by extending upwards 
to articulate with the acromial process of the scapula. The 
Monotreme humerus is, moreover, strikingly like the corresponding 
bone of many of the Anomodonts and of some of the allied 
Labyrinthodont Amphibians. 


Family ORNITHORHYNCHIDA. 


Ornithorhynchus.\—Cerebral hemispheres smooth. Premaxille 
and mandible expanded anteriorly and supporting a horny beak 


1 Blumenbach, Voigts Magazin, vol. ii. p. 205 (1800). 


120 MONOTREMATA 


something like that of a duck, bordered by a naked and‘very sensitive 
membranous expansion. The place of teeth in the adult is supplied 
functionally by horny structures, elongated, narrow, and sharp- 
edged, along the anterior part of the sides of the mouth, and broad, 
flat-topped or molariform behind. Functional molar teeth present 
in the young and adolescent condition. Legs short, fitted for 
swimming ; feet webbed, each with five well-developed toes armed 
with large claws, beyond which in the fore feet the interdigital 
membrane is extended. Vertebre: C 7, D 17, L 2,8 2, Ca 21. 
Acetabulum not perforated. Tongue not extensile. Mucous mem- 
brane of small intestine covered with delicate, close-set transverse 
folds or ridges. Tail rather short, broad, and depressed. Eyes 
very small. Fur close and soft. 

The Duck-billed Platypus (Platypus anatinus) was the name 
assigned to one of the most remarkable of known animals by 
Shaw, who had the good fortune to introduce it to the notice 
of the scientific world in the Naturalists Miscellany (vol. x., 1799). 
In the following year it was independently described by Blumenbach 
(Voigts Magazin, ii. p. 205) under the name of Ornithorhynchus 
paradoxus. Shaw’s generic name, although having priority to that 
of Blumenbach, could not be retained, as it had been used at 
a still earlier time (1793) by Herbst for a genus of Coleoptera. 
Ornithorhynchus is therefore now universally adopted as the scien- 
tific designation, although Duck-billed Platypus or Duck-bill may 
be conveniently retained as a vernacular appellation. By the 
colonists it is called “ Water-Mole,” but it need scarcely be said, 
its affinities with the true moles are of the slightest and most 
superficial description. Until the last few years the early stages 
of the development of the young were not fully known. It had, 
indeed, been repeatedly affirmed, in some cases by persons who 
have had actual opportunities of observation, that the Platypus lays 
eggs ; but these statements were generally received with scepticism 
and even denial. This much-vexed question was, however, settled 
by the researches of Mr. W. H. Caldwell in 1884, who found that 
these animals, although undoubtedly mammals throughout the 
greater part of their structure, are oviparous, laying eggs, which in 
the manner of their development bear a close resemblance to the 
development of those of the Reptilia. Two eggs are produced at 
a time, each measuring about three-fourths of an inch in its long, 
and half an inch in its short, axis, and enclosed in a strong, flexible, 
white shell. 

The Platypus is pretty generally distributed in situations 
suitable to its aquatic habits throughout the island of Tasmania 
and the southern and eastern portions of Australia. Slight variations 
in the colouring and size of different individuals have given rise to 
the idea that more than one species may exist; but all naturalists 


ORNITHORHVNCHIDZ 121 


who have had the opportunity of investigating this question by the 
aid of a good series of specimens have come to the conclusion that 
there is but one, and no traces of any extinct allied forms have yet 
been discovered. 

The length of the animal when full grown is from eighteen to 
twenty inches from the extremity of the beak to the end of the tail, 
the male being slightly larger than the female. The fur is short, 
dense, and rather soft to the touch, and composed of an extremely 
fine and close under-fur, and of longer hairs projecting beyond 
this, each of which is very slender at the base, and expanded, 


Fic, 32.—Platypus or Duck-bill (Ornithorhynchus anatinus), From Gould's Mammals of 
Australia. 


flattened, and glossy towards the free end. The general colour is 
deep brown, but paler on the under parts. The tail is short, broad, 
and depressed, and covered with coarse hairs, which in old animals 
generally become worn off from the under surface. The eyes are 
small and brown. There is no projecting pinna or ear-conch. The 
mouth, as is well known, bears a striking resemblance to the bill of 
a Duck. \It is covered with a naked skin, a strong fold of which 
projects outwards around its base. The nostrils are situated near 
the extremity of the upper surface. There are no true teeth in the 
adult, but their purposes are served by horny prominences, or 
cornules, two on either side of each jaw—those in the front narrow, 
longitudinal, sharp-edged ridges, and those behind broad, flattened, 


122 MONOTREMATA 


and molariform. The upper surface of the lateral edges of the, 
mandible has also a number of parallel fine transverse ridges, like 
those on the bill of a Duck. Until 1888 it was thought that true 
teeth were totally wanting throughout the life of this animal ; but in 
the spring of that year Mr. E. B. Poulton! announced the discovery 
in an embryo of teeth which were regarded as quite functionless. In 
the following year, however, Mr. O. Thomas ? was fortunate enough 
to find some young skulls with functional teeth in situ, and was thus 
enabled to give a detailed account of their structure and of their 
relations to the cornules. From these specimens it appears that 
the teeth are functional for a considerable part of the life of the 
animal, cutting the gum in the usual manner, and, after being worn 
down by friction with food and sand, are shed from the mouth 
in the same manner as are the milk-teeth of other mammals. The 
cornules are developed from the epithelium of the mouth under and 
around the teeth, and the hollows found in the middle of them are 
the vestiges of the alveoli from which the teeth have been shed. 
One of the skulls showed on either side, both above and below, two 
completely calcified teeth ; but in another example there were three 
teeth on either side of the lower jaw. According to Mr. Thomas’s 
account, ‘the teeth themselves are broad, flat, and low-crowned. 
The upper ones have each two high, conical, internal cusps, from 
which minute ridges run downwards and outwards to the outer 
borders of the crowns, where the edge is peculiarly crenulate rather 
than cuspidate, in the ordinary sense of the word. On the whole, 
the anterior and posterior upper teeth are essentially similar to one 
another, except that the former are narrower, and their outer edges 
are less markedly crenulated. In the lower jaw there is a greater 
difference between the two. The anterior is triangular in outline, 
its longest side is placed antero-externally, and its anterior and 
postero-external angles have each a high pointed cusp, ridged on 
its internal aspect, while the posterior and internal borders are 
indistinctly crenulated. The posterior tooth is broadly quadrangular 
in outline, with a projecting antero-internal angle. As in the cor- 
responding tooth above, there are two cusps on one side, and a series 
of crenulations on the other, but they are of course reversed, the 
cusps being external and the crenulations internal, The cusps are 
high, and connected with transverse ridges running across towards 
the internal border.” 

In trying to find any teeth like those of the Duck-bill amone 
other known mammals Mr. Thomas considers, as was first suggested 
by Professor Cope, that those of the Mesozoic Multituberculata (p.109) 
make the nearest approximation. He adds, however, that “it must 
be insisted that the resemblance between the Multituberculate 

1 Proceedings of the Royal Society of London, vol. xliii, p. 358 (1888), 
2 bid. vol. xlvi. p. 126 (1889). 


ORNITHORHYNCHIDA 123 


and the Ornithorhynchus teeth is of the most general character, 
and that the two are certainly widely separated generically, even if 
we do admit that they appear to possess a relationship nearer to 
each other. than to any other known groups of mammals.” 

Reverting to the description of the Duck-bill, we find that in 
the cheeks are tolerably capacious pouches, which appear to be used 
as receptacles for food. The limbs are strong and very short, each 
with five well-developed toes provided with strong claws. In the 
fore feet the web not only fills the interspaces between the toes, but 
extends considerably beyond the ends of the long,“broad, and some- 
what flattened nails, giving great expanse to the foot when used for 
swimming, though capable of being folded back on the palm when 
the animal is burrowing or walking on the land. On the hind foot 
the nails are long, curved, and pointed, and the web extends only 
to their base. On the heel of the male is a strong, curved, sharply 
pointed, movable horny spur, directed upwards and backwards, 
attached by its expanded base to the accessory bone of the tarsus. 
This spur, which attains the length of nearly an inch, is traversed 
by a minute canal, terminating in a fine longitudinal slit near 
the point, and connected at its base with the duct of a large gland 
situated at the back part of the thigh. The whole apparatus is so 
exactly similar in structure to the poison-gland and tooth of a 
venomous snake as to suggest a similar function, but evidence that 
the Platypus ever employs its spur as an offensive weapon has, at 
all events until lately, been wanting. A case is, however, related 
by Mr. Spicer in the Proceedings of the Royal Society of Tasmania 
for 1876 (p. 162), of a captured Platypus inflicting a severe wound by 
a powerful lateral and inward movement of the hind legs, which wound 
was followed by symptoms of active local poisoning. It is not improb- 
able that both the inclination to use the weapon and the activity of the 
secretion of the gland may be limited to the breeding season, and 
that their purpose may be, like that of the antlers of deer and 
many similar organs, for combat among the males. In the young 
female the spur is present in a rudimentary condition, but it dis- 
appears in the adult of that sex. 

The Platypus is aquatic in its habits, passing most of its time in 
the water or close to the margin of lakes and streams, swimming 
and diving with the greatest ease, and forming for the purpose of 
sleeping and breeding deep burrows in the banks, which generally 
have two orifices—one just above the water level, concealed among 
long grasses and leaves, and the other below the surface. The 
passage at first runs obliquely upwards in the bank, sometimes to 
a distance of as much as fifty feet, and expands at its termination 
into a cavity, the floor of which is lined with dried grass and 
leaves, and in which the eggs are laid and the young brought up. 
The food consists of aquatic insects, small crustaceans, and worms, 


124 MONOTREMATA 


which are caught under water, the sand and small stones at the 
bottom being turned over with the bill. The creatures appear 
at first to deposit what they have thus collected in their cheek 
pouches, and when these are filled they rise to the surface and 
quietly triturate their meal with the horny plates before swal- 
lowing it. Swimming is effected chiefly by the action of the 
broad forepaws, the hind feet and tail taking little share in 
locomotion in the water. When asleep they roll themselves into 
a ball, as shown in the figure. In their native haunts they are 
extremely timid and wary, and very difficult to approach, being 
rarely seen out of their burrows in the daytime. Mr. 
Crowther, who has supplemented the often quoted observations 
of Dr. Bennett upon the habits of these animals in confinement, 
says, ‘They soon become very tame in captivity ; in a few days 
the young ones appeared to recognise a call, swimming rapidly 
to the hand paddling the water; and it is curious to see their 
attempts to procure a worm enclosed in the hand, which they 
greedily take when offered to them. I have noticed that they 
appear to be able to smell whether or not a worm is contained in 
the closed hand to which they swim ; for they desisted from their 
efforts if an empty fist was offered.” When irritated they utter a 
soft low growl, resembling that of a puppy. 


Family ECHIDNIDZ. 


Cerebral hemispheres larger and well convoluted. Facial portion 
of skull produced into a long, tapering, tubular rostrum, at the 
end of which the anterior nares are situated. Rami of mandible 
slender, styliform. Opening of mouth small, and placed below the 
extremity of the rostrum. No teeth or laterally placed horny plates, 
though the palate and tongue are furnished with spines. Tongue 
very long, vermiform, slender, and protractile. Lining membrane 
of small intestine villous, but without transverse folds. Feet not 
webbed, but with long strong claws fitted for scratching and 
burrowing. The hinder feet with the ends of the toes turned 
outwards and backwards in the ordinary position of the animal 
when on the ground. Tail very short. Acetabulum with a large 
perforation, as in Birds. Calcaneal spur and gland of the male 
much smaller than in Ornithorhynchus. Fur intermixed with strong, 
sharp-pointed spines. Terrestrial and fossorial in habits, feeding 
exclusively on ants, and recalling in the structure of the mouth and 
various other parts relating to their peculiar mode of life the true 
Anteaters of the order Edentata. 

The Echidnas or Spiny Anteaters constitute a family which 
appears in some respects to be less specialised than the Ornitho- 
rhynchide. According to Mr. O. Thomas, all the living forms may 


ECHIDNIDE 125 


be included in two species, which, with some hesitation, are referred 
to two genera—Hechidna and Proechidna (Acanthoglossus). 

Echidna.i—In Echidna there are five toes, all of which are 
provided with claws, those of the fore feet being broad, slightly 
curved, and directed forwards, while the posterior ones are slender, 
more curved, and inclined outwardly. The beak is about as long 
as the rest of the head, and either nearly straight, or slightly curved 
upwards, while the palate is comparatively wide, and but slightly 
vaulted. The number of the vertebre is C 7,D 16, L 3,8 3, Ca 12. 
The one existing representative of the genus (Z. aculeata) occurs in 
New Guinea, Tasmania, and Australia. 

So much variation is displayed by this animal, that it has been 
divided into several species, but the latest researches tend to show 
that these variations cannot be regarded as indicating more than 
races, of which there are three well-marked types. 

The first race, or variety, has been termed the Port Moresby 
Echidna, and is only known from that Papuan locality. It is 
distinguished from the typical form by its smaller size, by the 
shorter spines on the back, which admit of the fur being seen, and 
by the more spinous covering of the head, belly, and limbs, as well 
as by the lighter skull and relatively larger beak. 

The typical variety is confined to the Australian mainland, and 
is of medium size. The spines of the back are very long and stout, 
often reaching a length of two inches, and almost completely con- 
cealing the hair. The colour of these spines varies from yellow at 
the roots to black at the tips, but some may be altogether yellow. 
The hair of the back is black or dark brown in colour, but it may 
be occasionally absent, or in the region of the loins may exceed the 
spines in length. The limbs and under surface of the body are 
covered with dark brown hair, thinly interspersed with short spines ; 
and the hair of the face is of the same general hue as that of the 
body. The skull has a slender rostrum and a flat and narrow 
brain-case. 

In the third or Tasmanian race, which is confined to Tasmania, 
the average size is somewhat larger than in the typical form. ‘The 
most characteristic feature is, however, the shortness of the spines 
of the back, which in the greater part of that region are almost or 
quite concealed by the hairs. The hairs of the back are dark 
brown, those of the under surface and sides of the head being 
generally rather paler. There is often a white spot on the chest. 
Very frequently there is a difference in the proportionate lengths 
of the hinder claws from those of the typical race. In the skull 
the beak is comparatively short and stout, and the brain-case large 
and wide. 

Echidnas are usually found in rocky districts, and more especially 


1 Cuvier, Tableau Elémentaire @ Hist. Nat. p. 148 (1798). 


126 MONOTREMATA 


in the mountains. In a wild state they live mainly on ants. Speci- 
mens have been brought to this country and kept in the Zoological 
Society’s Gardens ; and in captivity they will readily eat eggs, and 
bread-and-milk. They are able, however, to endure long fasts, an 
individual having been known to go without food for upwards of a 
month. 

These animals seem to be mainly of nocturnal habits, and if 
brought out during the day-time appear to be sluggish and stupid, 
crouching to the ground with the head between the legs, and thus 
presenting a mass of spines to an enemy. They burrow rapidly in 
soft ground, sinking directly downwards, and not going head for- 
wards. A specimen placed on a large chest of earth containing 
plants reached the bottom in less than two minutes; and it is said 
that the muzzle assists in the work of burrowing. 

Procchidna.1—The one known representative of the genus 
Proechidna (Fig. 33) attains dimensions about equal to those of 


Fia. 33.—The Three-toed Echidna (Procchidna bruijnii). From Gervais. 


the largest race of Echidna aruleata. The skull is less depressed 
than in the latter, with the anterior portion of the palate very 
concave, and the deflected beak nearly twice the length of the 
remainder of the skull. As a rule, there are only three claws to 
each foot; but the first and fifth digits are represented by several 
phalanges, and one instance is known where there are five complete 
claws on the anterior and four on the posterior feet. There are 
two more vertebre in the dorsal and lumbar region than in 
Echidna. 

The head and body are covered with a thick coat of hair 
among which there are a number of short spines in the region of 
the back, which are much less numerous than in the typical race of 
the last species. The colour of the fur is generally dark brown or 
black, but the head may be almost white; and the spines are 
usually entirely white, although in certain cases they may be brown 
at the root. 

* Gervais, Ostéographie des Monotremes, p. 43 (1877). 


ECHIDNIDA 127 


This species is known only from New Guinea, the recorded 
specimens being from the north-western regions of that country. It 
inhabits rocky ground, and dwells chiefly in the mountains, the 
specimens which were first described having been obtained at an 
elevation of about 3500 feet above the sea level. The Papuans capture 
it by digging trenches in the ground to a depth of about a yard, by 
which means they generally come upon its runs. 

Fossil Species.—Remains of a species of Echidna of very much 
larger size than the existing forms have been obtained from the 
cave-deposits of New South Wales, which appear to be of Pleisto- 
cene age. This species was named Echidna oweni by the late Mr. 
Krefft, but was subsequently called £. ramsayi by Sir R. Owen. 
In referring this species to the genus Echidna, that term must be 
regarded as including Proechidna. 


CHAPTER VI 
THE SUBCLASS METATHERIA OR DIDELPHIA 


General Characters.—The Metatheria or Didelphia are represented at 
present by numerous species, presenting great diversities of general 
appearance, structure, and habits, although all united by many 
essential anatomical and physiological characters, which, taken 
altogether, give them an intermediate position between the Proto- 
theria and the Eutheria. 

Although the striking differences in external form, in many 
anatomical characters, and in mode of life of various animals of this 
section might lead to their division into groups equivalent to the 
orders of the Eutheria, it is more convenient on the whole to adhere 
to the usual custom of treating them all as forming one order called 
Marsvupia.i4,! the limits of which are therefore equivalent to that 
of the subclass. The more essentially distinctive characters are as 
follows. 

In the structure of the brain and the presence of epipubic bones 
they agree with the Prototheria, while in the structure of the ear- 
bones and the shoulder-girdle and the presence of teats on the 
mammary glands they resemble the Eutheria, the reproductive 
organs belonging to neither one nor the other type, but having a 
special character representing an intermediate grade of develop- 
ment. The ureters open into the base of the bladder. The 
oviducts are differentiated into uterine and Fallopian portions, and 
open into a long and distinct vagina, quite separate from the cystic 
urethra. The penis is large, but its crura are not directly attached 
to the ischia. The spongy body has a large bifurcated bulb. The 
young are born in an exceedingly rudimentary condition, and are 
never nourished by means of an allantoic placenta, but are trans- 
ferred to the nipple of the mother, to which they remain firmly 


1 For the detailed characters of all the genera and species of Marsupials the 
reader should consult the British Museum Catalogue of Marsupialia and Mono- 
tremata, by Oldfield Thomas, 1888. 


GENERAL CHARACTERS 129 


attached for a considerable time, nourished by the milk injected 
into the mouth by compression of the muscle covering the 
mammary gland. ‘They are therefore the most typically mam- 
malian of the whole class. The nipples are nearly always concealed 
in a fold of the abdominal integument or “pouch” (marsupium) 
which serves to support and protect the young in their early 
helpless condition. 

Entering more fully into the characters of the subclass, which 
are also those of the order Marsupialia, it may be observed that the 
brain is generally small in proportion to the size of the animal, and 
the surface-folding of the cerebral hemispheres, though well marked 
in the larger species, is never very complex in character, and is 
absent in the medium-sized and smaller species. The arrangement 
of the folding of the inner wall of the cerebrum differs essentially 
from that of all known Eutheria, the hippocampal fissure being 
continued forward above the corpus callosum, which is of very 
small size. The anterior commissure is, on the other hand, greatly 
developed. 

The teeth are always divisible, according to their position and 
form, into incisors, canines, premolars, and molars; but they vary 
much in number and character in the different families. Except in 
the genus Phascolomys, the number of incisors in the upper and 
lower jaws is never equal. The true molars are very generally four 
in number on either side of each jaw. The chief peculiarity in the 
dentition lies, however, in the mode of succession. Thus there is no 
vertical displacement and succession of the teeth, except in the case 
of a single tooth on either side of each jaw, which is always the 
hindermost of the premolar series, and is preceded by a tooth 
having more or less of the characters of a true molar (see Fig. 34); 
this deciduous tooth 
being the only one 
comparable to the 
“milk-teeth ” of the 
diphyodont Eu- 
theria. In some 
cases (as in Poto- 
vous) this tooth re- 


tains its place and 

a = Fic. 34.—Teeth of upper jaw of Opossum (Didelphys mar- 
function until the supialis), all of which are unchanged, except the last premolar, 
animal has nearly, the place of which is occupied in the young animal by a molari- 
if not quite, attained form tooth, represented in the figure below the line of the other 


its full stature, and 

is not shed and replaced by its successor until after all the other 
teeth of the permanent series, including the posterior molars, are 
fully in place and use. In others, as the Thylacine, it is very 
rudimentary in form and size, being shed or absorbed before any 


130 METATHERIA 


of the other teeth have cut the gum, and therefore quite function- 
less. It must further be noted that there are some Marsupials, 
as the Wombat, Myrmecobius, and the Dasyures, in which no such 
milk-tooth, even in a rudimentary state, has yet been discovered, 
possibly in some cases from want of materials for observation at 
the right stage of development. 

Epipubic or marsupial bones are present in both sexes of nearly 
all species. In one genus alone, Thylacinus, they are not ossified. 
The number of dorso-lumbar vetebre is always nineteen, although 
there are some apparent exceptions caused by the last lumbar being 
modified into a sacral vertebra. The number of pairs of ribs is 
nearly always thirteen. The tympanic bone remains permanently 
distinct. The carotid canal perforates the basisphenoid. The 
lachrymal foramen is situated upon or external to the anterior margin 
of the orbit, and there are generally large vacuities in the bony 
palate. The angle of the mandible is (except in Tarsipes) more or 
less inflected. The hyoid bones have always a peculiar form, 
consisting of a small, more or less lozenge-shaped basi-hyal, broad 
cerato-hyals, with the remainder of the anterior arch usually 
unossified, and stout, somewhat compressed thyro-hyals. There are 
two anterior vene cave,' into each of which a “vena azygos” 
enters. In the male the testes are always contained in a scrotum, 
which is suspended by a narrow pedicle to the abdomen in front of 
the penis. The vasa deferentia open into a complete and continuous 
urethra, which is also the passage by which the urine escapes from 
the bladder, and is perfectly distinct from the passage for the feces, 
although the anus and the termination of the urethro-sexual canal 
are embraced by the same sphincter muscle. The glans is often 
bifurcated anteriorly. In the female the oviducts never unite to 
form a common cavity or uterus, but open separately into the 
vagina, which at least for part of its course is double. The 
mamme vary much in number, but are always abdominal in 
position, having long teats, and in most of the species are more 
or less enclosed in a fold of the integument forming a pouch 
or marsupium, though in some this is entirely wanting, and the 
newly-born, blind, naked, and helpless young, attached by their 
mouths to the teat, are merely concealed and protected by the 
hairy covering of the mother’s abdomen. In this stage of their 
existence they are fed by milk injected into their stomach by the 
contraction of the muscles covering the mammary gland, the 
respiratory organs being modified temporarily, much as they are 
permanently in the Cetacea—the elongated upper part of the 
larynx projecting into the posterior nares, and so maintaining a free 
communication between the lungs and the external surface 


1 Except in Petawrus (Belideus) breviceps (Forbes, Proc. Zool. Soc. 1881, 
p. 188). 


DISTRIBUTION 131 


independently of the mouth and gullet, thus averting the danger of 
suffocation while the milk is passing down the latter passage. 
Distribution.—The existing species of Marsupials are, with the 


Fic. 35.—Front view of skull of Surcophilus ursinus, showing polyprotodont and carnivorous 
dentition (Quart. Journ. Geol. Soc. vol. xxiv. p. 313). 


exception of one family (the Didelphyide), limited in geographical 
region,! 


distribution to the 
mammalian fauna of Australia, 
New Guinea, and some of the 
adjacent islands. The Didel- 
phyide are almost purely Neo- 
tropical, one or two species 
ranging northwards into the 
Nearctic region. Fossil re- 
mains of members of this 
family have also been found in 
Europe and America in strata 
of the Eocene and early Mio- 
cene periods ; and it is probable 
that at least many of the poly- 
protodont Mesozoic mammals 
noticed in Chapter IV. are 
referable to the Marsupialia. 
Classification.—In dividing 
the Marsupials into minor 
groups, it may be observed 
that one of the most obvious 
distinctive characters among 


Australasian 


forming the chief 


Nun 


Fig. 36.—Front view of skull of Koala (Phas- 


colarctus cinercus), showing diprotodont and 
herbivorous dentition (Quart. Journ. Geol. Soc. 
vol. xxiv. p. 313). 


them is derived from the form and arrangement of the teeth. 


1 Including the transitional Austro-Malayan region. 


132 MARSUPIALIA 


In certain species, as the Opossums, Dasyures, and Thylacine, 
the incisors are numerous, small, and subequal in size, and the 
canines large, as in the typical placental Carnivores (Fig. 35). 
To these the term “polyprotodont” is applied, and they are all 
more or less carnivorous in their habits. In others the central 
incisors are very prominent, and the lateral incisors and canines 
absent or subordinate in function (Fig. 36). These are called 
“ diprotodont,” and they are all wholly or in great part vegetable 
feeders. In one group of these, the Wombats, there are but two 
incisors above and the same number below; but all the others, in- 
cluding the Kangaroos, Koalas, and Phalangers, have two functional 
incisors below and as many as six above, three on each side, but 
of these the first or central pair is the most fully developed. 

Some hesitation has frequently been expressed as to whether the 
Polyprotodont and Diprotodont types are entitled to constitute 
distinct primary groups, owing to the presence of syndactylism 
among the Peramelide in the former, as well as in the latter ; but if 
Mr. O. Thomas is right in regarding this feature as acquired 
independently in the two groups we may safely adopt such a 
division. Taking various combinations into consideration, the 
existing Marsupials readily group themselves into six very natural 
families, the leading characters of which may be summarised as 
follows :— 


Order MARSUPIALIA. 


A, Potyproropontia.—Incisors numerous, small, subequal. Canines 
larger than the incisors. Molars with sharp cusps. 


a. Incisors 3. Hind feet with the four outer toes subequal, 


distinct, and a well-developed opposable hallux. Didel- 
phyide. 

. Incisors $. Hind feet with four outer toes distinct. Hallux 
small or rudimentary, rarely opposable. Dasyuride. 


(4—5) 
= 


y. Incisors Hind feet long and narrow. Fourth toe 


larger than the others. Hallux rudimentary or absent. 
Second and third toes very slender, and united in a 
common integument (syndactylous). Peramelide. 


B. Diproropontia.—Incisors not exceeding $, usually 3, but occasion- 
ally +. Central (first) upper and lower incisors large and 
cutting. Upper canines generally, and lower invariably, absent 
or small. Molars with bluntly tuberculated or transversely 
ridged crowns. 


a. Teeth with persistent pulps. Incisors 4, large, scalpriform, 
with enamel on the outer surface only. No canines. 
Hind feet with four subequal outer toes, partially 


syndactylous, and with rudimentary hallux. Phascolo- 
myidee. 


DIDELPHVIDA 133 


8. Teeth rooted. Three upper incisors and acanine. Hind 
limbs not disproportionately large. Feet syndactylous, 
broad, with four subequal outer toes, and a large 
opposable hallux. Phalangeride. 

y. Teeth rooted. Three upper incisors, and frequently a 
canine. Hind limbs disproportionately large, with 
syndactylous feet as in Peramelide. Macropodide. 


Suborder POLYPROTODONTIA. 


The leading characters of this group are given in the foregoing 
schedule. This group is the only one represented at the present 
day, and so far as we know also in past epochs, beyond the confines 
of the Australasian region and adjacent islands. 


Family DIDELPHYID. 


Dentition: i $,¢ 4, p 3, m 4; total 50. Incisors very small 
and pointed. Canines large. Premolars with compressed pointed 
crowns. Molars with numerous sharp cusps. The last premolar 
preceded by a deciduous multicuspidate milk-molar, which remains in 
place until the animal is nearly adult (Fig. 34). Limbs of moderate 
development, each with five complete and distinct toes, all of which 
are provided with short, compressed, 
curved, sharp claws of nearly equal 
size, except the first toe of the hind 
foot or hallux (Fig. 37), which is large, 
widely separable from the others, to 
which it is opposed in climbing, and 
terminates in a dilated rounded ex- 
tremity, without a nail. Tail gener- 
ally long, partially naked and prehen- 
sile. Stomach simple. Czcum of 
“small or moderate size. Pouch gener- 
ally absent, sometimes represented by 
two lateral folds of the abdominal 
integument, partially covering the 
teats, rarely complete. Vertebre : 
C7, D 13,L 6,8 2, C 19-35. 

The Didelphyide, or true Opos- 
sums, differ from all other existing Fic. 37.Skeleton of the right hind 
Marsupials in their habitat, being feotof the Virginian Opossum (Didelphys 

: : . marsupialis). 
peculiar to the American continent. 
They are mostly carnivorous or insectivorous in their diet, and 
arboreal in habits. 

Opossums occur throughout the greater part of the American 


134 MARSUPIALIA 


continent, ranging from the United States to Patagonia, the greater 
number of species being found in the warmer regions. In South 
America the opossums take the place of the Eutherian Insectivora, 
and the sharp cusps on their teeth are admirably adapted for crushing 
the insects on which they mainly subsist. 

Chironectes.\—The family comprises two genera only, namely 
Didelphys, containing all the species, with the exception of the curious 
Yapock, which forms by itself the genus Chironectes, and is distin- 
guished from all other Opossums by its webbed feet, non-tuberculated 
soles, and peculiar coloration. Its ground colour is light gray, with 
four or five sharply-contrasted brown bands passing across its head 
and back, and thus giving it a very peculiar mottled appearance. 
It is almost wholly aquatic in its habits, living on small fish, 
crustaceans, and water insects. Its range extends from Guatemala 
to southern Brazil. 

Didelphys.2—The type genus Didelphys is a very large one, con- 
taining, according to Mr. O. Thomas, twenty-three existing species. 
It may be divided into five groups, or sub-genera, all of which have 
received distinct names. The typical group is represented only by 
the common or Virginian Opossum (J. marsupialis), of which the 
numerous varieties have received separate specific names. This 
species is of large size, with a long, scaly, prehensile tail, and long 
bristle-like hairs mingled with the fur. The pouch is complete. 
It ranges over all temperate North America, and is also found in 
central and tropical South America, where it is commonly known 
as the Crab-eating Opossum. This animal is extremely common, 
being even found living in the towns, where it acts as a scavenger 
by night, retiring for shelter by day upon the roofs of the houses or 
into the sewers. The female produces in the spring from six to 
sixteen young ones, which are placed in her pouch immediately 
after birth, and remain there until able to take care of them- 
selves. 

The second or Metachirine group includes three species found 
all over the tropical parts of the New World. They are of medium 
size, with short close fur, very long, scaly, and naked tails, and 
less developed ridges on their skulls than in the type species. As 
a rule there is no pouch adapted to carry the young, which 
commonly ride on their mother’s back, holding on by winding 
their prehensile tails round hers. The Philanderine group is 
closely allied to the preceding, but is readily distinguished by the 
woolly hair, and the brown streak down the middle of the face. 
The Woolly Opossum (D. lanigera), which is represented in the 
accompanying woodcut (Fig. 38) carrying its young in the fashion 
mentioned above, is one of the two species of this group. In the 

* Hliger, Prod. Syst. Mami. et Aves, p. 76 (1811). 
* Linn. Syst, Nat. Ed. 12, vol. i. p. 71 (1766). 


DIDELPHVIDE 135 


fourth or Jficowreine group the numerous species are all smaller 
than in the preceding groups, and have short and close hair, and 
no dark streak down the face. The best known species is the 
Murine Opossum (D. murina), little larger than a House-Mouse, 
and of a bright red colour, which is found as far north as central 
Mexico, and extends thence right down to the south of Brazil. The 
last or Peramyne group contains several extremely shrew-like 
species, of very small size, with short, hairy, and usually non-pre- 
hensile tails, not half the length of the trunk, and with wholly 
unridged skulls. The most striking member of the group is the 
Three-striped Opossum (D. americana), from Brazil, which is of a 
reddish-gray colour, with three clearly-defined deep-black bands 


Fia. 38.—The Woolly Opossum (Didelphys lanigera). 


down its back, very much as in some of the striped mice of 
Africa. 

The numerous fossil species of Opossum found in the Upper 
Eocene and Lower Miocene of Europe are of especial interest from a 
distributional point of view, since they indicate how the Opossums of 
America may have been connected with the Australian Marsupials. 
These forms were originally referred to Didelphys, but have been 
subsequently described as Peratherium and Amphiperatherium. The 
characters of the molar teeth on which these genera are hased do 
not appear to be sufficiently important to justify their separation 
from Didelphys. Allied forms occur in the Tertiaries of North 
America, which were originally described under the name of Her- 
petotherium, but have been subsequently referred to Peratherium. 
Remains of many of the existing species of Opossum are found in 
a fossil condition in the Pleistocene cave-deposits of Brazil. 


136 MARSUPIALIA 


Funily DASYURID 


Dentition : i4,¢4,p and m numerous, variable. Incisors small ; 
canines well developed ; molars with pointed cusps. Limbs equal. 
Fore feet with five subequal toes terminating in claws. Hind feet 
with the four outer toes well developed, and distinct from each 
other and bearing claws; the first (or hallux) clawless, generally 
rudimentary, sometimes entirely wanting. Stomach simple. No 
cecum. Predatory carnivorous or insectivorous animals, inhabit- 
ants of Australia, Tasmania, and the southern parts of New Guinea 
and some of the adjacent islands. The aberrant genus J/yrimecobius, 
though clearly a member of this family, is so sharply distinguished 


Fic. 39.—The Thylacine (Thylacinus cynocephalus). 


from all the others as to render a division into two subfamilies 
necessary. 

Subfamily Dasyurinss.—This comprises the more typical Dusy- 
uride, in which the premolars and molars never exceed the normal 
number of seven on either side of each jaw, and in which the tongue 
is not specially extensile. 

Thylacinus..—Dentition : 24, ¢ }, $3, m4=46. Incisors small, 
vertical, the outer one in the upper jaw larger than the others. 
Summits of the lower incisors, before they are worn, with a deep 
transverse groove dividing them into an anterior and a posterior cusp, 
Canines long, strong, and conical. Premolars separated from one 
another by intervals, with compressed crowns, increasing in size 
from before backwards. True molars in general characters ye- 


1 Temminck, Monographies de Mammaloyie, vol. i, p. 60 (1827). 


DASVURIDE 137 


sembling those of Dasywrus, but of more simple form, the cusps 
being not so distinct nor sharply pointed. Milk-molar very small, 
and shed before the animal leaves the mother’s pouch. Humerus 
with an entepicondylar foramen. General form very Dog-like. 
Head elongated. Muzzle pointed. Ears moderate, erect, triangular. 
Fur short and closely applied to the skin. Tail of moderate length, 
thick at the base and tapering towards the apex, clothed with short 
hair. Hallux (including the metacarpal bone) wanting. Vertebre : 
C7, D138, L6, 82, C23. Marsupial bones represented only by 
small unossified fibro-cartilages. 

The only known existing species of this genus, 7. cynocephalus 
(Fig. 39), though smaller than a common Wolf, is the largest preda- 
ceous Marsupial at present living. It is now entirely confined to the 
island of Tasmania, although fragments of bones and teeth found in 
caves afford evidence that a closely allied species once inhabited the 
Australian mainland. The general colour of the Thylacine is 


Fic. 40.—Right lateral aspect of the skull of the Thylacine. 


grayish brown, but it has a series of transverse black bands on the 
hinder part of the back and loins, whence the name of “Tiger” 
frequently applied to it by the colonists. It is also called “ Wolf,” 
and sometimes, though less appropriately, “Hyzena.” Owing to 
the havoc it commits among the sheepfolds, it has been nearly 
exterminated in all the more settled parts of Tasmania, but still 
finds shelter in the almost impenetrable rocky glens of the more 
mountainous regions of the island. The female produces four 
young at a time. The pouch opens backwardly, and there are four 
mamme. The figure of the skull exhibits the peculiar Dog-like 
form so characteristic of the genus. 

Sarcophilus..—Dentition: 1 4,¢4, p2, m4. Upper incisors nearly 
equal, and placed vertically, the first not differentiated from the 
rest. Premolars rounded and closely crowded between the canine 
and molars, with broad crowns; molars broad and heavy, the last 
one without a distinct hind talon. Form thick and powerful ; 


1 F, Cuvier, Hist. Nat. des Mammiferes, iv. (1837). 


138 MARSUPIALIA 


head disproportionately large for the body; muzzle short and 
broad; ears broad and rounded; tail of moderate length, and 
evenly hairy. Hallux wanting ; soles of feet naked, without defined 
pads. Humerus with entepicondylar foramen. 

This genus is now represented only by a single species 
(S. ursinus) found in Tasmania, where, from its ferocious and des- 
tructive habits, it is commonly known under the name of the “ Devil.” 
A front view of the skull is shown in Fig. 35. 

The prevailing colour of this animal is black, and the size about 
equal to that of an English Badger ; its habits are fossorial, and it 
is very destructive to sheep. On account of the similarity in the 
number of its teeth this genus has been generally included in the 
next one, but in the structure of the teeth it is much nearer to 
Thylacinus. An extinct species is found in the Pleistocene deposits 
of the mainland of Australia. 

It may be observed that the two premolars missing from the 
typical series of four in this and the next genus are the second and 
the fourth; the fourth milk-molar being likewise absent. In 
Thylacinus and other Polyprotodonts with three premolars it is the 
second that is missing. 

Dasywrus..—Dentition: 1 4,¢4, p 2, m 4; total 42. Upper 
incisors nearly equal, and placed vertically ; first slightly longer, 
narrower, and separated from the rest. Lower incisors sloping 
forwards and upwards. Canines large and sharply pointed. Pre- 
molars with compressed and sharp-pointed crowns, and slightly 
developed anterior and posterior accessory basal cusps. True 
molars with numerous sharp-pointed cusps. In the upper jaw the 
first three with crowns having a triangular oral surface, the fourth 
small, simple, narrow, and placed transversely. In the lower jaw 
the molars more compressed, with longer cusps; the fourth not 
notably smaller than the others. Form viverrine. Ears long and 
narrow, prominent, and obtusely pointed. Hallux rudimentary, or 
absent ; its metatarsal bone always present. Tail long and well 
clothed with hair. Humerus without an entepicondylar foramen. 
Vertebre : C7, D13, L6, S 2, C 18-20. 

The Dasyures are small Civet-like animals with a gray or brown 
pellage profusely spotted with white; they are mostly inhabitants 
of the Australian continent and Tasmania, where in the economy of 
nature they take the place of the smaller predaceous Carnivora, the 
Cats, Civets, and Weasels of other parts of the world. They hide 
themselves in the daytime in holes among rocks or in hollow trees, 
but prowl about at night in search of the small living mammals 
and birds which constitute their prey. The species are not numer- 
ous, and include D. maculatus, about the size of a common Cat, 
inhabiting Tasmania and the southern part of Australia 3 D. viver- 


1 Geoffroy, Bull. Soc. Philom. vol. i. p. 106 (1796). 


DASVURIDE 139 


rinus, Tasmania and Victoria; D. geoffroyi, nearly all Australia ; 
D. hallucatus, North Australia; D. albopunctatus, New Guinea. 

Remains referred to D. viverrinus occur in the Australian Pleis- 
tocene deposits. 

Phascologale1—This genus comprises a considerable number of 
small Marsupials, none of them exceeding a common Rat in size, 
differing from the Dasyures in possessing an additional pre- 
molar—the dentition being 7 $,¢4+,p3, m4; total 46,—and having 
the teeth generally developed upon an insectivorous rather than a 
carnivorous pattern, the upper middle incisors being larger and 
inclined forwards, the canines relatively smaller, and the molars 
with broad crowns, armed with prickly tubercles. The muzzle is 
pointed. Ears moderately rounded and nearly naked. Feet broad 
and short. Fore feet with five subequal toes, having compressed, 
slightly curved, pointed claws. Hind feet with the four outer toes 
subequal, having claws similar to those in the fore feet; the hallux 
always distinct and partially opposable, though small and nailless. 
Tail long, very variable in its covering, being either bushy, crested, 
or nearly naked. Pouch represented merely by a few folds of skin. 
Mamme varying from four to ten in number. The food of these 
animals is almost entirely insects ; some species pursuing their prey 
among the branches of trees, while others are purely terrestrial. 
They are found throughout Australia, and also in New Guinea and 
the Aru and some of the adjacent islands. 

P. cristicaudata, a species with a thick compressed tail orna- 
mented upon its apical half with a crest of black hair, differs from the 
others by the very reduced size of the fourth premolar in the upper, 
and its complete absence in the lower jaw, thus forming an interest- 
ing transition in dentition towards Dasyurus. It constitutes the 
genus Cheetocercus of Krefft, but is included by Mr. O. Thomas in 
Phascologale, the frequent absence of the fourth lower premolar in 
P. thorbeckiana indicating that the total absence of this tooth in the 
known specimens of this species cannot be regarded as of generic 
importance. All the members of this and the two following genera 
can be at once distinguished from Dasyurus by the absence of white 
spots on the fur. 

Sminthopsis.2— The genus Sminthopsis includes several small 
species allied to Phascologale but characterised by the narrowness 
of the hind foot, and by the soles of the feet being either granulated 
or hairy, instead of naked. 

Antechinomys.2—The last genus of the Dasyurine is Antechinomys, 
represented only by 4. laniger of Queensland and New South Wales. 
This elegant little mouse-like creature, which has large oval ears and 

1 Temminck, Monographies de Mammalogiec, vol. i. p. 56 (1827). 


2 Thomas, dann. Mus. Genov. ser. 2, vol. iv. p. 503 (1887). 
3 Krefft, Proc. Zool. Soc. 1866, p. 434. 


140 MARSUPIALIA 


a long tail with the terminal part bushy, is distinguished from 
Sminthopsis by the absence of the hallux and the great elongation 
of the limbs. The tympanic bull of the skull are also unusually 
large, with the mastoid portion much swollen. A full account of 
the habits and anatomy of this animal, which appears to be of very 
rare occurrence, is given in the Proc. Zool. Soc. 1880, p. 454. 

Subfamily Myrmecobiinze.—Molars and premolars exceeding 
the normal number of seven on each side. Tongue, long cylindrical, 
and extensile. 

Afyrmecobius.\—Dentition : i 4,¢4, p 2 


,m£or &; total 52 or 56, 


Fic. 41.—Myrmecobius fusciatus. From Gould. 


being the largest number of teeth in any existing Marsupial. The 
distinction between the molars and premolars is founded not on 
a knowledge of the succession of the teeth, but on their form. The 
teeth are all small and (except the four posterior inferior molars) 
separated from each other by an interval. Head elongated, but 
broad behind. Muzzle long and pointed. Ears of moderate size, 
ovate, and rather pointed. Fore feet with five toes, all having 
strong, pointed, compressed claws, the second, third, and fourth 
nearly equal, the fifth somewhat, and the first considerably, shorter. 
Hind feet with no trace of hallux externally, but the metatarsal bone 


1 Waterhouse, Proc. Zool. Suv. 1836, p. 69. 


PERAMELIDA 141 


present. Tail long, clothed with long hairs. Fur rather harsh and 
bristly. Female without any pouch, the young when attached to 
the nipples being concealed only by the long hair of the abdomen. 
Vertebre: C 7, D 13, L 6, 8 3, C 23. A gland on the under 
surface of the body just in advance of the sternum. 

Of this singular genus but one species is known, M. fasciatus 
(Fig. 41), found in western and southern Australia. It is about the 
size of an English squirrel, to which animal its long bushy tail 
gives it some resemblance; but it lives entirely on the ground, 
especially in sterile, sandy districts, feeding on ants. Its pre- 
vailing colour is chestnut-red, but the hinder part of the back 
is elegantly marked with broad, white, transverse bands on a dark 
ground. 

The special interest of this form lies in its apparent relationship 
to those Mesozoic mammals which possess a large number of true 
molars (see p. 114); and it is suggested by Thomas that it may 
eventually be found advisable to include some of the latter in the 
present subfamily. 


Family PERAMELIDA. 


(4—5) 1 3 
g Pale 
small, with short broad crowns. Lower incisors moderate, nar- 
row, proclivous. Canines well developed. Premolars compressed, 
pointed. Molars with quadrate tuberculated crowns. Fourth pre- 
molar preceded by a small molariform tooth, which remains in place 
until the animal is nearly full grown. Fore feet with two or 
three of the middle toes of nearly equal size, and provided 
with strong, sharp, slightly curved claws; the other toes rudi- 
mentary. Hind feet long and narrow; the hallux rudimentary 
or absent; the second and third toes very slender, and united in a 
common integument; the fourth very large, with a stout elongated 
conical claw ; the fifth smaller than the fourth (see Fig. 43). The 
ungual phalanges of the large toes of both feet cleft at their ex- 
tremities (as in Manis among the Edentata, but in no other 
Marsupials). Head elongated. Muzzle long, narrow, and pointed. 
Stomach simple. Czcum of moderate size. Pouch complete, 
opening backwards. Alone among Marsupials they have no clavicles. 
The Peramelide form a very distinct family, in some respects 
intermediate between the sarcophagous Dasyuride and the 
phytophagous Macropodide. In dentition they resemble the former, 
but they agree with the latter in the peculiar structure of the hind 
feet. In the construction of the fore feet they differ from all other 
Marsupials. 
The Bandicoots, as these Marsupials are popularly termed, are 


m : ; total 46 or 48. Upper incisors 


Dentition : 7 


142 MARSUPIALIA 


of fossorial habits, and subsist either on an insectivorous or omni- 
vorous diet. It has been generally considered that their syndac- 
tylous feet indicate direct affinity with the Diprotodonts, but owing 
to the essentially Polyprotodont character of the organisation— 
which extends even to their carpal and tarsal bones—Thomas 
dissents from this view, and concludes that their syndactylism is an 
independently acquired character, and that they are really a direct 
offshoot from the Dasyuride. Some individuals are remarkable for 
the presence of a longitudinal groove in the root of the canines, by 
which feature they approximate to some of the Mesozoic Polypro- 
todont forms. They may be divided into three genera. 
Perameles.1— Anterior and posterior extremities not differing 
greatly in development. Fore feet with the three middle toes well 


Fic, 42.—Perameles gunni. From Gould. 


developed, the third slightly larger than the second, the fourth 
somewhat shorter, provided with long, strong, slightly curved, 
pointed claws. First and fifth toes very short and without claws. 
Hind feet with hallux of one or two phalanges, forming a distinct 
tubercle visible externally ; the second and third toes very slender, 
of equal length, joined as far as the ungual phalanges, but with 
distinct claws ; the fifth intermediate in length between these and 
the largely developed fourth toe. Ears of moderate or small size, 
ovate, pointed. Tail rather short, clothed with short adpressed 
hairs. Fur short and harsh. Vertebre ; C 7, D 13, L 6,8 1,C 17. 
Skull long and narrow, with the bulla single, and its mastoid portion 
not inflated. 

The animals of this genus are all small, and live entirely on the 
ground, making nests composed of dried leaves, grass, and sticks in 

’ Geoffroy, Bull. Soc. Philom. vol. iii, p. 249 (1803). 


PERAMELIDZ 143 


hollow places. They are rather mixed feeders; but insects, worms, 
roots, and bulbs constitute their ordinary diet. The various species 
are widely distributed over Australia, Tasmania, New Guinea, and 
several of the adjacent islands, as Aru, Kei, and New Ireland. The 
best known are—P. gunni (Fig. 42), bougainvillei, nasuta, obesula, and 
macrura from Australia, and P. doreyana, raffrayana, and longicaudata 
from New Guinea. 

Remains apparently referable to existing species are found in 
the cave-deposits of New South Wales. 

Peragale1—Molar teeth curved, typically with longer crowns 
and shorter roots than in the last. Hinder extremities proportionally 
longer, and hallux without claw. Muzzle much elongated and 
narrow. Fur soft and silky. lars very large, long, and pointed. 
Tail long, its apical half clothed on the dorsal surface with long 
hairs which form a crest. Vertebre: C7, D 13, L 6,8 2, C 23. 
Skull distinguished from that of Perameles by the large size and 
double structure of the auditory bulla, of which the mastoid portion 
is inflated. There is also an abrupt contraction of the muzzle at 
the third premolar. 

The type species of Rabbit- Bandicoot (P. lagotis), as these 
animals are called, is found in Western Australia, and also occurs 
fossil in the cave-deposits of New South Wales. It is the largest 
member of the family, being about the size of the common Rabbit, 
to which animal it bears sufficient superficial resemblance to have 
acquired the name of “Native Rabbit” from the colonists. It 
burrows in the ground, but in other respects resembles the true 
Bandicoots in its habits. 

The smaller P. leucura has short-crowned molars, with distinct 
cusps, which are almost obsolete in the type species. 

Cheropus.*—Dentition generally resembling that of Perameles, 
but the canines are less developed, and in the upper jaw two-rooted. 
Limbs very slender ; posterior nearly twice the length of the anterior. 
Fore feet with the functional toes reduced to two, the second and 
third, of equal length, with closely united metacarpals and short, 
sharp, slightly curved, compressed claws. First toe represented by 
a minute rudiment of a metacarpal bone ; the fourth by a metacarpal 
and two small phalanges without a claw, and not reaching the 
middle of the metacarpal of the third ; fifth entirely absent. Hind 
foot (Fig. 43) long and narrow, mainly composed of the strongly 
developed fourth toe, terminating in a conical pointed nail, with a 
strong pad behind it; the hallux absent or represented by a rudi- 
mentary metatarsal ; the remaining toes completely developed, and 
with claws, but exceedingly slender; the united second and third 
reaching a little way beyond the metatarso-phalangeal articulation of 

1 Gray, in Grey’s Australia, vol. ii. p. 401 (1841). 
2 Ogilby, Proc. Zool. Soc, 1838, p. 25. 


144 


MARSUPIALIA 


the fourth ; the fifth somewhat shorter. 
the body, and covered with short hairs forming a slight crest. 


Fic. 43.—Skele- 
ton of right hind 
foot of Cheropus 
castanotis. c, Cal- 
caneum ; a, astra- 
galus ; cb, cuboid ; 
n, navicular ; c3, 
ectocuneiform ; IT 
and III, the con- 
joined second and 
third digits; IV, 
the large and only 
functional digit ; 
V, the rudiment- 
ary fifth digit. 


equal, stout, and short. 


Tail not quite so long as 
Ears 
large and pointed, and folded down when the animal 
is at rest. Fur soft and loose. Vertebre: C 7, D 
13, L 6,81, C 20. Skull short and wide, with a 
small and single bulla, and a contraction of the 
muzzle at the third premolar. 

The only known species of this genus (Fig. 44), 
chiefly remarkable for the singular construction of 
its limbs, is an animal about the size of a small 
Rat, found in the interior of the Australian continent. 
Its general habits and food appear to resemble those 
of the other Peramelide. It was first described as 
C. ecaudatus by Ogilby from a mutilated specimen, 
but the specific name was afterwards changed, as being 
inappropriate, by Gray to castanotis. 


Suborder DIPROTODONTIA. 


For the leading characters of this group, see 
page 132. 


Fumily PHASCOLOMYID 


Dentition: ¢ 4,72, 94,m4=24. Allthe teeth 
with persistent pulps. The incisors large, scalpriform, 
with enamel only on the front surface, as in the 
Rodentia. The molars strongly curved, forming from 
the base to the summit about a quarter of a circle, 
the concavity being directed outwards in the upper 
and inwards in the lower teeth. The first of the 
series, or premolar, appears to have no milk-prede- 
cessor, and is single-lobed; the other four composed 
of two lobes, each subtriangular in section. Limbs 
Fore feet with five distinct toes, each 


furnished with a long, strong, and slightly curved nail, the first and 
fifth considerably shorter than the other three. Hind feet with a very 
short nailless hallux, the second, third, and fourth toes partially 
united by integument, of nearly equal length, the fifth distinct 
and rather shorter ; all four provided with long and curved nails, 
In the skeleton of the foot, the second and third toes are distinctly 
more slender than the fourth, showing a slight tendency towards 
the peculiar character so marked in the next two families. Tail 
rudimentary. Stomach simple, provided with a special gland 
situated near the cardiac orifice. Caecum very short, wide, and with 
a peculiar vermiform appendage. Pouch present. The auditory 
bulla of the skull are imperfect, open behind, with their anterior 


PHASCOLOM VIDE 145 


wall formed by a descending process of the squamosal, instead of the 


Fic, 44.—Chwropus castunotis, From Gould. 


alisphenoid. Masseteric fossa of mandible with a perforation and 
a deep pit. 


Fic. 45.—Common Wombat (Phascolomys ursinus). 


PhascolomysA—The existing Wombats (Fig. 45) comprise three 
1Geoffroy, dan. du Muséum, vol. ii. p. 365 (1803). 


10 


146 ILARSUPLALLA 


species, all of whieh are included in the one genus Phascolonus, 
and all of which date from the Pleistocene. 

In the typical group we find the following characters, luv 
rough and coarse, Kars short and rounded. Mufile naked. Post- 
orbital process of the frontal bone obsolete. Ribs fifteen pars. 
Vertebre: G7,D15,L 4,8 4, 10-12. The Wombat of Tas- 
mania and the islands of Bass’s Straits (2) wrsivus) and the closely 
similar but larger animal of the southern portion of the mainland of 
Australia (2. witchelli) belong to this group. 

In the second group the characters are as follows. Fur smooth 
and silky. Ears large and more pointed.  Mutlle hairy. Frontal 
region of skull broader than ino the other group, with well- 
marked postorbital processes. Ribs thirteen.  Vertehre: C7, D 
13, Lb 6,8 4, C 15-16. One species, 2. latifrons, the Hairy -nosed 
Wombat of Southern Australia, 

In their general form and actions the Wombats resemble small 
bears, having a somewhat similar shallling manner of walking, but 
they are still shorter in the legs, and have broader, latter backs Chan 
bears. They live entirely on the ground, or in burrows or holes 
among voeks, never climbing trees, and feed entirely on) grass, 
roots, and other vegetable substances. ‘They sleep during the day, 
and wander forth at might in search of food, and are shy and 
gentle in their habits generally, though they can bite strongly when 
provoked. The only noise the common Wombat makes is a low 
kind of hissing, but the Hairy-nosed Wombat is said to emit a short 
quick grunt when annoyed. The prevailing colour of the last- 
named species, as well as of 7. vesieus of Tasmania, is a brownish 
gray. The large wombat of the mainlund is very variable in colour, 
some individuals being found of a pale yellowish brown, others 
dark gray, and some quite black. The length of head and body is 
about three fect. 

It is noteworthy that Po mitehelli was first’ described) from the 
evidence of fossil remains, the living form subsequently deseribed as 
P. platyrhinus being found to be indistingnishable. Other extinet 
species oceur in the Pleistocene of Austratia. 

Phaseolones.$—Rennins of a large extinet Wombat, whieh must 
have nearly equalled the dimensions of a ‘Tapir, occur in the 
Pleistocene of Queensland, and have been described as Phaseolonis, 
It is probable that the expanded and thittened upper incisors from 
the same deposits upon the evidence of which the presumed genus 
Neceparnodon was founded, are likewise referable to the same form, 
The characters of both the upper and lower incisors distingnish 
Phaseolonus from Phescolomys, ; 


TOwen, Phil, Trans, IST, p. 257, 


PHALANGERID A. 147 


Manly PUALANGERID.E, 


Dentition oxtromely variable, owing to the presence of minute 
rudimental teoth not constant in the same species, or even in the 
two sides of the jaws of the same individual; exclusive, however, of 


Torsipes, the formuky ¢ a ¢ io p | me a represents fairly the 
general condition of the functional teeth. First incisors long and 
stout; the lower pair very huge and pointed, but without the scissor- 
like action fornd ino tho existing Mueropadide ¢ second and third 
lower incisors minute and probably fiuctionless, Fourth premolar 
generally secant; milk-moku generally minute and deciduons at an 
oarly period. Molars cither with sharp cutting-crests or bluntly 
tuboreulate ; fourth sometimes absent, Mandible without pit, and 
at most a very minute perforation in the masseteric fossa, Limbs 
subequal, Fore feet with tive distinct, subequal toes, furnished with 
claws. Hind feet short and broad, with tive well-developed toes ; the 
hallux large, nailless and opposable ; the second and third slender, 
and united by a common integument as far as the claws. Tail 
generally long, and frequently more or less prehensile, Stomach 
simple. Cree present (except ino Zursipes), and usually large. 
Pouch complete, Animals of small or moderate size and arboreal 
habits, usually feoding on a vegetable or mixed diet, inhabiting 
Australia and the Papuan Iskunds. 

The homologies of the lower functionless teeth between the first 
incisor and fourth premolar are very ditticult to determine, but 
it is probablo that one represents a canine only when the largest 
known number is present; this tooth, according to Mr. Thomas, 
boing the tirst to disappear, 

Phalangers are small woolly-couted animals, with long, power- 
ful, and often prehensile tails, large claws, and, as in the American 
oposstms, with opposable natiless great toes. Their expression 
seoms in the day to be dull and. sleepy, but by night they 
appear to decidedly greater advantage. They live mostly upon 
fruit, leaves, and blossoms, although some few feed habitually upon 
inseets, and all relish, when in confinement, an occasional bird 
or other small animal Several of the Phalangers possess tying 
membranes stretched between their fore and hind limbs (Hig. £8), 
by the help of which they cau make long and. sustained leaps 
through the air, like the Flying Squirrels, but it is interesting to 
notice that the possession of these flying membranes does not seem 
to bo any indication of special attinity, the characters of the skull 
and teeth sharply dividing the tlying forms, and uniting them with 
other species of the non-tlying groups. Their skulls (Pig. £7) 
areas atule broad aud: tlattened, with the posterior part swollen 


148 MARSUPIALIA 


out laterally, owing to the numerous air-cells situated in the 
substance of the squamosal. 

The Phalangers are interesting from an historical point of 
view, since the Gray Cuscus (Phalanger orientalis) was the first of 
the Marsupials of the eastern hemisphere brought to the notice of 
Europeans, having been described in a work published at Leyden 
in 1611, from an account of a specimen seen at Amboyna during 
the third expedition of Admiral Van der Hagen. 

The present family corresponds to the Dasyuride among the 


Fic. 46.—Tarsipes rostratus. From Gould. 


Polyprotodonts as presenting, on the whole, the most generalised 
types of the suborder. The existing forms may be divided into 
three subfamilies. 

Subfamily Tarsipedinze.—Cheek-teeth almost rudimentary and 
variable in number. Tongue long, slender, pointed, and very ex- 
tensile. Tail long. Ceacum absent. 

Tarsipes.\—So named from some supposed resemblance of its 
foot to that of the Lemurine genus Tarsius; but it must be remarked 
that it has none of the peculiar elongation of the caleaneum and 
navicular so characteristic of that genus, Head with elongated 


* Gervais and Verraux, Proc. Zool. Soc. 1842, p.1. 


PHALANGERIDE 149 


and slender muzzle. Mouth-opening small. The two lower 
incisors are long, very slender, sharp-pointed, and horizontally 
placed. All the other teeth are simple, conical, minute, and placed 
at considerable and irregular intervals apart in the jaws, the number 
appearing to vary in different individuals and even on different 
sides of the same individual. The formula in a specimen in the 
Museum of the Royal College of Surgeons is 7 2, ¢ 4, p and m 3 on 
one side, and # on the other; total 20. Rami of the mandible 
extremely slender, nearly straight, and without coronoid process or 
inflected angle. Fore feet with five well-developed toes, furnished 
with small, flat, scale-like nails, not reaching to the extremity of 
the digits. Hind feet rather long and slender compared with those 
of the Phalangerine, having a well-developed opposable and nailless 
hallux ; second and third digits syndactylous, with sharp compressed 
curved claws; the fourth and fifth free, and with small flat nails. 
Ears of moderate size and rounded. Tail longer than the body and 
head, scantily clothed with short hairs, prehensile. Vertebree: C 7, 
D13,L 5,8 3, C 24. 

Of this singular genus but one species, 7’. rostratus (Fig. 46), is 
known, about the size of a common Mouse. It inhabits Western 
Australia, lives in trees and bushes, uses its tail in climbing, and 
feeds on honey, which it procures by inserting its long tongue into 
the blossoms of Melaleuce, etc. One kept in confinement by Mr. 
Gould was also observed to eat flies. 

Subfamily Phalangerinz.— Teeth normal. One or more 
rudimentary teeth between the upper canine and fourth premolar, 
and between the first lower incisor and fourth premolar. Tongue 
of ordinary structure. No cheek-pouches. Stomach and ascending 
colon simple. Cezecum long, simple. Tail well-developed, generally 
prehensile. 

A numerous group of animals, varying from the size of a mouse 
to that of a large cat, arboreal in their habits, and abundantly 
distributed throughout the Australian region. The members of 
this group are the typical representatives of the family, and are 
commonly known to the colonists as Opossums. 

Phalanger.—The typical genus Phalanger (Cuscus) presents the 
following characters. No flying membrane; size large or medium, 
and build stout and clumsy; fur thick and woolly. Ears short 
or medium, hairy externally, and in some cases also internally. 
Toes of fore feet subequal, their relative lengths in the order 4, 3, 
5, 2,1. Claws long, stout, and curved. Soles of feet naked and 
striated, with large ill-defined pads. Tail stout and markedly 
prehensile, with the proximal half furred like the body, and the 
terminal portion entirely naked. Four mammez. Skull (Fig. 47) 

1 Storr, Prodromus Meth. Mam. p. 38 (1780). Syn. Phalangista, Geoffroy, 
Bull. Soc. Philom. vol. i. p. 106 (1796). 


150 MARS UPIALIA 


stout and strong, with large vacuities in the hinder half of the 
palate, and the auditory bulle thick and inflated. Dentition usually 
7 3,¢4, 94, m4. First upper incisor with nearly circular section, 
or only slightly flat- 
tened in front; can- 
ine more or less 
closely approximated 
to third incisor 
(which is very small), 
and situated partly 
in front of the suture 
between the pre- 
maxilla and maxilla. 
Fourth premolar 
large, secant, and 
placed obliquely to 
line of molars. 


7 Molars four-cusped, 
Fie 47.—Left lateral view of skull of Gray Cuscus (Phal- with the inner cusps 
anger orientalis). After Peters. 


of the upper ones 
crescentoid, and imperfect transverse ridges connecting each pair 
of cusps. 

The Cuseuses are curious sleepy-looking animals, inhabiting the 
various islands of the East Indian Archipelago as far west as Celebes, 
and being the only Marsupials found west of New Guinea. As 
already noted, it was a member of this genus, the Gray Cuscus 
(P. orientalis), a native of Amboyna, Timor, and the neighbouring 
islands, which was the first Australasian Marsupial known to European 
naturalists. There are altogether five species known, all of about 
the size of a large cat; their habits resemble those of other Phalan- 
gers, except that they are said to be somewhat more carnivorous. 

Trichoswrus..The members of the genus Jrichosurus are of 
relatively large size, and are distinguished from Phalanger by the 
following characters. Ears more or less hairy behind. Relative 
lengths of toes of fore feet in the order 4, 3, 2, 5, 1. Hair on the 
soles of the hind feet beneath the heel, but not elsewhere. Tail 
thick, not tapering, covered with bushy hair up to the extreme tip, 
which is naked, but with a naked strip on the inferior surface in 
the distal third or half. A gland on the chest. Dentition usually 
a 3,¢4,p2,m 4. Upper incisors of nearly uniform length, the 
first much flattened in front. Canine situated some distance behind 
the third upper incisor, which it scarcely exceeds in size. Last 
premolar and molars very similar to those of Phalunger. 

The true Phalangers comprise two species, of which the best 
known is the Vulpine Phalanger (7. vulpecula), so common in 


1 Lesson, Dict. Class, v’ Hist. Nut. vol. xiii, p. 833 (1828). 


PHALANGERIDE I51 


zoological gardens, where, however, it is seldom seen, owing to 
its nocturnal habits. It is of about the size and general build of 
a small fox, whence its name. In the typical variety the colour 
is gray, with a yellowish white belly, white ears, and a black tail. 
This variety is a native of the greater part of the continent of 
Australia, but is replaced in Tasmania by the closely allied Brown 
Phalanger (var. fuliginosa). Its habits are very similar to those of 
the Yellow-bellied Flying-Phalanger (Petawrus australis) described 
below, except that it is unable to take the flying leaps of that animal. 
Like all the other phalangers, its flesh is freely eaten both by the 
natives and the lower class of settlers. 

Pseudochirus1—The genus Pseudochirus agrees with the pre- 
ceding in the absence of a flying membrane, and presents the 
following leading characters. Size large or medium. Fur com- 
paratively short and woolly. Ears medium or short, hairy 
behind, although seldom closely furred over all this aspect. 
Claws medium. Fore toes subequal, the first two distinctly 
opposable to the other three. Soles of feet naked, with large, 
striated, round pads, and hair beneath the heels. Tail tapering, 
markedly prehensile, with its distal third and the whole of the 
under surface short-haired; tip naked underneath for a short 
distance. Four mamme. No gland on chest. Skull with larger 
nasals than in the preceding genera; the posterior part of the 
palate in most cases fully ossified, and the auditory bulle generally 
(2—3) (0O—1) 3 4 

eee Ne a Vag 
Upper teeth nearly uniform in length, but the first incisor distinctly 
longer than second. Upper premolars variable. Molars with both 
inner and outer cusps distinctly crescentoid, and recalling those 
of the Selenodont Artiodactyle Ungulates. 

Range.—Tasmania, Australia, and New Guinea. 

There are about ten species of this genus known, of which the 
commonest is Cook’s Ring-tailed Phalanger (Pseudochirus peregrinus), 
an animal discovered by Captain Cook during his first voyage, at 
Endeavour river, North Queensland. 

The complex and sub-selenodont character of the molars of this 
and the following genus readily distinguish them from the more 
typical Phalangers, and show an approximation to the type of 
dentition prevailing in Phascolarctus ; according, however, to Mr. 
O. Thomas, a tendency towards the same structure is observable 
in unworn molars of young Cuscuses. The genus may be divided 
into three groups, of which the first, as typified by the common P. 
peregrinus, is restricted to Australia and Tasmania, while the third, 
as represented by P. canescens, is only found in New Guinea. P. 
albertisi may be taken as the type of the second group, which is 


somewhat inflated. Dentition (at most) i 


1 Ogilby, Proc. Zool. Soc. 1836, p. 26. 


152 JLTARSUPIALIA 


represented by that species in New Guinea, and by P. archeri in 
Queensland. With the exception of P. peregrinus, the species have 
a more or less restricted range. Remains of Pseudochirus, probably 
referable to existing species, are found in the cave-deposits of New 
South Wales. 

Petauroides.A— With the genus Petauroides, containing only the 
single species P. vulans, we come to the first of the Flying-Phalangers, 
characterised by the possession of a flying membrane along the flanks. 
The characters of this genus are as follows. Size large. Fur very 
long and silky: Ears large and oval, thickly furred on the back, 
but naked internally. Flying-membrane reaching from wrist to 
ankle, but very narrow along the sides of the forearm and lower 
leg. Fore toes subequal, their relative lengths in the order 4, 3, 5, 
2,1. Claws long, curved, and sharp. Tail long, cylindrical, and 
bushy, except near its tip, where it is naked and prehensile. Skull 
short and broad, with the nasals short, and not extending nearly as 
far forwards as the premaxille. Large vacuities in hinder part of 
palate. Auditory bulle inflated and smooth. Dentition usually 
73, ¢%4,p%, m4. General characters of teeth very similar to those 
of Pseudochirus, but the first upper incisor scarcely longer than the 
second. 

The single species is found in Australia, from Queensland to 
Victoria, and is commonly known as the Taguan Flying-Phalanger. 
The structure of the skull and teeth indicates close affinity with 
Pseudochirus, although the external form is widely different in the 
two genera. This Phalanger seems, indeed, to be, so to speak, a 
very specialised Pseudochirus, in which the teeth have become 
somewhat further diminished and the flying membrane has been 
developed. 

Dactylopsila.2—The genus Dactylopsila is one of the forms with- 
out any trace of a flying membrane, its characters being as follows. 
Size medium. Body striped black and white. Ears oval, nearly 
naked at the ends. Fore toes of very unequal length, the fourth 
being enormously elongated; fourth and fifth toes of pes also 
markedly elongated. Claws long, moderately curved. Tail long, 
cylindrical, and evenly bushy, with the extremity more or less 
naked below. Skull narrow, but with the zygomatic arches greatly 
expanded ; palate fully ossified. Dentition: i 3, ¢ wpa, m 4. 
Upper incisors very large, the third being directed horizontally 
forwards; canine small and approximated to the third incisor, which 
it resembles. The fourth premolar of moderate size, with its longer 
axis placed obliquely. First lower incisor longer than in any other 
genus. Molars oblong, with four cusps. 

The typical D. trivirgatu, or Striped Phalanger, inhabits the 

1 Thomas, Cat. Marsupials Brit. Aus. p. 163 (1888). 
* Gray, Proc. Zool. Soc. 1858, p. 109. 


PHALANGERIDE 153 


Papuan and North Anstralian sub-region; a second species (D. 
palpator), characterised by the still greater clongation of the fourth 
finger, ocewrring in South New Guinea. These animals are said 
to be of insectivorous habits, the clongated fourth finger, as in the 
analogous instance of the Lemuroid genus Chiroiys, being appar- 
ently specially adapted for extractings inseets and larva from their 
hiding places. 

PetaurusA—Size medium or small. Fur very soft and. silky. 
A broad flying membrane extending from the outer side of the fifth 
digit of the manus to the ankle. Fore toes usually increasing 
regmlarly in length from the tirst to the fifth, but in some of the 
smaller species the fourth is the longest. Claws strong, sharp, and 
much curved. Tail long, evenly bushy to the extremity. Glands 
on the chest and between the ears. Skull short and wide, with 
the nasals expanded posteriorly, and usually two small palatal 
yacuities near the second molars, Auditory bulle inflated, and 
variable in size. Dentition: ¢8,¢), p.m +4. First upper incisors 
very large, and taller than canine. Molars with square crowns 
rounded at the angles, and four cusps, except in the last, which is 
triangular. 

This genus, which ranges from New Ireland to South Australia, 
but is not found in Tasmania, contains three species, the largest of 
which is the Yellow-bellied Flying-Phalanger (P. australis), whose 
habits are recorded by Mar. Gould as follows. “This animal is 
common inall the brushes of New South Wales, particularly those 
which streteh along the coast from Port Philip to Moreton Bay. 
In these vast forests trees of one kind or another are perpetually 
flowering, and thus offer a never-faling supply of the blossoms 
upon which it feeds; the flowers of the various kinds of gums, 
some of which are of great magnitude, are the principal favourites. 
Like the rest of the genus, it is nocturnal in its habits, dwelling in 
holes and in the spouts of the larger branches during the day, and 
displaying the greatest activity at might while running ever the 
small leafy branches, frequently even to their very extremities, in 
seaveh of insects and the honey of the newly-opened blossoms. — Its 
structure being il adapted for terrestrial habits, it seldom descends 
to the ground except for the purpose of passing to a tree too dis- 
tant to be reached by flight. When chased or foreed to fight it 
aseonds to the highest branch and performs the most enormous 
leaps, sweeping from tree to tree with wonderful address: a slight 
elevation gives its body an impetus which, with the expansion of 
its membrane, enables it to pass to a considerable distance, always 
ascending a little at the extremity of the leap: by this ascent the 
animal is prevented from receiving the shock which it would other- 
wise sustain,” 

V Shaw, Naturalists Miscellany, vol. ii, pl. lx. (1791. 


154 MARSUPIALIA 


A second species, P. sciurews, in some ways one of the most 
beautiful of all mammals, has been chosen for the accompanying 
woodcut. : 

Gymnobelideus.'—Like Petuurus in every respect, but. without 
any trace of a flying membrane, and with the fifth digit of the 
manus slightly shorter than the third. This genus is represented 
only by G. leadbeatert of Victoria, and according to Mr. Thomas, 
may be regarded as the primitive form from which the specialised 
Petaurus has been developed. 


\ 


Fic, 48.—Squirrel Flying-Phalanger (Petaurvs sciureus). 


Dromiciu.2—Size small, and general appearance dormouse- 
like. Ears large and thin, almost naked, and without internal 
or basal tufts. No flying membrane. Digits of normal propor- 
tions, the relative lengths of those of the manus in the order 
3, 4, 2, 5, 1; fore claws rudimentary, hind ones long and sharp. 
Tail mouse-like, cylindrical, furry at base, the remainder sealy, 
with fine hairs, except at the tip, which is naked and prehensile. 


1 Moy, Ann. Mag. NV. H. (3) xx. p. 287 (1867). 
” Gray, in Grey’s Australia, appendix, vol. ii. p. 407 (1841). 


PHALANGERIDE 155 


Skull short and broad, with the hinder part of the palate in- 
complete, and the auditory bulle large, much inflated, and trans- 
parent. Dentition: 2 :, c a p . m = First upper incisor spat- 
ulate, and much longer than either of the others. Canine large, 
placed at some distance behind the third incisor. Molars (except the 
last) with evenly rounded crowns, carrying four small smooth cusps. 

This genus, which occurs in New Guinea, Western Australia, and 
Tasmania, is represented by four species. It seems to be inter- 
mediate between Petawrus and Acrobates, and it has apparently had 
to yield place to those more highly organised types in regions where 
they have come in contact with one another. 

Distechurus.1—Size small. Ears rather short, thinly covered 
with hair, but with small tufts at the base. No flying membrane. 
Digits of normal proportions, without expanded terminal pads. 
Claws curved and sharp. Tail, skull, and dentition as in Acrobutes, 
with the exception that the fourth premolar is small in the upper, 
and absent in the lower jaw. 

The one species of Feather-tailed Phalanger (D. pennatus) is 
found in New Guinea. 

Acrobates.2—Size very small. Ears moderate, thinly covered 
with hair, but with small tufts round the base and on the internal 
prominences. A narrow flying membrane, fringed with long hairs, 
running from the elbow to the flank, and from the latter to the 
knee. Four mamme. Digits furnished with expanded and striated 
terminal pads, the relative length of those of the manus being in the 
order 4, 3, 5, 2, 1. Claws sharp, although somewhat concealed by 
the terminal pads. Tail short-haired above and below, with a broad 
fringe on either side. Skull short, wide, and depressed. Posterior 
portion of palate very imperfectly ossified ; anterior palatal vacuities 
almost confined to the maxille. Auditory bulle low, rounded, and 
but slightly prominent. Dentition: 7 3,¢3,p 4, m2. Teeth sharp, 
and of an insectivorous type. Upper canine long, and approximated 
to third incisor. The three upper premolars large, functional, and 
taller than the molars. Molars small and rounded, with smooth 
unridged cusps. 

There is only one species in this genus, the beautiful little 
Pigmy Flying-Phalanger (4. pygmeus), not so big as a Mouse, which 
is found in Queensland, New South Wales, and Victoria, and feeds 
on the honey it abstracts from flowers, and on insects. Its agility 
and powers of leaping are exceedingly great, and it is said by 
Mr. Gould to make a most charming little pet. 

Subfamily Phasecolaretine.—Teeth large, normal; no rudi- 
mentary premolars before the last upper premolar, or any teeth 


1 Peters, dnn. Mus. Genov. vol. vi. p. 303 (1874). 
2 Desmarest, Nouv, Dict. d’ Hist. Nat. sér. 2, vol. xxv. p. 405 (1817). 


156 JLARSUPIALIA 


between the first lower incisor and fourth premolar. Tongue 
of ordinary structure. Distinct cheek-pouches. Stomach with a 
special gland near the cardiac orifice. Caecum very long, and (with 
the upper portion of the colon) dilated and provided with numerous 
longitudinal folds of mucous membrane. In many anatomical 
characters, especially the possession of a special gastric gland, this 
group resembles the Phascolomyide.1 

Phascolarctus.2—Dentition : 12, ¢4, p+, m4; total 30. Upper 
incisors crowded together, cylindroidal, the first much larger than 
the others, with a bevelled eutting edge (Fig. 36). Canine very 
small ; a considerable interval between it and the premolar, which 
is as long from before backwards but not so broad as the true 
molars, and has a cutting edge, with a smaller parallel inner ridge. 
The molars slightly diminishing in size from the first to the fourth, 
with square crowns, each bearing four pyramidal cusps, with curved 
ridges radiating from them, and having a structure very similar to 
these of Pseudochirus. The lower incisors are semiproclivous, com- 
pressed and tapering, bevelled at the ends. Premolars and molars 
in continuous series, as in the upper jaw. Milk-tooth very minute, 
and almost functionless. Fore feet with the two inner toes slightly 
separated from and opposable to the remaining three, all with strong, 
curved, and much compressed 
claws. Hind foot (Fig. 49) with 
the hallux placed very far back, 
large and broad, the second and 
third (united) toes considerably 
smaller than the other two ; the 
fourth the largest. No external 
tail. Fur dense and woolly. 
Ears of moderate size, thickly 
clothed with long hairs. Verte- 
bre: C7,D11,L8,82, Cs. 
Ribs eleven pairs, a rare excep- 
tion to the usual number (13) 
in the Marsupialia. 

There is but one species, 
the Koala or Native Bear of 
£Fic. 49,—Skeleton of right hind foot of Koala the Australian colonists (P. cin- 


(Phascolarctus cinereus), showing the stout op- @réus), an animal of compar- 
posable hallux, followed by two slender toes, atively large size and heavy 


which in the living animal are enclosed as far * . ie 4 

as the nails in a common integument. build (Fig. 50), found in the 
; ; ; south-eastern parts of the Aus- 

tralian continent. It is about two feet in length, and of an ash- 

gray colour, an excellent climber, and residing generally in lofty 


1 cy, W. A, Forbes, ‘ Anatomy of the Koala,” Proc. Zool. Soc. 1881, p. 180 
? Blainville, Bull. Soc. Philom, 1816, p. 116. 


PHALANGERIDE 157 


Eucalyptus trees, on the buds and tender shoots of which it feeds, 
though occasionally descending to the ground in the night. 


EXTINCT PHALANGEROIDS. 


Numerous imperfect remains recently described by De Vis are 
regarded as indicating large extinct types of Phalangeride, but 
further evidence is required before all these determinations can be 
definitely accepted. Thus part of an upper jaw is provisionally 
referred to a large species of Pseudochirus, while part of a scapula 
is made the type of a genus Archizonurus which appears to be 


Fic, 50.—The Koala (Phascolarctus cinereus). From Selater, Proc. Zool. Soc. 1880, p. 355. 


allied to the former. Another fragmentary scapula is considered to 
indicate a large Phalanger. Finally, part of a fibula described under 
the name of Koalemus is regarded as affording evidence of the 
former existence of a large ancestral form allied to the Koala, and 
it is suggested that an upper jaw with teeth may belong to the 
same or an allied type. 

Thylacoleo.1—Dentition of adult: 13, ¢3, p%, m4; total 28, 
First upper incisor much larger than the others; canine and first 
two premolars rudimentary. In the lower jaw the two small 
anterior premolars are functionless, and often deciduous ; posterior 
premolars of both jaws formed on the same type as those of Potorous, 
but relatively much larger; true molars rudimentary, tubercular. 
One species, 7. carnifer. This animal presents a most anomalous 


1 Owen, in Gervais’s Zool. et Pal. frangaises, Ist ed. pt. i. p. 192 (1849-52). 


158 MARSUPIALIA 


condition of dentition, the functional teeth being reduced to one 
pair of large cutting incisors situated close to the median line, and 
one great, trenchant, compressed premolar, on each side above and 
below. It was first 
described as a car- 
nivorous Marsupial, 
and named, in ac- 
cordance with its 
presumed _ habits, 
“as one of the fel- 
lest and most de- 
structive of preda- 
tory beasts”; but, 
as its affinities are 
certainly with the 
Phalangeride and 
Macropodide, and 
its dentition com- 
pletely unlike that 
of any known pre- 
daceous animal, this 
view has been called 
——— in question. 
Fig. 51.—Front view of skull of Thylacoleo carnifee, restored. The dentition is 
3 natural size. From Quart. Journ. Geol. Soc. vol. xxiv. p. 312. nearer to that of the 
existing Phalangeride than to that of the Mfucropodide, and the 
genus may be provisionally regarded as the type of a distinct 
subfamily of the former. 


Family MAcRopopID&. 


ae 3 0—1 2 4 F 
Dentition 2 > ¢ ( 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 <round when the animal is at rest in the upright position. 

The stomach is of large size. and very complex, its walls being 
puckered up by longitudinal muscular bands into a great number ot 
sacculi, like those of the human colon. The alimentary canal Is 
long. and the cxcum well developed. All the species have a 
marsupium or pouch formed by a fold of the skin of the abdomen 
covering the mammary glands with their four nipples. In this 
pouch the young are placed as soon as they are born: there their 
growth and development proceeds; and to it they resort tempor- 
arily for the purpose of shelter, concealment, or transport, for some 
time after they are able to run and jump about the ground and 
feed upon the same herbage which forms the nourishment of the 
parent. During the early period of their sojown in the pouch, 


MACROPODIDE 161 


the blind, naked, helpless young creatures (which in the Great 
Kangaroo (Fig. 53) scarcely exceed an inch in length) are attached 
by their mouths to the nipples of the mother, and are fed by 
milk injected into their stomach by the contraction of the muscle 
covering the mammary gland. 

The Kangaroos are all vegetable feeders, browsing on grass and 
various kinds of herbage, the smaller species also eating roots. 
They are naturally timid, inoffensive creatures ; but the larger ones 
when hard pressed will turn and defend themselves, sometimes 
killing a dog by grasping it in their fore paws, and inflicting 
terrible wounds with the sharp claws of their powerful hind legs, 
sustaining themselves meanwhile upon the tail. A few aberrant 
forms are arboreal. The great majority are inhabitants of Australia 
and Tasmania, forming one of the most prominent and characteristic 
features of the fauna of these lands, and in the scenery of the 
country, as well as the economy of nature, performing the part of 
the deer and antelopes of other parts of the world, which are 
entirely wanting in Australia. Kangaroos were very important 
sources of food-supply to the natives, and are hunted by the colon- 
ists, both for sport and with a view to their destruction, on account 
of the damage they naturally do in consuming the grass, now 
required for feeding cattle and sheep. Notwithstanding this, they 
have in some districts increased in numbers, owing to the sup- 
pression of their former enemies, the aborigines and the Dingo or 
native dog. A few species are found in New Guinea and the 
adjacent islands, which belong, in the zoological sense, to the 
Australian region. 

Before noticing the various generic types of the IMucropodide, a 
few words are necessary in respect of the tooth-change, and we may 
here quote the observations of Mr. O. Thomas on this subject. 
“The full dentition of the members of this family consists, in the 
upper jaw, first of three incisors, then of a small canine (often, 
however, suppressed, as in Fig. 55), and then of six cheek-teeth, 
of which the second in the series is the only one which has a milk 
or deciduous predecessor, and is therefore the one to be regarded 
as the last premolar of the typical mammalian dentition, The 
special characteristics that render the development and succession of 
the teeth in the Macropodide, and especially in the genus Macropus, 
so puzzling to systematic zoologists, are: firstly, a general pro- 
gression forwards in the jaw of the whole tooth-row, comparable to 
that found elsewhere only in the Elephants and some Sirenians ; 
and, secondly, the fact that before the tooth-change the first tooth 
of the series (p 3) and the single milk-tooth (dm 4) placed next to 
it, both of which fall out at the change, are respectively so very 
similar in shape and size to the first and second teeth of the 
permanent series, viz. the permanent premolar (p 4) and the first 

11 


162 MARSUPIALIA 


molar (m 1), as to be most naturally mistaken for, or compared with, 
them in specific descriptions. . . . The necessary knowledge as to 
the stage of dentition in which any skull may he, can often be 
gained only by cutting open the bone either above and behind the 
first tooth of the series to see if the true permanent p 4 be still 
buried there (in which case, of course, that first tooth is only p 3), 
or behind the last visible molar to see if there be yet another tooth 
behind it, showing it to be m3 and not m4. The first plan is, 
as a rule, the better, since » 4 is generally by far the most 
important tooth for diagnostic purposes, and its characters have, 
therefore, in any case to be taken into account.” 

The Macropodide are divided into three well-marked sections : 
(1) the true Kangaroos (Macropodine) ; (2) a group consisting of 
smaller animals, commonly called Rat Kangaroos, or (improperly) 
“ Kangaroo Rats,” or sometimes Potoroos; and (3) the Hypsiprym- 
nodontince, now represented only by a single species. 

Subfamily Hypsiprymnodontinz.—Size very small. Claws 
small, feeble, and subequal. Hind feet with an opposable hallux. 
Tail naked and scaly. The fourth premolar twisted obliquely out- 
wards, as in Phalanger. Other teeth as in the Potorvine. 

This subfamily is now represented only by the genus Hyps?- 
prymnodon, which is a form of great interest, as showing a structure 
of foot connecting that of the Kangaroos with that of the Phalan- 
gers. The single known species, H. moschatus, was described by 
Ramsay from specimens discovered in north-east Australia. It 
was described almost simultaneously by Owen under the name of 
Pleopus nudicaudatus. From the resemblance in the structure of the 
foot and the obliquity of the premolars to the Phalangers Mr. 
Thomas has some hesitation as to which family should receive this 
genus, but the macropine characters of the mandible preponderate 
in favour of the Macropodide. 

Trichis2—A lower jaw of a much larger form from the Pleisto- 
cene deposits of Australia apparently indicates another member of 
this subfamily, having the outwardly directed and grooved pre- 
molar characteristic of Hypsiprymnodon. It differs, however, from 
that genus, and also from all other known Macropodide, in having 
a small tooth between the incisor and fourth premolar, which 
apparently represents a canine, or perhaps an anterior premolar. 
This form indicates, therefore, a closer connexion between the 
Phalangeride and Macropodide than any other. 

Subfamily Potoroinz.—The second section or subfamily, the 
Potoroine, have the first upper incisor narrow, curved, and much 
exceeding the others in length (Fig. 54). Upper canines always 
persistent, flattened, blunt, and slightly curved. Premolars of both 

1 Ramsay, Proc, Linn, Soc, N.S. Wales, vol. i. p. 33 (1876). 
* De Vis, Proc. Roy. Soc. Queensland, sev. 2, vol. iii, p. 8 (1888). 


MACROPODIDA 163 


jaws always having large, simple, compressed crowns, with a nearly 
straight or slightly concave free cutting edge, both outer and inner 
surfaces usually marked by a series of parallel, vertical grooves and 
ridges, these teeth being either set in the same line with the 
molars, or slightly bent outwards. Molars with quadrate crowns, 
having a blunt, conical cusp at each corner, the fourth notably 
smaller than the third, sometimes rudimentary, and appearing early. 
Fore feet narrow; three middle toes considerably exceeding the 
first and fifth in length; their claws long, compressed, and but 
slightly curved. Hind feet as in Macropus. Tail long and hairy, 
sometimes partially prehensile, being used for carrying bundles of 
grass with which these animals build their nests. 

The Potoroos or Rat Kangaroos are all small animals, none of 
them exceeding a common rabbit in size. They inhabit Australia 
and Tasmania, are nocturnal, and feed on the leaves of various 


Fia. 54.—Skull and Teeth of Rat Kangaroo (Bettongia lesuewiri). vc, Upper canine. 
The other letters as in Fig. 51. 


kinds of grasses and other plants, as well as roots and bulbs, which 
they dig up with their fore paws. Nine species are known, present- 
ing a considerable range of diversity in minor characters, and 
admitting of being grouped in four principal sections, which may 
be allowed the rank of genera. These are: 

Potorous.\—Head long and slender. Auditory bulls some- 
what inflated. Ridges on premolars few and_ perpendicular. 
Large palatine foramina. Tarsus short. Muffle naked. Three 
species, viz. P. triductylus, P. gilberti, and P. platyops ; the last two 
being confined to West Australia. 

Bettongia.2—Head comparatively short and broad. Ears short 
and rounded. Auditory bulle generally much inflated. Large 
palatine foramina. Tarsus long. Ridges on premolars numerous 


1 Desmarest, Nouv. Dict. d’Hist. Nat. ser. 1, vol. xxiv. Table Meth. ». 20 
(1804). Syn. Hypsiprymnus, Miger, Prodromus Syst. Mam. p. 79 (1811). 
* Gray, Charlesworth’s Mag. Nat. Hist. vol. i, p. 584 (1837). 


164 WARSUPIALIA 


and oblique. Tail more or less prehensile, thickly haired, and 
the hairs on the upper surface longer than those on the lower, and 
forming a crest. Muffle naked. Four species, viz. B. penicillata, 
B. cuniculus, B. gaimardi, B. lesueuiri. 

Caloprymnus.rMufile naked, as in Bettongia, but the edge of the 
hairy part less emarginate backwards in the middle line. Ears 
short, rounded, and hairy. Auditory bulle much inflated, and of 
large size. Nasals larger and wider behind than in the other 
genera. Very long anterior palatine foramina. Limbs as in 
Bettongia. Tail thin, cylindrical, evenly coated with short hair, 
without trace of a crest. Skull broad and flat, with a remarkably 
short and conical muzzle. The sole representative of this genus is 
C. campestris of South Australia, originally referred to Bettongia. 


sil, 


pm | H H ; 

m mn? ne? iit 

Fig. 55.—Skull and Teeth of the Red-necked Wallaby (Macropus rwficollis). i, i, i3, First, 
second, and third upper incisors ; pm, fourth or posterior premolar (the penultimate or third 


having been already shed); m1, m2, m3, m4, the four true molars. The last, not fully de- 
veloped, is nearly concealed by the ascending rainus of the jaw. 


Aipyprymnus.,—Head short and broad. Auditory bulle not 
inflated. No palatine foramina. Tarsus long. Mufile partially 
hairy. ‘Tail evenly hairy, not crested above. Molars oblong, less 
distinctly quadritubercular, and not decreasing so much in size pos- 
teriorly as in the other genera. Represented only by -E. rufescens. 

Remains of 4. rufescens occur in the Pleistocene cave-deposits 
of New South Wales. 

Subfamily Maeropodinsze.—This subfamily includes the largest 
forms. The cutting edges of the upper incisors are nearly level, or 
the first pair but slightly longer than the others (Fig. 55). The 
canines are rudimentary and often wanting. The premolars are 
usually not longer (from before backwards) than the true molars 


1 Thomas, Cat. Marsup. Brit. Mus. p. 114 (1888). 
* Garrod, Proc. Zool. Soc, 1875, p. 59. 


MACROPODIDA 165 


and less compressed than in the last subfamily; they are placed 
in precisely the same line with the molars. The crowns of the 
molars always have two prominent transverse ridges; and these 
teeth increase in size from before backwards, the fourth molar 
appearing very late. The fore limbs are small, with subequal toes 
armed with strong, moderately long, curved claws. Hind limbs 
very long and strongly made. Head small, with more or less 
elongated muzzle. Ears generally rather long and ovate. 

Upwards of forty-four existing species of this group have been 
described, and many attempts have been made to subdivide them into 
smaller groups or genera for the convenience of arrangement and 
description, but these have generally been based upon such trivial 
characters that it is preferable to speak of many of them as sections 
of the genus Macropus, reserving generic rank only to forms some- 
what aberrant in structure. According to this arrangement the 
genera will be as follows: 

Lagostrophus..—Represented only by the Banded Wallaby 
(L. fasciatus) of Western Australia, which presents the following 
distinctive features. Size small. Mufile naked. Hind feet covered 
with long bristly hairs, concealing the claws. Lower part of back 
marked by dark cross-bands. Skull with a narrow pointed muzzle 
and inflated auditory bulle ; symphysis of mandible firmly united. 
No canine. Upper incisive series meeting at a sharp angle, and 
diverging but slightly behind. First incisor smaller in section than 
either of the others and scarcely longer, bluntly pointed ; second 
with a flattened oral surface ; third smaller, similarly flattened, but 
with a groove on oral surface forming a notch at its postero- 
external angle. Fourth premolar short, with a distinct inner ledge. 
Molars as in Macropus. 

Dendrolagus.2—General proportions of limbs and body normal 
and unlike those of other members of the family. Muffle broad and 
only partly naked. Fur on nape, and sometimes on back, directed 
forwards. Fore limbs nearly as large as the hind; hind feet with 
the syndactylous second and third digits relatively large; claws of 
fourth and fifth hind digits curved like those of the manus. Tail 
very long, and thickly furred. Skull stout, with a short and wide 
muzzle; the posterior part of the palate fully ossified, and the 
auditory bulle not inflated. A small canine. Fourth premolar 
large, but much shorter antero-posteriorly than in the next genus ; 
molars as in the latter. 

This genus includes four species of Tree-Kangaroos, three of 
which occur in New Guinea, while D. dumholtz is found in North 
Queensland. They differ greatly from all the other forms in being 
chiefly arboreal in their habits, climbing with facility among the 

1 Thomas, Proc. Zool. Soc. 1886, p. 544. 
° Schlegel and Miiller, Verh. Nat. Ges. Nederland, p. 138 (1839-44). 


_ 166 MARSUPIALIA 


branches of large trees, and feeding on the bark, leaves, and fruit. 
They are confined to the tropical forests of the regions mentioned ; 
and it would appear that we must regard their resemblance in the 
proportions of the limbs and habits to the Phalangers as having 
been independently acquired. 

Dorcopsis..—Hind limbs relatively less large than in Muacropus. 
Muffle large, broad, and naked. Ears small. Fur on nape directed 
wholly or partially forwards. Hind claws not concealed by hair. 
Tail with a nearly naked tip. Skull long and narrow, with the 
auditory bulle not inflated. A well-developed canine. First upper 
incisor somewhat short; second and third nearly equal, notched 
externally. Fourth premolar greatly elongated antero-posteriorly, 
its length generally exceeding the united lengths of the first and 
second molars ; a distinct inner ledge, and vertical grooves on both 
sides. Molars low and rounded, with the median longitudinal 
bridge between the ridges almost or quite aborted, and the talon in 
front of the first transverse ridge very narrow, and not extending 
to the inner side. The two series of cheek-teeth parallel, or nearly 
so, instead of converging at the extremities. 

Three species of this genus are known, all of which are from 
New Guinea ; the type being D. muelleri. In the characters of the 
dentition, the forward inclination of the fur on the nape, and other 
points, this genus is allied to Dendrolagus ; but Dorcopsis macleayt 
connects the other species with Afucropus. 

Lagorchestes.2-—Muffle entirely or partially covered with hair. 
Fourth hind digit with a long claw, not concealed by hair. Tail 
rather short, evenly furred, without a spur. Skull with short 
muzzle and diastema, and inflated auditory bulla. Canine present, 
sometimes very small. Fourth premolar large, not constricted in 
the middle, with a continuous inner ledge. 

This genus includes the Hare-Kangaroos, a group of small 
hare-like animals, great leapers and swift runners, which mostly 
affect the open grassy ridges, particularly those of a stony character, 
sleeping in forms or seats like the common hare. Their limbs are 
comparatively small, their claws sharp and slender, and their muffle 
is clothed with velvet-like hairs. Three species—J/. leporoides, ML. 
hirsutus, M. conspicillatus. 

The range extends over the whole of Australia, but does not 
embrace Tasmania. 

Onychogale.2— Muffle hairy. Fourth hind claw long, narrow, 
compressed, and sharp. Tail long and tapering, covered with short 
hair, and furnished at the tip with a horny spur. Skull nearly as in 
Macropus, with the auditory bulle more or less inflated. Canine 


? Schlegel and Miiller, Verh. Nat. Ges. Nederland, p. 180 (1839-44), 
? Gould, Monograph of Macropodide, pl. xiii. (1841), 
* Gray, in Grey's Australia, vol. ii. appendix, p. 402 (1841). 


MACROPODIDA 167 


small or wanting. Upper incisors small, decreasing in size from first 
to third. Fourth premolar small, hour-glass shaped, and without 
inner ledge. Molars as in Macropus. 

This genus contains three species, having the same distribution 
as Lagorchestes. Mr. O. Thomas observes: “The spur-tailed Wallabies 
form a natural little group, distinguished both by the shape of the 
incisors and the peculiar horny excrescence at the tip of the tail. 
The latter character is altogether unique among Marsupials, and is 
only found among other mammals in the Lion, which occasionally 
has a somewhat similar horny spur at the end of its tail. In the 
case of the Wallabies it is difficult to conceive what can be the 
use of this spur; and observations on the living animal are much 
needed with regard to this interesting point.” 

Petrogale._— Muffle naked. Fur of nape directed backwards. 
Claw of fourth hind digit very short. Tail long, cylindrical, thinner 
than in Macropus, and thickly haired and pencilled at the extremity. 
Skull as in the smaller species of Macropus, with large posterior 
palatal vacuities, and the bulle sometimes inflated. No canine. 
Upper incisors small, the third resembling that of Macropus. Fourth 
premolar large and stout, as in some of the Wallabies, with a con- 
tinuous inner ledge, and two or three indistinct vertical ridges 
externally. Molars as in the Wallabies. 

This genus is represented by six species, of which P. penicillata 
is a well-known example, ranging over the whole of the mainland of 
Australia. The Rock-Wallabies, as its members may be called, are 
very closely allied to some of the true Wallabies ; and some hesitation 
may be expressed as to the advisability of accepting their generic 
separation from Jfacropus. They inhabit rocky regions, making 
their retreats in caverns and crevices, leaping with surprising agility 
from one narrow ledge to another, and browsing upon the scanty 
herbage that the neighbourhood of such situations affords. The 
species are P. xanthopus, P. penicillata, P. lateralis, P. concinna, P. 
brachyotis, P. inornata. 

Remains of P. penicillata are found in a fossil state in the 
Pleistocene cave-deposits of New South Wales. 

Macropus.2—Muffie generally completely naked. Lars large. 
Fur on nape (with an occasional exception in two species) directed 
backwards. Claw of fourth hind digit very long. Tail thick, 
tapering, and evenly furred. Four mamme. Skull (Fig. 55) long, 
smooth, and rounded ; the nasals expanded behind ; generally large 
palatal vacuities; and the auditory bulle not inflated. Canine 
minute, and shed at an early period. JIncisor series forming an 
open curve ; the first the tallest, and the third nearly always the 
longest antero-posteriorly, and generally with an infolding of enamel 


1 Gray, Charlesworth’s Mag. Nat. Hist. vol. i. p. 583 (1837). 
2 Shaw, Naturalist’s Miscellany, vol. i. pl. xxxiii. (1790). 


168 JARSUPIALIA 


near its postero-external angle. Fourth upper premolar with a 
secant edge, and an inner basal ledge or tubercle ; corresponding 
lower tooth secant ; both may be longer or shorter than first molar. 
Molars (except very occasionally) with a distinct longitudinal bridge 
connecting transverse ridges. Lower incisors long and scalpriform, 
with inner secant edges opposable, owing to the loose articulation of 
the mandibular symphysis. 

This genus includes the true Kangaroos and Wallabies, the size 
of the individual existing species varying from that of a Rabbit 
to that of a Man. There are no less than twenty-three existing 
species, which may be divided into three groups, as well as many 
extinct ones. The genus is found in Australia and New Guinea, 
as well as in the eastern half of the Austro-Malayan transitional 
region. 

The first group, or true Kangaroos, comprises the largest 
existing forms, which are generally of a uniform and sombre colour. 

The skull is of a large and massive type, with the palate more 
or less well ossified posteriorly, while the molars frequently have 
a median longitudinal bridge connecting the first transverse ridge 
with the anterior talon, and no antero-external bridge between the 
same ridge and talon. The history of the discovery of the typical 
representative of this group, as being of considerable interest, may 
be given at some length. When Captain Cook, during his first 
memorable voyage of discovery, was detained for the purpose of 
refitting his ship at Endeavour river on the north-east coast of 
Australia, a strange-looking animal, entirely unknown to them, was 
frequently seen by the ship’s company; and it is recorded in the 
annals of the voyage that, on the 14th of July 1770, “Mr. Gore, 
who went out this day with his gun, had the good fortune to kill 
one of the animals which had been so much the subject of our 
speculation, . . . and which is called by the natives kanguroo,” a 
name which, though it does not appear to be now known to any of 
the aboriginal tribes of the country, has been adopted for ‘this 
animal in all European languages, with only slight modifications of 
spelling. With the exception of a passing glimpse in the beginning 
of the sume century by the Dutch traveller Bruyn of some living 
examples of an allied species, this was the first introduction to the 
civilised world of any member of a group of animals now so 
familiar. The affinities of the species, skins of which were brought. 
home by Captain Cook and subsequent voyagers, were recognised 
by Schreber as nearer to the American opossums (then the only 
known Marsupials) than to any other mammals with which zoologists 
were acquainted, and consequently it was placed by him, in. his 
great work on the Mammalia, then in the course of publication, in the 
genus Didelphys, with gigantes for a specific designation,x—the latter 
having heen bestowed upon it by Zimmermann under the impression 


MACROPODIDZ 169 


that it was a huge species of jerboa. Soon afterwards (1791) Dr. 
Shaw very properly formed a new genus for its reception, which 
he named Mucropus, in allusion to the peculiar length of its hind 
foot. By the name thus formed, Macropus giganteus, this kind of 
Kangaroo has ever since been known in zoological literature. It is 
the common Gray Kangaroo, called “boomer,” “forrester,” or “old 
man” by the colonists, and frequents the open grassy plains of the 
greater part of eastern Australia and Tasmania; a figure being 
given in the woodcut on p. 160. The muffle is partly covered 
with hair, and the fourth premolar very short. Several varieties 
are known. 

A sub-group, distinguished from the above by the naked 
muffle, includes some very large and handsome species, which prin- 
cipally dwell in rocky mountain ranges, as M/. rufus, the great Red 
Kangaroo, M. antilopinus, and Jf. robustus. The fourth premolar is 
of large or medium size in these forms. Remains of J/. giganteus 
occur fossil in the Pleistocene of Australia, where we also find the 
allied extinct Jf. titan, which attains somewhat larger dimensions. 
Af. robustus also dates from the same geological epoch, where it was 
accompanied by two allied types known as JV. altus and JJ. cooperi. 

The second group includes the larger Wallabies, which are 
smaller than the true Kangaroos, with a brighter and more 
variegated coloration. The palate is generally more incomplete 
than in the typical group; and in the molars the anterior talon is 
connected with the first transverse ridge by an external instead of 
a median longitudinal bridge. The members of this group are 
frequenters of forests and dense impenetrable brushes and scrubs, 
and hence are often called Brush Kangaroos, though a native name, 
“Wallaby,” is now generally applied to them. There are several 
species, of which MV. ruficollis, MM. ualabatus, M. perry, and AL. agilis 
are the best known. 

M. ualabatus and AM. parryi ave found fossil in the Pleistocene 
deposits of Australia. In those beds we also meet with remains of 
several very large extinct species, which appear to be allied to those 
Wallabies in which the fourth premolar is large and elongated, all 
of them agreeing with the Wallabies in the absence of the median 
bridge between the first ridge and talon of the molars. These fossil 
forms comprise M. brehus, in which the skull was probably about 
one foot in length, and JZ. rechus, and Af. anak, which were of some- 
what inferior dimensions. In the last-named species the length of 
the fourth upper premolar is equal to that of the first and half of 
the second molar.+ 

The third and last group of the genus includes the small 


1 For the characters of these species and the undermentioned distinct genera, 
see Owen’s Extinct Mamials of Australia (1877), and Lydekker’s Catalogue of 
Fossil Mammalia in the British Muscum, pt. v. (1887). 


170 MWARSUPIALIA 


Wallabies, which are small and lightly-built animals, in some 
instances not larger than a Rabbit. Their muffles are always naked, 
and in the skull the anterior palatine foramina are small and the 
posterior vacuities very large, while the posterior expansion of the 
nasals is very marked. The third upper incisor is smaller than in 
the last group. This group extends farther into the tropics than 
either of the others, being found in the New Britain and Aru 
islands, as well as in New Guinea. J. brachywrus is remarkable for 
its comparatively short and slender tail and small ears. The earliest 
known species of Kangaroo, referred to before, I. brunt, belongs to 
this section. Several examples were seen by Bruyn in 1711 living 
in captivity in the garden of the Dutch governor of Batavia, and 
described and figured in the account of his travels (Reizen over 
Moskovie, ete.) under the name of “Filander.” It was quite lost 
sight of, and its name even transferred by S. Miiller to another 
species (Dorcopsis muelleri), until rediscovered in 1865 by Rosenberg, 
who sent a series of specimens to the Leyden Museum from the 
islands of Aru and Great Key, thus determining its true habitat. 
M. thetidis is a well-known Australian representative of this 
group. 

Extinct genera.—In addition to the fossil forms already mentioned 
which can be referred to existing genera, there are others from the 
Australian Pleistocene indicating extinct generic types of Macropod- 
ide, to which brief reference may now be made. The first of these 
is Sthenurus,) represented by a single large species (S. aélas), and 
characterised by the presence of a complete inner lobe to the fourth 
upper premolar, and of an outer one in the opposing lower tooth, 
so that these teeth present a flat and oval grinding surface when 
worn. The median longitudinal bridge connecting the transverse 
ridges of the molars is very imperfect; and in the upper molars 
there is no bridge between the first ridge and talon. In Procoptodon? 
the premolars resemble those of Sthenurus, but the molars are 
elongated, and usually have their enamel thrown into numerous 
vertical foldings. The most distinctive feature is, however, the 
complete ankylosis of the mandibular symphysis; the mandibular 
rami being deep, and the diastema in the dental series short. The 
lower incisors are nearly cylindrical, and the palate has large 
vacuities. Three species are known. The largest representation of 
the whole family is the type of the genus Pulorchestes® (P. azael), in 
which the length of the skull is estimated at sixteen inches. It is 
distinguished from Procoptodon by the longer mandibular symphysis 
and diastema, and the spatulate lower incisors. The true molars 
have no distinct anterior talon, and are not grooved, while the 
palate was fully ossified. 


1 Owen, Phil. Trans, 1874, p. 264. 
2 Owen, op. cit. p. 788. 5 Owen, op. cit. p. 797. 


EXTINCT FAMILIES 171 


EXxtTINcr FAMILIES. 


Here may be noticed two genera of extinct Marsupials, the remains 
of which have been found in the Pleistocene deposits of Australia, 
which agree with the Macropodide and the Phalangeride in having 
2. incisors, those of the lower jaw being very large and proclivous. 
As the whole of their structure, especially that of the hind feet, is 
not yet known, their precise affinities cannot be determined. 

Diprotodon.t—Dentition : 1 3, ¢ 2, p 4,m 4; total 28. The first 
upper incisor very large and scalpriform (Fig. 56). True molars 
with prominent transverse ridges, as in Macropus, but wanting 
the longitudinal connecting bridge. Anterior and posterior limbs 
less disproportionate than in the Kangaroos. Humerus elongated, 
and differing from that of nearly all Marsupials in the absence of an 


Fic. 56.—Left lateral aspect of the skull of Diprotodon australis ; from the Pleistocene of 
Australia, jp; natural size. i, Incisors; p, premolar; m, molars. (After Owen.) 


entepicondylar foramen. ‘The palate is fully ossified, and there is 
no pit or perforation in the masseteric fossa of the mandible. D. 
australis is the largest known Marsupial, being fully equal in bulk 
to a Rhinoceros. It may be regarded as the type of a family— 
Diprotodontide—having affinity on the one hand with the Phalangers 
and on the other with the Kangaroos. 

Nototherium.2—Represented by a species of somewhat smaller 
size than the type of Diprotodon, with a shorter skull, in which the 
zygomatic arches are very wide and the nasals curiously expanded 
at their extremities. The mandibular symphysis is ankylosed ; 

1 Owen, in Mitchell’s Eastern Australia, 2d ed. vol. ii. p. 362 (1838). 

2 Owen, Cat. Mamm. and Aves, Mus. R. Coll. Surgeons, p. 314 (1845). 


172 MARSUPIALIA 


and, as in Diprotodon, there appears to have been no tooth-change. 
The humerus probably referable to Nototherium is of a short and 
widely expanded type, with a large entepicondylar foramen, and 
coming nearer to that of the Wombat than to that of any other 
existing form. The Nototheriide may apparently be regarded as a 
distinct family connecting the Diprotodontide with the Phasco- 
lomyide and Phalangeride. 


Bibliography of Marsupialia.—G. R. Waterhouse, Nat. Hist. of the Mammalia, 
vol. i, ‘‘Marsupiata,” 1846 ; J. Gould, Mammals of Australia, 1863 ; R. Owen, 
article ‘‘Marsupialia,” in Cyclop. of Anatomy and Physiology, and various 
memoirs ‘‘On Extinct Mammals of Australia” in Philosophical Transactions ; 
W. H. Flower, ‘‘On the Development and Succession of the Teeth in the Mar- 
supialia,” Phil. Trans. 1867 ; O. Thomas, ‘‘On the Homologies and Succession 
of the Teeth in the Dasyuride,” Phil. Trans. 1887; and ‘‘Catalogue of Mar- 
supialia and Monotremata in the British Museum,” 1888. 


CHAPTER VII 
THE SUBCLASS EUTHERIA AND THE ORDER EDENTATA 


Tue whole of the remaining groups of mammals are included in a 
single subclass, known by the names Eutheria, Monodelphia, or 
Placentalia.! The one distinctive feature they have in common 
(from which the last-mentioned name is derived) is the presence of 
an allantoic placenta by means of which the foetus is nourished within 
the uterus of the mother. Throughout the entire subclass, as a general 
rule, the urino-genital organs open quite independently of the rectum ; 
the corpus callosum of the brain is well developed ; the mandible does 
not show a marked inflection of its angle; and distinct epipubic 
bones are not attached to the anterior margin of the pubic symphysis. 
In those cases where there is a heterodont and diphyodont dentition 
the dental formula can be reduced to some modification of the one 
given on p. 25, there being only one known genus where four 
true molars occur, and even that not invariably. As in the 
Metatheria, the coracoid is reduced to a mere appendage of the 
scapula, and the acetabular cavity of the pelvis is imperforate. 
While the survivors of the other subclasses have probably been 
for a long time in a stationary condition, these have, as there is 
already good evidence to show throughout all the Tertiary 
geological age, and by inference for some time before, been multi- 
plying in numbers and variations of form, and attaining higher 
stages of development and specialisation in various directions. 
They consequently exhibit far greater diversity of external or 
adaptive modification than is met with in either of the other sub- 
classes,—some being fitted to live as exclusively in the water as 
fishes, and others to emulate the aerial flight of birds. 

To facilitate the study of the different component members 
of this large group, it is usual to separate them into certain 


1 The characters of the chief groups of the Eutheria here given are, in some 
measure, a fuller recapitulation of those already detailed in Chapter III, pp. 
83-88. 


174 EUTHERIA 


divisions which are called “orders.” In the main zoologists 
are now of accord as to the general number and limits of these 
divisions among the existing forms, but the affinities and relation- 
ships of the orders to one another are far from being understood, and 
there are very many extinct forms already discovered which do not 
fit at all satisfactorily into any of the orders as commonly defined. 

Commencing with the most easily distinguished, we may first 
separate a group called Edentata, composed of several very distinct 
forms, the Sloths, Anteaters, and Armadillos, which under great 
modifications of characters of limbs and digestive organs, as well as 
habits of life, have just enough in common to make it probable that 
they are the very specialised survivors of an ancient group, most 
of the members of which are extinct, although the researches of 
paleontology have not yet revealed them to us. The characters of 
their cerebral, dental, and in many cases of their reproductive organs 
show an inferior grade of organisation to that of the generality of 
the subclass. The next order, about the limits of which there is no 
difficulty, is the Sirenia,—aquatic vegetable-eating animals, with 
complete absence of hind limbs, and low cerebral organisation,— 
represented in our present state of knowledge by but two existing 
genera, the Dugongs and Manatees, and by a few extinct forms, 
which, though approaching a more generalised mammalian type, 
show no special characters allying them to any of the other orders. 
Another equally well-marked and equally isolated, though far more 
numerously represented and diversified order, is that of the Cetacea, 
composed of the various forms of Whales, Dolphins, and Porpoises. 
In aquatic habits, external fish-like form, and absence of hind limbs, 
they resemble the last, though in all other characters they are 
as widely removed as are any two orders among the Eutheria. 

All the remaining orders are more nearly allied together, the 
steps by which they have become modified from one general 
type being in most cases not difficult to realise. Their dentition 
especially, however diversified in detail, always responds to the 
formula already alluded to, and, although the existing forms are 
broken up into groups in most cases easy of definition, the discoveries 
already made in paleontology have in great measure filled up the 
gaps between them. 

Very isolated among existing Eutheria are the two species of 
Elephant constituting the group called Proboscidea. These, however, 
are now known to be the survivors of a large series of similar animals, 
Mammoths, Mastodons, and Dinotheres, which as we pass backwards 
in time gradually assume a more ordinary or generalised type ; and 
the interval which was lately supposed to exist between even these 
and the rest of the class is partially bridged over by the discovery 
in American Eocene and early Miocene formations of the gigantic 
Dinocerata, evidently offshoots of the great group of hoofed animals, 


ORDERS 175 


or Ungulata, represented in the actual fauna by the Horses, 
Rhinoceroses, Tapirs, Swine, and Ruminants. Almost as isolated 
as the Proboscidea among existing mammals are the few small 
species constituting the family Hyracide, and in their case palxon- 
tology affords no help at present, and therefore, pending further dis- 
coveries, it has been thought advisable in most recent systems to 
give them the honour of an order to themselves, under the name of 
Hyracoidea. But the number of extinct forms already known allied 
to the Ungulata, though not coming under the definition of either 
of the two groups (Artiodactyla and Perissodactyla) under which all 
existing species range themselves, is so great that either many new 
orders must be made for their reception or the definition of the old 
order Ungulata so far extended as to receive them all, in which 
case both Proboscidea and Hyracoidea may be included within it. 
Again, the Rodentia or gnawing animals—Rabbits, Rats, Squirrels, 
Porcupines, Beavers, etc.—are, if we look only at the present state 
of the class, most isolated. No one can doubt what is meant by a 
Rodent animal, or have any difficulty about defining it clearly, at 
least by its dental characters ; yet owr definitions break down before 
the extinct South American Typotherium, half Rodent and_ half 
Ungulate, which leads by an easy transition to the still more truly 
Ungulate Towodon, for the reception of which a distinct order 
(Toxodontia) has been proposed. It has also been suggested that 
the Rodents are connected by some of the extinct Tillodontia (or 
Teeniodontia) with the Edentates. The Insectivora and the 
Carnivora again are at present quite distinct orders, but they merge 
into one another through fossil forms, and are especially connected 
by the large group of primitive Carnivora, so abundantly repre- 
sented in the Eocene deposits both of America and Europe, to which 
Cope has given the name of Creodonta. The Carnivora also appear 
to have been closely connected with the primitive Ungulates as repre- 
sented by the extinct group called Condylarthra. In another 
direction the step from the Insectivores to the Lemurs is not great, 
and in past times the transition was probably complete. The Bats 
or Chiroptera are allied to the Insectivora in all characters except the 
extraordinary modification of their anterior extremities into wings ; 
but this, like the want of the hind limbs in the Cetacea and Sirenia, 
makes such a clear distinction between them and all other mammals 
that, in the absence of any knowledge of any completely inter- 
mediate or transitional forms, they can be perfectly separated, and 
constitute as well-defined an order as any in the class. We have, 
however, an inkling of the mode in which the Insectivora were 
modified into Chiroptera shown us by the so-called Flying Lemur 
(Galeopithecus). Finally, we have the important and well-character- 
ised group called Primates, including all the Monkeys and Man ; 
and the question is not yet solved as to how and through what 


176 EDENTATA 


forms this is linked on to the other groups. It is commonly assumed 
that the Lemurs are nothing more than inferior Primates, but the 
interval between them in the actual fauna of the world is very great, 
and our knowledge of numerous extinct types recently discovered 
in America, said to be intermediate in characters, is not yet 
sufficient to enable us to form a definite opinion upon the subject. 
The Edentata may be taken first as standing in some respects 
apart from all the others; and the Primates must be placed at the 
head of the series. The position of the others is quite arbitrary, as 
none of the hitherto proposed associations of the orders into larger 
groups stand the test of critical investigation, and paleontological 
researches have already gone far to show that they are all modifica- 
tions of a common heterodont, diphyodont, pentadactylate form. 


Order EDENTATA. 


The name assigned to this group (which some zoologists think 
ought rather to be ranked as a subclass! than an order) by Cuvier 
is often objected to as inappropriate—for although some of the 
members are edentulous, others have very numerous teeth—and the 
Linnean name Bruta is occasionally substituted. But that term is 
quite as objectionable, especially since the group to which Linnzus 
applied it is by no means equivalent to the order as now understood, 
as the names of the genera contained in it, viz. Elephas, Trichechus, 
Bradypus, Myrmecophaga, Manis and Dasypus, indicate. It contained, 
in fact, all the animals then known which are comprised in the 
modern groups of Proboscidea, Sirenia and Edentata together with 
the Walrus, one of the Carnivora. If retained at all, it should 
rather belong to the Proboscidea, as Elephas stands first in the 
list of genera in the Systema Nature. Cuvier’s order included the 
Ornithorhynchus and Echidna, the structure of which was then im- 
perfectly known, and which are now by common consent removed 
to an altogether different section of the class; but otherwise its 
limits are those now adopted. The name Edentata is so generally 
used, and its meaning so well understood, that it would be un- 
desirable to change it now ; in fact similar reasons might be assigned 
for ceasing to use nearly all the other current ordinal designations, 
for it might be equally well objected that all Carnivora are not 
flesheaters, many of the Marsupialia have not pouches, and so 
forth. 

If the teeth are not always absent, they invariably exhibit 
certain imperfections, which are indeed almost the only common 
characters binding together the various extinct and existing members 
of the order. These are—that they are homodont and, with the 


1 The name Paratheria has heen suggested for this proposed subclass 


GENERAL CHARACTERS 177 


remarkable exceptions of Tatusta and Orycteropus, monophyodont ; 
they are never rooted, but have persistent pulps; except in some 
fossil forms, they are always deficient in one of the constituents 
which enter into the formation of the complete mammalian 
tooth, the enamel; and, at least among living forms, are never 
present either in the upper or lower jaw in the fore part of 
the mouth, the situation occupied by the incisors of other 
mammals.1 

The peculiar nature of the dentition in the aberrant Orycteropus 
will be noticed under the heading of that genus. As a rule, the 
coracoid process of the scapula of the Edentates is more developed 
than in other Eutheria. 

The degree of development of the brain varies considerably in 
the different families, the 
hemispheres being in some 
cases almost or quite smooth 
(Fig. 57), with a small corpus 
callosum, and large anterior 
commissure ; while in other 
instances the hemispheres 
are convoluted, and the 
corpus callosum is larger. 

There is so great a differ- 
ence in structure and habits 
between some of the existing 
animals assigned tothis order 


Fic. 57.—Upper surface of the brain of the Broad- 
7 banded Armadillo (Xenurus wnicinctus). The large 
that, beyond the negative olfactory lobes are seen at the anterior extremity 


characters just mentioned, (left of figure); the hemispheres have only three 


there seems little to connect sulci. (From Garrod, Proc. Zool. Soc. 1878, p. 230.) 


them. The Sloths and Anteaters, for instance, in mode of life, 
general conformation of limbs, structure of digestive organs, etc., 
appear at first sight almost as widely separated as any mammals. 
Paleontology has, however, thrown great light upon their relations, 
and proved their real affinities. Perfectly intermediate forms have 
been discovered in the great Ground Sloths of America, which have 
the dentition and general form of the head of the Sloths, combined with 
the limbs and trunk of the Anteaters. It is, indeed, highly probable 
that the existing members of this order are very much differentiated 
representatives of a large group, the greater number of which are 
now extinct, and have become so without ever attaining a high 
grade of organisation. The great diversity of structure in the 
existing families, the high degree of specialisation to which many 
have attained, the paucity of species and even of individuals, their 


1 In some few Armadillos the suture between the premaxilla and maxilla 
passes behind the first upper tooth ; but in all other known members of the order 


all the teeth are implanted in the maxilla. 
12 


178 EDENTATA 


limited area of distribution, and their small size compared with 
known ancestral forms, all show that this is an ancient and a waning 
group, the members of which seem still to hold their own either by 
the remoteness and seclusion of their dwelling-places, by their 
remarkable adaptation of structure to special conditions of life, or 
by aid of the peculiar defensive armature with which they are 
invested. Their former history can, however, only be thus surmised, 
rather than read, at present ; for, though we have ample evidence 
of the abundance and superior magnitude of certain forms in the 
most recent or Pleistocene geological age, yet we have at present 
no definite evidence as to their origin, or relationship to other 
orders of mammals. 

The existing members of the order readily group themselves 
into five distinct families, the limits of which are perfectly clear. 
These are (1) Bradypodide, or Sloths; (2) Myrmecophagide, or Ant-. 
eaters ; (3) Dasypodide, or Armadillos; (4) Manide, Pangolins or 
Scaly Anteaters; and (5) Orycteropodide, Aard-varks or African 
Anteaters. The geographical distribution of these families coincides 
with their structural distinction, the first three being inhabitants of 
the New and the last two of the Old World. It has been usual to 
arrange these families into two large groups or suborders: (1) the 
Phyllophaga, leaf-eaters, also called Tardigrada, containing the 
Bradypodide alone; and (2) the Entomophaga, insect-eaters, or 
Vermilingua, containing all the other families, from which some- 
times the Orycteropodide are separated as a third suborder under 
the name of Effodientia, or Tubulidentata. Such an arrangement 
is, however, an artificial one, founded on superficial resemblance. 
The bonds which unite the Munidw to the Myrmecophagide are 
mainly to be found in the structure of the mouth, especially the 
extensile character of the tongue, the great development of the sub- 
maxillary glands, and the absence of teeth. These characters are 
exactly analogous to those found in the Echidna among Monotremes, 
the Woodpeckers among Birds, and the Chameleon among Reptiles, 
—the fact probably being that in countries where Termites and 
similar insects flourish various distinct forms of vertebrates have 
become modified in special relation to this abundance of nutritious 
food, which could only be made available by a peculiar structure of 
the alimentary organs. A close study of the more essential 
portions of the anatomy of these animals! leads to the belief 
that all the American Edentates at present known, however di- 
versified in form and habits, belong to a common stock. Thus the 
Bradypodide, Megatheriide, and Myrmecophagide are certainly allied, 
the modifications seen in the existing families relating only to food 
and manner of life. The ancestral forms may have been omni- 


1 See Flower, ‘‘On the Mutual Affinities of the Animals composing the 
Order Edentata,” Proceedings of the Zoological Society, 1882, p. 358. 


BRADVPODIDE 179 


vorous, and gradually separated into the purely vegetable and 
purely animal feeders; from the former are developed the modern 
Sloths, from the latter the Anteaters. The Armadillos (Dasypodide) 
are another modification of the same type, retaining some 
generalised characters, as those of the alimentary organs, but in 
other respects, as in their defensive armature, remarkably special- 
ised. The two Old World families Manide and Orycteropodide are 
so essentially distinct, both from the American families and from 
each other, that it may even be considered doubtful whether they 
are derived from the same primary branch of mammals, or whether 
they may not be offsets of some other branch, the remaining 
members of which have been lost to knowledge. Further remarks on 
this point are recorded under the description of the Orycteropodide.+ 


Family BRADYPODIDZ. 


Externally clothed with long, coarse, crisp hair. Head short 
and rounded. External ears inconspicuous. Teeth $ in each jaw, 
subcylindrical, of persistent growth, consisting of a central axis of 
vaso-dentine, with a thin investment of hard dentine, and a thick 
outer coating of cement ; without (so far as is yet known) any suc- 
cession. Clavicles present. Fore limbs greatly longer than the 
hind limbs. All the extremities terminating in narrow, curved 
feet; the digits never exceeding three in number, encased for 
nearly their whole length in a common integument, and armed 
with long strong claws. Tailrudimentary. Stomach complex. No 
cecum. Uterus simple and globular. Placenta deciduate, dome-like, 
composed of an aggregation of numerous discoidal lobes. Strictly 


1 An attempt has been made to represent these views by the following 
classification : 
Order EDENTATA, 

Suborder Pinosa. 
Bradypodide. 
Megatheriide. 
Myrmecophagide. 

Suborder Loricara. 
Dasypodide. 

Suborder SQUAMATA. 
Manide. 

Suborder TUBULIDENTATA. 
Orycteropodide. 


It may be objected to this arrangement that the present divergence between 
the Sloths and Anteaters is hardly sufficiently indicated by their association in 
one suborder.—Flower, ‘‘On the Arrangement of the Orders and Families of 
Mammals,” Proc. Zool. Soc. 1883, p. 178. 


180 EDENTATA 


arboreal in habits, vegetable feeders, and limited geographically to 
the forest regions of South and Central America. 

The Sloths, as the animals of this family are called on account 
of the habitual sluggishness of their movements, are the most strictly 
arboreal of all mammals, living entirely among the branches of 
trees, usually hanging under them, with their backs downwards 
(Fig. 58), and clinging to them with the simple hook-like organs to 
which the terminations of all their limbs are reduced. When they 
are obliged from any cause to descend to the ground, which they 
rarely, if ever, do voluntarily, their limbs, owing to their unequal 
length and the peculiar conformation of the feet—which allows 
the animals to rest only on the outer edge—are most inefficient 


Fic. 58.—Two-toed Sloth (Cholepus hoffmanni). 


for terrestrial progression, and they crawl along a level surface 
with considerable difficulty. Though generally slow and inactive, 
even when in their natural haunts, Sloths can on occasions travel 
with considerable rapidity along the branches; and, as they do not 
leap, like most other arboreal creatures, they avail themselves of 
the swaying of the boughs by the wind to pass from tree to tree. 
They feed entirely on leaves and young shoots and fruits, which 
they gather in their mouth, the fore limbs aiding in dragging 
boughs within reach, but not being used like hands, as they are by 
monkeys, squirrels, etc. When sleeping they roll themselves up in 
a ball, and, owing to the dry shaggy character of their hair, are 
very inconspicuous among the mosses and lichens with which the 


BRADYPODIDE 181 


trees of their native forests abound ; the concealment thus afforded 
being heightened in some species by the peculiar greenish tint 
of the outer covering—very uncommon in mammals. This is not 
due to the colour of the hair itself, but to the presence upon its 
surface of an alga, the lodgment of which is facilitated by the fluted 
or rough surface of the exterior of the hair, and the growth of which 
is promoted by the dampness of the atmosphere in the gloomy 
tropical forests, as it soon disappears from the hair of animals kept 
in captivity in England. Sloths are nocturnal, silent, inoffensive, and 
solitary animals, and usually produce but one young at birth. They 
appear to show an almost reptilian tenacity of life, surviving the 
most severe injuries and large doses of poisons, and exhibiting 
longer persistence of irritability of muscular tissue after death than 
other mammals. 

In the Bradypodide, as well as in the Myrmecophagide, the 
testes are placed close to each other, lying on the rectum between 
it and the bladder; the penis is quite rudimentary, consisting 
of a pair of small corpora cavernosa, not directly attached by their 
crura to the rami of the ischia, and having a glans scarcely larger 
than that of the clitoris of most mammals, and, as in birds and 
reptiles, without any true corpus spongiosum. In the females of 
both families the uterus is simple and globular ; and the vagina, at 
least in the virgin state, is divided into two channels by a strong 
median partition. The deciduate placenta of Cholepus is composed 
of a number of lobes aggregated into a dome-like mass; and it 
does not appear that the placenta of the Anteaters departs in any 
important characters from this type. According to the late Pro- 
fessor W. K. Parker, the embryos of the Sloths, Anteaters, and 
Pangolins have the stapes of the middle ear in the form of a rod, 
thus showing affinities with a very primitive type of mammalian 
organisation. 

The Sloths were all included in the Linnean genus Bradypus, 
but Illiger very properly separated the species with but two claws 
on the fore feet, under the name of Cholepus, leaving Bradypus 
for those with three. 

Bradypus..—Three-toed Sloths. Teeth usually 2 on each side ; 
no tooth projecting greatly beyond the others; the first in the 
upper jaw much smaller than any of the rest; the first in the 
lower jaw broad and compressed ; the grinding surfaces of all much 
cupped. Vertebre: C9, D and L 20 (of which 15 to 17 bear ribs), 
$6, Cll. All the known species present the remarkable pecu- 
liarity of possessing nine cervical vertebra, 7.c. nine vertebrae 
in front of the one which bears the first thoracic rib (or first 
rib connected with the sternum, and corresponding in its general 
relations with the first rib of other mammals); but the ninth. 


1 Linn. Syst. Nat, 12th ed. vol. i. p. 50 (1766), 


182 EDENTATA 


and sometimes the eighth, bears a pair of short movable ribs. 
The arms or fore limbs are considerably longer than the hind 
legs. The bones of the fore arm are complete, free, and capable of 
pronation and supination. The hand is long, very narrow, habit- 
ually curved, and terminates in three pointed curved claws, in 
close apposition with each other. The claws are, in fact, incapable of 
being divaricated, so that the hand is reduced to the condition of a 
triple hook, fit only for the function of suspension from the boughs 
of trees. The foot closely resembles the hand in its general struc- 
ture and mode of use; the sole being habitually turned inwards, so 
that it cannot be applied to the ground in walking. The tongue is 
short and soft, and the stomach large and complex, bearing some 
resemblance to that of the ruminating Ungulates. The windpipe 
or trachea has the remarkable peculiarity among mammals—not 
unfrequent among birds and reptiles—of being folded on itself 
before it reaches the lungs. The mamme are two, and pectoral in 
position. 

“ Ai” is the common name given in books to the Three-toed 
Sloths. They were all comprised by Linnzus under the species 
Bradypus tridactylus. More recently Dr. Gray described as many 
as eleven species, ranged in two genera, Bradypus and Arctopithecus ; 
but the distinctions which he assigned both to species and genera do 
not bear close examination. Some are covered uniformly with a 
gray or grayish-brown coat; others have a dark collar of elongated 
hairs around the shoulders (B. forquatus); some have the hair of 
the face very much shorter than that of the rest of the head and 
neck ; and others have a remarkable-looking patch of soft short hair 
on the back between the shoulders, consisting, when best marked, 
of a median stripe of glossy black, bordered on each side by bright 
orange, yellow, or white. There are also structural differences in 
the skulls, as in the amount of inflation of the pterygoid bones, 
which indicate real differences of species; but the materials in our 
museums are not yet sufficient to correlate these with external 
characters and geographical distribution. The habits of all are 
apparently alike. They are natives of Guiana, Brazil, and Peru, 
and one if not two species (B. infuscatus and B. castuneiceps) extend 
north of the Isthmus of Panama as far as Nicaragua. Of the 
former of these Dr. Seeman says that, though generally silent, 
a ‘specimen in captivity uttered’ a shrill sound like a monkey 
when forcibly pulled away from the tree to which it was 
holding. 

Choleyus..—Teeth $; the most anterior in both jaws separated 
by an interval from the others, very large, caniniform, wearing 
to a sharp, bevelled edge against the opposing tooth, the upper 
shutting in front of the lower when the mouth is closed (Fig. 59), 

' Wliger, Prodromus Syst. Mumm. ct Avium, p. 108 (1811). 


MEGATHERIUD.E 183 


unlike the true canines of heterodont mammals. Vertebre: C 6 
or 7, D23-24, L3, 87-8, C 4-6. One species (C. didactylus) has 
the ordinary number of vertebra in the neck; but an otherwise 
closely allied form (C. hoffmanni) has but six. The tail is very 
rudimentary. The hand generally resembles that of Bradypus ; but 
there are only two functional digits with claws—those answering 
to the second and third of the typical pentadactylate manus. The 
structure of the hind limb generally resembles that of Bradypus, 
the appellation “two-toed” referring only to the anterior limb, 
for in the foot the 
three middle toes 
are functionally 
developed and of 
nearly equal size. 
C. didactylus, which 
has been longest 
known, is com- 
monly called by 
the native name 
of Unau. It in- 
habits the forests 
of Brazil. C. hoff- 
mann (Fig. 58) 
has a more north- 
ern geographical Fic. 59.—Skull Of Bwo-toed Sloth (Cholospus didactylus). From 
< Proc. Zool. Soc. 1871, p. 432. 

range, extending 

from Ecuador through Panama to Costa Rica. Its voice, which 
is seldom heard, is like the bleat of a sheep, and if the animal is 
seized it snorts violently. Both species are very variable in 
external coloration. 

Nothropus.1—The only fossil form which has been referred to 
this family is indicated by a lower jaw, described by Dr. Burmeister, 
from the Pleistocene of Argentina, which appears to have belonged 
to an animal of about double the dimensions of Cholapus didactylus. 
Professor Cope states, however, that this jaw really belongs to a 
Glyptodont; while it is referred by Dr. Ameghino to the next 
family. 


Family MEGATHERIID-£. 


The members of this family are all extinet. Their characters, 
so far as is known from the well-preserved remains of many species 
found abundantly in deposits of Pleistocene age in both North and 
South America, were intermediate between those of the existing 
Bradypodide and the Myrmecophagide, combining the head and 

1 Burmeister, Sitzb. lk. Berlin, vol. xxviii, p. 613 (1882). 


184 EDENTATA 


dentition of the former with the structure of the vertebral column, 
limbs, and tail of the latter. Almost all the known species are of 
comparatively gigantic size, the smallest, Nothrotheriwm escrivanense, 
exceeding the largest existing Anteater, and the Megatherium 
being larger than a Rhinoceros. The femur has no third trochanter, 
and the odontoid process of the axis vertebra has a peculiar facet 
on the ventral surface. The dentition is usually = on each side, as 
in the Sloths, but 4 in Nothrotherium.! This genus, and in a still 
more marked degree Megatherium, differ from all the others in the 
details of the structure of the teeth. They are very deeply 
implanted, of prismatic form (quadrate in transverse section), and 
the component tissues—hard dentine (Fig. 60, d), softer vaso-dentine 


5 


EES 


PED iS SYA 


[Ltt Pee 


Fic. 60.—Section of upper molar teeth of Megatherium americanum. x}. 
p, pulp-cavity ; the other letters explained in the text. (After Owen.) 


(v), and cement (c)—are so arranged that, as the tooth wears, the 
surface always presents a pair of transverse ridges, thus producing 
a triturating apparatus comparable to the “bilophodont” molar of 
Dinotherium, Tapirus, Manatus, Macropus, and others, though pro- 
duced in a different manner. In all the other genera the teeth are 
more or less cylindrical, though sometimes laterally compressed or 
even longitudinally grooved on the sides, and on the erinding 
surface the prominent ridge of hard dentine follows the eden 
contour, and is surrounded only by a thin layer of cement, as 
in the existing Sloths. The Ground Sloths, as the mem heve 


1 Lydekker, in Nicholson and Lydekker’s Manual of Paleontology, vol. ii. 
p. 1299 (1889). Originally described under the preoccupied name Cwlodon. 


MEGATHERIIDA 185 


of this family may be conveniently designated, agree with the 
Sloths and Anteaters, and thereby differ from all other mammals, 
in that the coracoid process of the scapula and the coracoidal 
border of the same unite over the coraco-scapular notch, 
which is thus converted into a foramen. Large clavicles are 
present. 

Megatherium..cThe typical genus Megatherium, as being the 
longest known representative of the family, may be noticed in some 
detail. A nearly complete skeleton, found on the banks of the 
River Luxan, near Buenos Ayres, and sent in 1789 to the Royal 
Museum at Madrid, long remained the principal if not the only 
source of information with regard to the species to which it belonged, 
and furnished the materials for many descriptions, notably that of 
Cuvier, who determined its affinities with the Sloths.2 In 1832 an 
important collection of bones of the Megatherium was discovered 
near the Rio Salado, and secured for the Museum of the College 


Fic. 61.—Oral surface of mandible of Megatherium americanum. 
a, Condyle ; b, masseteric process; c, angle; d, symphysis. (After Owen.) 


of Surgeons of England ; and these, with another collection found 
at Luxan in 1837, and now in the British Museum, supplied the 
materials for the complete description of the skeleton published 
by Sir R. Owen in 1861. Other skeletons have subsequently been 
received by several of the Continental museums, as Milan and Paris, 
and also by those in South America; and consequently our know- 
ledge of the organisation of the Megatherium, so far as it can be 
deduced from the bones and teeth, is as complete as that of any 
other animal, recent or extinct. 

The remains hitherto spoken of are all referred to one species, 
Megatherium americanum of Blumenbach (Jf. cuviert of Desmarest), 
and are all from the newest or Pleistocene geological formations of 
the Argentine Republic and Paraguay, or the lands forming the 


1 Cuvier, Tableaw Elém. d’ Hist. Nat. des Animaux, p. 146 (1798). 

? An excellent figure of this skeleton, which unfortunately was incorrectly 
articulated, and wanted the greater part of the tail, was published by Pander 
‘and D’Alton in 1821, and has been frequently reproduced in subsequent 
works. 


186 EDENTATA 


basin of the Rio de la Plata. Dr. Leidy has described, from similar 
formations in Georgia and South Carolina, bones of a closely allied 
species, about one-fourth smaller, which he has named MZ. mirabile. 
Three other South American species have been described; but JZ. 
lawrillardi, of Lund, founded upon remains found in Brazil, has 
been made the type of the genus Ocnopus. 

The following description will apply especially to the best-known 
South American form, Megatherium americanum. In size it exceeded 
any existing land animal except the elephant, to which it was 
inferior only in consequence of the comparative shortness of its 
limbs ; for in length and bulk of body it was its equal, if not 


Fic. 62.—Skeleton of Megatherium, from the specimen in the Museum of the Royal College 
of Surgeons. x 34. 


superior. The full length of a mounted skeleton (Fig. 62), from 
the fore part of the head to the end of the tail, is 18 feet, of which 
the tail occupies 5 feet. The head, which is small for the size of 
the animal, presents a general resemblance to that of the Sloth ; 
the anterior part of the mouth is, however, more elongated, and the 
jugal bone, though branched posteriorly in the same way as that of 
the Sloth, meets the zygomatic process of the squamosal, thus 
completing the arch. The lower jaw has the middle part of its 
horizontal ramus curiously deepened, so as to admit of im- 
plantation of the very long-rooted teeth, the peculiar structure 
of which has been already described. A skull recently discovered 
shows that, instead of the wide gap between the extremity of 
the nasals and the premaxille exhibited in Fig. 62, there was 
a prenasal bone, towards which a process extended upwards and 


MEGATHERIDA 187 


backwards from the extremity of the upper surface of the pre- 
maxille. 

The vertebral column consists of seven cervical, sixteen dorsal, 
three lumbar, five sacral, and eighteen caudal vertebre. The 
spinous processes are much better developed than in the Sloths, 
and are all directed backwards, there being no reversing of the 
inclination near the posterior end of the dorsal series, as in most 
active-bodied mammals. In the lumbar region, the accessory zyga- 
pophyses, rudimentary in Sloths, are fully developed, as in the 
Anteaters. 

The tail is large, and its basal vertebrae have strong lateral and 
spinous processes and chevron bones, indicating great muscular 
development. The scapula resembles that of the Sloths in the 
union of the acromion with the coracoid, and in the bridging over 
of the supra-scapular notch. The clavicle is complete and very 
large, much resembling that of man on a large scale. The fore 
limbs are longer than the hind limbs. The humerus has no ent- 
epicondylar foramen. The radius and ulna are both well developed, 
and have a considerable amount of freedom of movement. The 
hand is singularly modified. The pollex is represented only by a 
rudimentary metacarpal, but the next three digits are large, and 
terminate in phalanges adapted for the support of immense claws, 
the middle one being especially large. The outer or fifth digit has 
no claw, and it may be considered as certain that the weight of the 
foot was, in standing and walking, chiefly thrown upon this one, 
which was protected by a callous pad below, as in the existing 
great Anteater, while the other toes were curved inwards towards 
the palm, and only came in contact with the ground by their outer 
surfaces. The mechanical arrangements by which the weight of the 
body was thrown entirely upon the outer side of the foot are very 
curious, and are fully described in Owen’s memoir. The pelvis is 
remarkably wide, even more so than that of the Elephant, but it is 
formed on the same principle as in the Sloths. The femur is 
extremely broad and flattened; the tibia and fibula are short and 
strong, and united together at each end. The hind foot, contrary 
to the usual rule in the Edentata, is even more singularly modified 
than the hand. Thus the ankle-joint is formed upon a peculiar 
plan, quite unlike that of the Sloths, or of any other mammal, except 
the Megatherium’s nearest allies ; and the caleaneum projects nearly 
as far backwards as the fore part of the foot does forwards. There 
is no trace of great toe or hallux, or of its corresponding cuneiform 
bone ; the second toe is rudimentary ; while the third has an enor- 
mous ungual phalanx, which, as in those of the hand, is remarkable 
for the immense development of the bony sheath reflected from 
its proximal end around the base of the claw. The two outer toes 
have large and very peculiarly-shaped metatarsals, but only small 


188 EDENTATA 


phalanges, and no claws. The creature probably walked upon the 
outer edge of the sole, so that the great falcate claw of the third 
toe did not come into contact with the ground, and so was kept in 
a state of sharpness ready for use. The foot was therefore formed 
upon quite a different principle from that of the Anteaters or 
Sloths, though somewhat like the latter in having two of the toes 
aborted. 

Taking all the various points of its structure together, they 
clearly indicate affinities both with the existing Sloths and with 
the Anteaters, the skull and teeth more resembling those of the 
former, and the vertebral: column and limbs the latter. It is also 
not difficult to infer the food and habits of this enormous creature. 
That it was a leaf-eater there can be little doubt; but the greater 
size and more complex structure of its teeth might have enabled it 
to crush the smaller branches as well as the leaves and succulent 
shoots which form the food of the existing Sloths. It is, however, 
very improbable that it climbed into the branches of the trees like 
its diminutive congeners, and it is far more likely that it obtained 
its subsistence by tearing them down with the great hook-like claws 
of its powerful prehensile fore limbs, being easily enabled to reach 
them by raising itself up upon the massive tripod formed by the 
two hind feet, firmly fixed to the ground by the one huge falcate 
claw, and the stout, muscular tail. The whole conformation of 
the hinder part of the animal is strongly suggestive of such an 
action. There can also be little doubt but that all its move- 
ments were as slow and deliberate as those of its modern repre- 
sentatives. 

An idea at one time prevailed that the Megatherium was 
covered externally with a coat of bony armour like that of the 
Armadillos ; but this originated in dermal plates belonging to the 
Glyptodon having been accidentally associated with bones of the 
Megatherium. Similar plates, on a smaller scale, have indeed been 
found in connection with the skeleton of the Mylodon, but never 
yet with the Megatherium, which we may therefore imagine with 
a covering of coarse hair like that of its nearest living allies, the 
Sloths and Anteaters. 

Scelidotherium, Mylodon, etc.—Of the more important remaining 
genera of this family a briefer notice will suffice. Scelidotherium (in 
which Platyonyx may be included) comprises several species of 
considerably smaller dimensions than the Megatherium, and is in 
some respects intermediate between that genus and Alylodon. The 
teeth have an oval cross-section, like those of the Sloths, while the 
skull, in which the length of the nasals is subject. to great variation 
in the different species, approximates more or less closely to that 
of the Myrmecophagide. The humerus generally has an ent- 
epicondylar foramen ; and the form and relations of the bones of 


MEGATHERIDA 189 


the feet differ considerably from those obtaining in the type genus. 
S. leptocephalum, the type of the genus, occurs in Patagonia and 
Argentina but 
other species are 
found in Brazil 
and Chili, The 
genus Mylodon, in 
its widest sense, 
may be taken to 
include a number 
of comparatively 
large _Edentates, 
some of which have 
been described 
under the names of 
Grypotherium, Lest- 
odon, and Pseudo- 
lestodon. The teeth 
of the upper jaw 
are generally of an 
oval or subtriangu- 
lar section ; and in 
the more typical forms the first and second teeth are separated 
by a short interval, the ‘former being horizontally worn. In 
other species, however, like Jf. (Lestodon) armatus, there is a 
considerable space between the first and second teeth, and the 
first is worn obliquely. The skull is exceedingly like that of 
the Sloths in general contour; and there is not the descending 
process at the angle of the mandible found in Megatheriwm. 
The humerus has no entepicondylar foramen. The species 
represented in Fig. 63 is from the Pleistocene of South America ; 
but the type of the genus is M. harlam, from beds of corre- 
sponding age in Kentucky. The Patagonian JM. (Grypotherium) 
darwini is a remarkable form, characterised by the presence of a 
bony arch connecting the premaxille with the nasals, of which, as 
already mentioned, there is an {incomplete development in 
Megatherium.  Megalonyz, from the Pleistocene of Kentucky, differs 
from Mylodon by the long interval between the first and second 
teeth, and also by the presence of an entepicondylar foramen in 
the humerus. Nothrotheriwm is a smaller form, occurring in the 
deposits of the Brazilian caves, of which the dental features have 
been already mentioned. The osteological characters of these and 
other allied genera have been fully described in the works of 
Cuvier, Owen, Burmeister, Leidy, Ameghino, Gervais, Reinhardt, 
and others. 

Promegatherium.Two genera from the infra-Pampean beds 


Fic. 63.—Skeleton of Mylodon robustus (Pleistocene, South 
America). From Owen. 


190 EDENTATA 


of Argentina, described as Promegatherium and Promylodon, are 
respectively distinguished from Jegatherium and \ylodon by 
the presence of bands of enamel on the teeth, which points 
to the descent of the Edentates from mammals with enamelled 
teeth. 

The Tertiary North American forms described as l/oropus and 
Morotherium,! and originally regarded as Edentates, would appear to 
be aberrant Ungulates. 


Family MYRMECOPHAGID. 


Externally clothed with hair. No teeth. Head elongated. 
Mouth tubular, with a small terminal aperture, through which the 
long, vermiform tongue, covered with the viscid secretion of the 
enormous submaxillary glands, is rapidly protruded in feeding, and 
withdrawn again with the adhering particles of aliment, which are 
then sucked into the pharynx. Clavicles rudimentary. In the 
manus, the third toe is greatly developed, and has a long falcate 
claw; the others are reduced or suppressed. The pes has four or 
five subequal digits with claws. Posterior dorsal and lumbar 
vertebr, with additional interlocking zygapophyses. Tail long, 
sometimes prehensile. Uterus simple. Placenta dome-like or 
discoidal. Brain fairly convoluted, and with a large corpus cal- 
losum and anterior commissure. The animals of this family are 
the “Anteaters” par evrcellence. They feed exclusively on animal 
substances, mostly insects. One species is terrestrial, the others 
arboreal ; none burrow in the ground. They are all inhabitants of 
the Neotropical region. 

The reproductive organs, as noticed on p. 181, are of the 
same general type as in the Bradypodide. 

Myrmecophaqu.n—Skall greatly elongated and narrow, its upper 
surface smooth and cylindriform. Anteriorly the face is produced 
into a long, tubular rostrum, rounded above and flattened below, 
with terminal nares, and composed of the mesethmoid ossified 
for more than half its length, the vomer, the maxillez, and the long 
and narrow nasal bones, the premaxille being extremely short and 
confined to the margin of the anterior nares. The zygomatic arch 
is incomplete, the styliform jugal only articulating with the maxilla 
in front, and not reaching to the very short zygomatic process of 
the squamosal. The lachrymal foramen is in front of the margin of 
the orbit. There are no postorbital processes to the frontals, i any 
other demarcation hetween the orbits and the temporal fosse. Palate 
extremely elongated, and produced backwards as far as the level of 


1 See E. D. Cape, licr. Naturalist, vol. xxiii. p. 152 (1889, 
* Linn. Syst. Vat, 12th ed. vol. i. p. 51 (1766) 


MYRMECOPHAGIDA I9I 


the external auditory meatus by the meeting in the middle line of 
the largely developed pterygoids. The glenoid fossa a shallow oval 
facet, with its long diameter from before backwards. Mandible very 
long and slender, with an exceedingly short symphysis, no distinct 
coronoid process, and a slightly elevated, elongated, flattened, con- 
dylar articular surface. Vertebre: C7, D 15-16, L 3-2,8 6, C31. 
Clavicles rudimentary. In the manus the first digit is very 
slender, the second also slender, with compressed phalanges of nearly 
equal length. The third digit is immensely developed ; though its 
proximal phalanx is extremely short, its ungual phalanx is so long 
that the entire length of the digit exceeds that of the second. The 
fourth has a long and rather slender metacarpal, and three 
phalanges diminishing in size, the ungual phalanx being very 
small. The fifth has the metacarpal nearly as long, but not so 
stout, as the fourth, and followed by two small phalanges, the last 
rudimentary and conical. Claws are developed upon all but the fifth. 
In walking the toes are kept strongly flexed, and have their points 
turned upwards and inwards, the weight being supported upon a 
callous pad over the end of the fifth digit, and by the dorsal sur- 
faces of the third and fourth digits. The hind feet are short and 
rather broad, with five subequal claws, the fourth the longest, the 
first shortest ; the whole sole is placed on the ground in walking. 
Body rather compressed, clothed with long, coarse hair. Tail 
about as long as the body, and covered with very long hair ; not 
prehensile. Ears small, oval, erect. Eyes very small. Stomach 
consisting of a subglobular, thin-walled, cardiac portion, and a 
muscular pyloric gizzard with dense epithelial lining. No ileo- 
colic valve, and a short wide ill-defined cecum. Mammez two, 
pectoral. 

There is one species,! J/. jubata, the Great Anteater, or Ant 
Bear (Fig. 64), measuring 4 feet in length without the tail, and 
upwards of 2 feet in height at the shoulder. Its prevailing colour 
is gray, with a broad black band, bordered with white, commencing 
on the chest, and passing obliquely over the shoulder, diminishing 
gradually in breadth as it approaches the loins, where it ends in a 
point. It is extensively distributed in the tropical parts of South 
and Central America, frequenting low swampy savannas along the 
banks of rivers, and the depths of the humid forests, but is nowhere 
abundant. Its food consists mainly of termites, to obtain which it 
opens their nests with its powerful sharp anterior claws, and as the 
insects swarm to the damaged part of their dwelling, it draws them 
into its mouth by means of its long, flexible, rapidly-moving tongue 
covered with glutinous saliva. The Great Anteater is quite terres- 
trial in its habits, being never known to climb trees, nor does it 


1 Professor Cope has recently come to the conclusion that there are three 
species ; but further evidence is required in support of this view. 


192 EDENTATA 


burrow underground like the Armadillos. Though generally an 
inoffensive animal, when attacked it can defend itself vigorously and 
effectively with its sabre-like anterior claws. The female bears but 
a single young at a birth. 

The union of the pterygoids in the middle line to prolong the 
narial passage is a character found elsewhere among existing mam- 
mals only in the next genus, in one Armadillo (Yatusia), and in 
certain Cetacea. The contrast in length between the skull of the 
Great Anteater and that of the Sloth is, as Professor Parker observes, 
very marked indeed ; the one being relatively the longest and the 


Wy 


WN X 


ANN a 


Fic. 64.—The Great Anteater (Myrmecophaga jubuta). (From Sclater, List of Animals in 
Zoological Society's Gardens, 18S3, p. 190.) 


other almost the shortest in the whole class. The small size and 
incomplete development of the jugal bone in the zygomatic arch 
affords another striking contrast to the Sloths (Fig. 59). 
Tamandua.1—This genus closely resembles the last in anatomical 
structure, but the head is much less elongated, the fur is short and 
bristly, the tail tapering, prehensile, with the under side through- 
out and the whole of the terminal portion naked and scaly. The 
stomach is similar to that of J/yrmecophaga, but with the muscular 
pyloric gizzard not quite so strongly developed. There is a distinct 
ileo-colic valve and a short globular cecum. The fore foot has a very 
large claw on the third toe, moderate-sized claws on the second and 


1 Gray, Annals of Philosophy, new series, vol. x. p. 343 (1825). 


MYVRMECOPHAGIDA 193 


fourth, a very minute one on the first, and none on the fifth, which 
is entirely concealed within the skin. The hind foot has five 
subequal claws. Vertebre: C7, D17,L2,85, C37. There are 
very rudimentary clavicles. 

The Tamandua (Fig. 65) is much smaller than the Great 
Anteater, and differs essentially from it in its habits, being mainly 


Fic. 65.—Tamandua Anteater (Tamandua tetradactylu). From Proce. Zool. Soc. 1871, pl. xliii. 


arboreal. It is an inhabitant of the dense primeval forests of 
South and Central America. As different individuals vary much 
in their coloration, it is possible that there may be more than one 
species. The usual colour is yellowish-white, with a broad black 
lateral band, covering nearly the whole of the side of the body. 
Cycloturus.1—The skull is much shorter even than in Tamandua, 
and is arched considerably in the longitudinal direction. It differs 
from that of the other members of the family mainly in the long 
canal for the posterior nares not being closed by bone below, as 
the greater part of the palatines and the pterygoids do not meet in 
the middle line. The mandible has a prominent, narrow, recurved 
coronoid, and a well-developed angular process ; it is strongly de- 
curved in front. Vertebre: C7, D16, L2, 84, C40. Ribs 
remarkably broad and flat. Clavicles well developed. Manus 
remarkably modified, the third digit being greatly developed at the 
expense of all the others, and having a stout short metacarpal and 
but two phalanges, of which the most distal is large, compressed, 
pointed, and much curved, and bears a very strong hook-like claw. 
The second digit has the same number of phalanges, and bears a 
claw, but is very much more slender than the third. The fourth 
is represented only by the metacarpal and one nailless phalanx, 
the first and fifth only by very rudimentary metacarpals. The pes 


1 Gray, Annals of Philosophy, new series, vol. x. p. 343 (1825). 
13 


194 EDENTATA 


is also completely modified into a climbing organ. The hallux is 
rudimentary, consisting of a metatarsal and one phalanx, concealed 
beneath the skin; but the other four toes are subequal and much 
curved, with long pointed compressed claws. The tuber calcanei is 
directed towards the plantar surface, and parallel with it and 
extending to about double its length is a greatly elongated sesamoid 
ossicle. These together support a prominent calcarine cushion, to 
which the nails are opposed in climbing. Stomach pyriform, with 
muscular walls, but no distinct gizzard-like portion, as in the 

oP foregoing genera. Commence- 
ment of the colon provided with 
two small ceca (Fig. 66), resem- 
bling those of many birds, narrow 
at the base, and rather dilated 
at their terminal blind ends, and 
communicating with the general 
cavity by very minute apertures. 
Tail longer than the body, taper- 
ing, bare on the under surface, 


and very prehensile. Fur soft 
Fic. 66.—Ceca of the Two-toed Anteater and silky. 


paces didactylus). i, Tleum ; ¢, colon. This genus has also but one 

species certainly known, the Little or Two-toed Anteater (C\ di- 
dactylus), an animal not larger than a Rat, of a general yellowish- 
colour, and exclusively arboreal in its habits. It is a native of 
the hottest parts of South and Central America. 


Family DASYPODID. 


The greater part of the skin strongly ossified. On the back 
and sides the union of numerous quadrate or polygonal scutes forms 
a hard shield, usually consisting of an anterior (scapular) and 
posterior (pelvic) solid portion (which overhang on each side the 
parts of the body they respectively cover, forming chambers into 
which the limbs are withdrawn), and a variable number of rings 
between, connected by soft flexible skin so as to allow of curvature 
of the body. The top of the head has also a similar shield 
(cephalic), and the tail is usually encased in bony rings or plates. 
The outer or exposed surfaces of the limbs are protected by irregular 
bony scutes, not united at their margins; but the skin of the a 
surface of the limbs and under side of the body is soft, and more or 
less clothed with hair. Hairs also in many species project through 
apertures between the bony scutes of the back. The aagitied 
dermal scutes are everywhere covered by a layer of horny epi- 
dermis. Teeth numerous, simple, of persistent growth, and usually 


DASVPODIDE 198 


monophyodont, but in one genus (Tatusia) a succession of teeth has 
been observed. Zygomatic arch of skull complete. Cervical vertebrae 
with extremely short, broad, and depressed bodies. The atlas free, 
but the second and third, and often several of the others, anky- 
losed together both by their bodies and arches. Lumbar vertebre 
with accessory zygomatic processes, and very large metapophyses, 
supporting the bony carapace. Clavicles well developed. A third 
trochanter on the femur. Tibia and fibula ankylosed at their distal 
extremities. Fore feet with strongly developed, curved claws, 
adapted for digging and scratching—three, four, or five in number. 
Hind feet plantigrade, with five toes, all provided with nails. 
Tongue long, pointed, and extensile, though to a less degree than 
in the Anteaters. Submaxillary glands largely developed. Stomach 
simple. Uterus simple. Placenta discoidal, deciduate. The brain 
is generally characterised by the large size of the olfactory lobes 
(Fig. 57), and the slight development of sulci on the hemi- 
spheres ; the sylvian fissure being represented only by a very open 
and shallow angle. From the earliest stage of development the 
stapes is stirrup-shaped, thus showing a nearer affinity to the higher 
mammals than is presented by the Sloths. 

The animals of this family are commonly called Armadillos, 
a word of Spanish origin, having reference to their armour-like 
covering. The existing species are all of small or moderate size. 
They are mostly, though not universally, nocturnal in their 
habits, and are all omnivorous, feeding on roots, insects, worms, 
reptiles, and carrion. Armadillos are harmless and inoffensive 
creatures, offering no resistance when caught, their principal means of 
escape from their enemies being the extraordinary rapidity with which 
they can burrow in the ground, and the tenacity with which they re- 
tain their hold in their subterranean retreats. Notwithstanding the 
shortness of their limbs they can run with great rapidity. Most of 
the species are esteemed good eating by the natives of the countries 
in which they live. They are all inhabitants of the open plains or 
the forests of the tropical and temperate parts of South America, 
with the exception of one species (Tatusia novem-cincta), which 
ranges as far north as Texas. Of the existing genera, Chlamy- 
dophorus stands apart from the rest in the formation of its external 
covering ; but in all other respects Tutusta is the most aberrant 
form, exhibiting a peculiar type of structure of the fore feet, which 
in all the others show modifications, though in very varying degrees, 
of a single and different type. 

The reproductive organs of the Dasypodide differ from those of 
the Sloths and Armadillos in the presence of a largely developed 
copulating organ in the male, and of a simple vagina of correspond- 
ing length in the female. The testes are still abdominal, although 
not in the same position; and the penis still wants both the glans 


196 EDENTATA 


and bulb. The uterus is nearly or quite as simple as in the Sloths 
and Anteaters; and there is no reason to believe that the placenta- 
tion is essentially different from that obtaining in the other groups. 

Subfamily Chlamydophorinz.—In most anatomical characters, 
especially the structure of the fore foot, this little group resembles 
the Dasypodine ; but it differs remarkably from all other known 
Armadillos, living or extinct, in the peculiar modification of the 
dermal armour. 


Chlamydophorus.A—Teeth ae 


pressed, moderate in size, smaller at each end (especially in front) 
than at the middle of the series. Skull broad and rounded behind, 
pointed in front. Muzzle subcylindrical and depressed. A con- 
spicuous rounded, rough prominence on the frontal bone, just before 
each orbit. Tympanic prolonged into a tubular auditory meatus, 
curving upwards round the base of the zygoma. Vertebre: C7, 
D11,L3,810,C15. Upper part of head and trunk covered with 
four-sided horny plates (with very small thin ossifications beneath), 
forming a shield, free, and overhanging the sides of the trunk, and 
attached only along the middle line of the back. The plates are 
arranged in a series of distinct transverse bands, about twenty in 
number between the occiput and the posterior truncated end, and 
not divided into solid thoracic and pelvic shields with movable 
bands between. The hinder end of the body is abruptly truncated 
and covered by a vertically-placed, strong, solid, bony shield, of an 
oval (transversely extended) form, covered by thin epidermic plates. 
This shield is firmly ankylosed by five bony processes to the hinder 
part of the pelvis. Through a notch in the middle of its lower 
border the tail passes out. The latter is rather short, cylindrical 
in its proximal half, and expanded and depressed or spatulate in 
its terminal portion, and covered with horny plates. The dorsal 
surfaces of the fore and hind feet are also covered with horny 
plates. The remainder of the limbs and under surface and sides 
of the body beneath the overlapping lateral parts of the dorsal 
shield are clothed with rather long, very soft, silky hair, Eres and 
ears very small, and concealed by the hair. Extremities short. 
Feet large, each with five well-developed claws, those on the fore 
feet very long, stout, and subcompressed, the structure of the digits 
being essentially the same as those of Xenurus and Priodun. Nipples 
two, pectoral. Visceral anatomy closely resembling that of Dasypus, 
the cecum being broad, short, and bifid. 

The Pichiciago (C. truncatus), a small burrowing animal, about 
5 inches long, inhabits the sandy plains of the western part of the 
Argentine Republic, especially the Vicinity of Mendoza. — Its 


subcylindrical, somewhat com- 


) Harlan, Ann. New York Lyceum Nat. Hist. vol. i. p. 237 (1824),— 
Amended from Chiamyphorus. 


DASYVPODIDAZ 197 


horny covering is of a pinkish colour, and its silky hair snow 
white. It is rare, and its habits are but little known. A second 
species, C. retusa, from Bolivia, has been described by Burmeister. 
It is of rather larger size, and has the dorsal shield attached: to the 
skin of the back as far as its edge, instead of only along the median 
line. 

Subfamily Dasypodinz.—Fore feet usually with all five digits 
developed and with nails, though the first and fifth may be 
suppressed. The first and second long and slender, with the 
normal number and relative length of phalanges. The others stout, 
with short broad metacarpals, and the phalanges greatly reduced 
in length and generally in number by coalescence. The ungual 
phalanx of the third very large, that of the others gradually 
diminishing to the fifth. Dasypus, as now restricted, has the 
most normal form of manus, but the modifications so markedly 
developed in all the others (and culminating in Tolypeutes) are fore- 
shadowed, as it were, in it. Ears wide apart. Mamme one pair, 
pectoral. 

Dasypus..—Teeth 7% or #, of which the anterior in the upper 
jaw is usually implanted in the premaxillary bone. The series of 
teeth extends posteriorly some distance behind the anterior root of 
the zygoma, almost level with the hinder edge of the palate. They 
are large, subcylindrical, slightly compressed, diminishing in size 
towards each end of the series; the anterior two in the mandible 
much smaller, and more compressed than the others. Cranial 
portion of the skull broad and depressed. Facial portion triangular, 
broad in front and much depressed. Auditory bulla completely 
ossified, perforated on the inner side by the carotid canal, and 
continued externally into an elongated bony meatus auditorius, with 
its aperture directed upwards and backwards. (In all the remain- 
ing genera of Dasypodine the tympanic bone is a mere half ring, 
loosely attached to the cranium.) Mandible with a high ascending 
ramus, broad transversely-placed condyle, and high slender coronoid 
process. Vertebre: C7, D11-12,L3,88, 017-19. Head broad 
and flat above. Muzzle obtusely pointed. Ears of moderate size or 
rather small, placed laterally, far apart. Body broad and depressed. 
Carapace with six or seven movable bands between the scapular 
and pelvic shields, each plate, or scute, being marked by a regular 
ellipse formed of widely separated punctures. Tail shorter than 
the body, tapering, covered with plates forming distinct rings near 
the base. Fore feet with five toes; the first much more slender 
than the others, and with a smaller ungual phalanx and nail; the 
second, though the longest, also slender. The third, fourth, and 
fifth gradually diminishing in length, all armed with very strong, 
slightly curved, compressed claws, sloping away from an elevated 


1 Linn. Syst. Nat., 12th ed. vol. i. p. 54 (1766). 


198 EDENTATA 


rounded inner border to a sharp, outer, and inferior edge. The 
hind foot rather short, with all five toes armed with stout, 
compressed, slightly curved, obtusely pointed claws—the third the 
longest, the second nearly equal to it, the fourth the next, the first 
and fifth shorter, and nearly equal. 

To this genus belongs one of the best-known species of the 
group, the Six-banded Armadillo or Encoubert (J. sexcinctus) of 
Brazil and Paraguay. A very similar species, D. villosus, the Hairy 
Armadillo, replaces it south of the Rio Plata. There are also two 
very small species—D. vellerosus, from the Argentine Republic and 
North Patagonia, and D. minutus from La Plata. The latter differs 
from the other three in having no tooth implanted in the pre- 
maxillary bone. Remains apparently referable to D. villosus occur 
in the Pleistocene cavern-deposits of Brazil. 

Xenwrus.—Teeth $% or 8, of moderate size and subcylindrical. 
The most posterior placed a little way behind the anterior root of the 
zygoma, but far from the hinder margin of the palate. Cranium 
somewhat elongated, much constricted behind the orbits, and 
immediately in front of the constriction considerably dilated. 
Mandible slender ; coronoid process very small and sharp-pointed, 
sometimes obsolete. Vertebre: C7, D12-13, L3, 810, C18. 
Head broad behind. Ears rather large and rounded, wide apart. 
Movable bands of carapace 12-13; the scutes being marked by an 
obscurely granular sculpture. Tail considerably shorter than the 
body, slender, and covered with nearly naked skin, with but a few 
small, scattered, dermal bony plates, chiefly on the under surface 
and near the apex. On the fore feet the first and second toes are 
long and slender, with small claws and the normal number of 
phalanges ; the other toes have but two phalanges; the third has 
an immense falcate claw ; the fourth and fifth similar but smaller 
claws. The hind feet are comparatively small, with five toes, bearing 
small, triangular, blunt nails; the third longest, the first shortest. 
The best known species of this genus, the Tatouay or Cabassou, _V. 
unicinetus, is, after Priodon gigas, the largest of the group. It is 
found, though not abundantly, in Surinam, Brazil, and Paraguay, 
its remains occurring in the Pleistocene cavern-deposits of Brazil. 
Others, X. hispidus and lugubris, have been described, but little is as 
yet known of them. 

Priodon.2—Teeth variable in number, and generally differing on 
the two sides of each jaw, usually from 20 to 25 on each side 
above and below, so that as many as 100 may be present alto- 
gether ; but as life advances the anterior teeth fall out, and all 
traces of their alveoli disappear. The series extends as far back as 
the hinder edge of the anterior root of the zygoma. The teeth are 

1 Wagler, Syst. Amphibien, etc., p. 36 (1830). 
° FP. Cuvier, Hist. Nat. des Manvmiferes (1822).—Priodontes. 


DASVPODIDZ 199 


all very small ; those in the anterior half of each series being strongly 
compressed, with flat sides and a straight free edge ; the posterior 
ones are more nearly cylindrical, with flat truncated, free surfaces. 
Vertebre: C7, D12, L3, 810, C23. Head small, elongated, 
conical. Ears moderate, ovate. Carapace with 12-13 movable 
bands. Tail nearly equal to the body in length, gradually tapering, 
closely covered with quadrangular scales, arranged in a quincunx 
pattern. Fore feet with five toes, formed on the same plan as those 
of Xenurus, but with the claw of the third of still greater size, and 
that of each of the others, especially the fifth, proportionately reduced. 
Hind foot short and rounded, with five very short toes, with short, 
broad, flat, obtuse nails. The only known species, the Great 
Armadillo (P. gigas), is by far the largest of existing members of the 
family, measuring rather more than 3 feet from the tip of the nose 
to the root of the tail, the tail being about 20 inches long. It 
inhabits the forests of Surinam and Brazil. The powerful falcate 
claws of its fore feet enable it to dig with great facility. Its food 
consists chiefly of termites and other insects, but it is said to attack 
and uproot newly-made graves for the purpose of devouring the 
flesh of the bodies contained in them. 

Tolypeutes..—Teeth % or 8, rather large in proportion to the size 
of the skull, the hinder end of the series reaching nearly to the 
posterior margin of the palate. Vertebre: C7, D11, L3, 8 12, 
C13. Ears placed low on the sides of the head, rather large, 
broadly ovate. Carapace with its scapular and pelvic shields very 
free at the sides of the body, forming large chambers into which the 
limbs can be readily withdrawn. Only three movable bands ; 
sculpture of scutes in the form of subconcentrically arranged 
granules. Tail short, conical, covered with large bony tubercles. 
The fore feet formed on the same type as in the last genus, but the 
peculiarities carried out to a still greater extent. The claw of the 
third toe is very long and falcate, the first and fifth greatly reduced 
and sometimes wanting. On the hind foot the three middle toes 
have broad, flat, subequal nails, forming together a kind of tripartite 
hoof; the first and fifth much shorter, with more compressed 
nails. 

The Armadillos of this genus have the power of rolling them- 
selves up into a perfect ball, the shield on the top of the head and 
the tuberculated dorsal surface of the tail exactly fitting into and 
filling up the apertures left by the notches at either end of the 
carapace. This appears to be their usual means of defence when 
frightened or surprised, as they do not burrow like the other 
species. They run very quickly, with a very peculiar gait, only 
the tips of the claws of the fore feet touching the ground. Three 
species are described:—T. tricinctus, the Apar; T. conurus, the 

1 Mlliger, Prodromus Syst. Manum. et Aviwm, p. 111 (1811). 


200 EDENTATA 


Matico ; and 7. mwrici. Remains apparently referable to 7. conurus 
are of not uncommon occurrence in the Brazilian cavern-deposits. 
Subfamily Tatusiine.—This group contains but one genus, 
Tatusia.1 Teeth £ or 4, very small subcylindrical. The first and 
second subcompressed, the last considerably smaller than the others. 
They present the remarkable peculiarity (elsewhere found among 
Edentates, so far as is yet known, only in Orycteropus) of all being, 
with the exception of the last, preceded by two-rooted milk teeth, 
which are not changed until the animal has nearly attained its full 
size. Vertebree: C7, D 9-11, L5, 88, C 20-27. Head narrow, 
with a long, narrow, subeylindrical, obliquely-truncated snout; 
pterygoids meeting in the middle line below the nasal passage. Ears 
rather large, ovate, and erect, placed close together on the occiput. 


Fic. 67.—The Peba Armadillo (Tutusia novemcincta). 


Carapace with seven to nine distinct movable bands : sculpture on 
scutes consisting of pits arranged in a V-shape. Body generally 
elongated and narrow. Tail moderate or long, gradually tapering ; 
its dermal scutes forming very distinct rings for the greater part ‘ot 
its length. Fore feet with four visible toes, and a concealed clawless 
rudiment of the fifth. Claws all long, slightly curved, and very 
slender, the third and fourth subequal and alike, the first and fourth 
much shorter. Hind feet with five toes, all armed with strong 
slightly curved, conical, obtuscly-pointed nails. The third longest, 
then the second and fourth; the first and fifth much shorter than 
the others. 

This genus differs from all the other Armadillos in having a pair 
of inguinal mamme, in addition to the usual pectoral pair, and in 


1 Lesson, Man. de Mammatoyir, p. 309 (1827); ex. F. Cuvier, Tatusic 


DASYPODID4 201 


producing a large number (four to ten) of young at a birth, all the 
others having usually but one or two. 

The Peba Armadillo, 7. novemcincta (Fig. 67), is a well-known 
species, having an extensive range from Texas to Paraguay. It is 
replaced in the more southern regions of South America by a smaller 
species, with shorter tail, the Mulita (T. hybrida), so called from the 
resemblance of its head and ears to those of a mule. T. kappleri is 
a large species from Surinam. 

A rare Armadillo from Peru described under the names of Crypto- 
phractus pilosus and Praopus hirsutus, but which evidently belongs to 
Tatusia, is of some interest owing to the thick coat of hair with 
which it is covered. This animal appears to be closely allied to 
LT. novemeineta, from which it mainly differs by having the whole of 
the carapace covered with a thick coating of light brown, fine, but 
rather stiff hair, about an inch and a half in length. Similar hair 
is found on the cheeks, the proximal portions of the limbs, and 
(although less abundantly and shorter) on the under surface of the 
body. The cephalic shield, snout, feet, and the tail, with the 
exception of the root, are bare. The coating of hair on the back 
and sides completely conceals the carapace, except near the margin 
of the scapular region ; but by separating the hairs the bands and 
scutes are rendered visible.t 

In the Pleistocene cavern-deposits of Brazil have been found 
remains of 7. novemcincta, and also of T. punctata, which appears to 
be an extinct form nearly allied to 7. kappleri, but of somewhat 
larger size. 

Extinct genera.—In addition to remains referable to existing 
genera, the above-mentioned deposits have also yielded evidence 
of the former existence of extinct generic types of Armadillos, 
some of which attained very large dimensions. Of these Eutatus 
was a large form distinguished from all existing genera by the 
circumstance that the whole of the carapace was composed of mov- 
able bands, which were thirty-three in number. Dasypotherium 
was a still larger form, furnished with eight teeth, of which the 
second seems to have been larger than the others; this genus is 
regarded as connecting the modern Armadillos with the next one. 
The gigantic Chlamydotherium, the scutes of which are common in 
the Brazilian caves, is considered to have been as large as a 
Rhinoceros ; the carapace has several movable bands, but the teeth 


1 A single imperfect skin, brought from the province of Ceara in Brazil, indi- 
cates a very remarkable form of Armadillo, named by A. Milne-Edwards Sclero- 
pleura brunetti (Ann. Se. Nat. xvi. p. 8, 1872). The dermal scutes are said to 
be much less developed than in other members of the family, and confined to the 
sides, all the median portion of the back being clothed with a flexible hairy skin. 
The head is broad and short, the ears small and far apart. The tail is long, and 
almost entirely devoid of scutes. The feet are unknown. 


202 EDENTATA 


approximate in structure to those of the next family, so that the 
genus tends to connect the Armadillos with the Glyptodonts. 


Family GLYPTODONTID. 


In the Pleistocene cavern-deposits of Brazil, but still more 
abundantly in the fluviatile deposits which cover the country in the 
neighbourhood of Buenos Ayres, are found the remains of some of the 
most remarkable forms of mammals yet discovered, the Glyptodonts, 
which may be regarded as forming a separate 
extinct family. They differ from the existing 
Dasypodide in their large size, and in having the 
carapace composed of a solid piece (formed by 
the union of a multitude of bony dermal scutes) 
without any movable rings, and in usually hav- 
ing also a ventral piece or plastron. The facial 
portion of the skull is very short. <A long 
process of the maxillary bone descends from 
the anterior part of the zygomatic arch. The 
ascending ramus of the mandible is remarkably 
high. The teeth are 8 in the known species, 
all much alike, having two deep grooves or 
flutings on each side, so as to divide them into 
three nearly distinct lobes (Fig. 68). The verte- 
bral column is almost entirely ankylosed into 
a solid tube, and there is a complex joint at the 
base of the neck, to allow of the head being 
retracted within the carapace. The limbs are 
very strong, and the feet short and _ broad, 
resembling externally those of an elephant or 
tortoise. This family is mainly characteristic 
of the southern half of the American continent, 
but some species of the type genus ranged into 

Fic. 68.—Tooth of Glyp, Texas and Mexico. Many species of the family 
todon from the side, and have been described and figured, especially by 
aoe surface. Burmeister Gn the fnnles del Museo publica de 

Buenos Aires), among which the following may 
be noticed. Hoplophorus is characterised by the sculptured and 
frequently thin scutes of the carapace, those of the periphery being 
flat, and not raised into prominences. The caudal sheath has 
several overlapping movable rings at the base, and ends in a lone 
subcylindrical terminal tube similar to the one represented with the 
carapace of Gilyptodon in Fig. 69, which in all probability really belongs 
to the genus under consideration. Each foot has four complete 
digits, and the humerus has an entepicondylar foramen. Most of the 


GLYPTODONTIDE 203 


species are of medium size. Part of a caudal tube from Uruguay 
described as Eleutherocercus indicates, however, a much larger allied 
form, in which the tail appears to have had a number of stout bristles 
protruding from the joints between the scutes. Panochthus com- 
prises very large Glyptodonts, distinguished by the great thickness 
of the scutes of the carapace, which are ornamented with tubercles. 
The termination of the caudal sheath forms a tube bearing large 
radiated tubercles. Euryurus is distinguished by the radiate 
sculpture of the scutes of the carapace. Dedicurus, of which one 
species was about twelve feet in length, also has a rugose 
sculpture on the carapace ; but the termination of the caudal tube is 
expanded into a club-like shape, flattened from above downwards, 


ISCO OS, 


Fic. 69.—Glyptodon clavipes (Pleistocene, South America). From Owen. The tail is incorrectly 
restored, and it is probable that the figured portion belongs to Hoplophorus. The left lower 
corner shows an upper and a lower view of the skull, and the right a section of the caudal 
sheath. 


and covered with tubercles mingled with a few large radiate discs, 
which, as in Panoehthus, probably carried horny spines in the living 
condition. The typical genus Glyptwlon has each scute of the 
carapace ornamented with a rosette-like sculpture, the peripheral 
scutes being raised into conical prominences (Fig. 69). The caudal 
sheath, instead of being like the one represented in the figure, was 
entirely composed of a series of movable rings, ornamented with 
large tubercles. The manus had five digits, and the pes four; and 
there was an entepicondylar foramen to the humerus. A species of 
this genus, which attained very large dimensions, was made the 
type of Schistopleurum, on the supposition that the tail of Glyptodon 
was of the type represented in Fig. 69. The genus Thvracophorus, 


204 _ EDENTATA 


of the Pleistocene of South America, as well as Carioderma, of the 
Pliocene of Texas, differ from all the preceding in having the scutes 
of the carapace in the form of disconnected nodules. Glyptodonts 
also occur in South American beds of earlier age than the Pleistocene, 
some of these forms having enamel bands on the teeth. ‘“ Why such 
a form as the Glyptodon should have failed to keep his ground is,” 
as the late Professor W. K. Parker remarks, “a great mystery ; 
nature seems to have built him, as Rome was built, for eternity.” 


Family MANID#&. 


Covered externally (except the under surface of the body and 
inside of the limbs) with large imbricated horny scales, and 
scattered hairs growing in the intervals. No teeth. Tongue long, 
vermiform, and protractile. No accessory articular processes to 
the lumbar vertebre, but the anterior zygapophyses largely de- 
veloped and deeply concave, completely embracing the semicylindri- 
cal surfaces of the posterior zygapophyses. Limbs short, with five 
complete digits on each foot. Scaphoid and lunar bones of carpus 
united. Uterus bicornuate. Placenta diffused and non-deciduate. 
All the existing forms belong to the Ethiopian and Oriental regions 
of the Old World. The absence of additional articular processes to 
the lumbar vertebree is a character in which this and the following 
family differ from all the preceding forms. 

Manis..—Skull somewhat of the form of an elongated cone, with 
the smal] end turned forwards ; very smooth and free from crests 
and ridges. No distinction between the orbits and temporal fosse. 
The zygomatic arch usually incomplete, owing to the absence of 
the jugal bone. No distinct lachrymal bone. Palate long and 
narrow. The pterygoids extend backwards as far as the tympanics, 
but do not meet in the middle line below. Tympanic ankylosed to 
the surrounding bones, and more or less bullate, but not produced 
into a tubular auditory meatus. Rami of mandible very slender 
and straight, without any angle or coronoid process. From near 
the anterior extremity of the upper edge a sharp, conical, tooth-like 
process projects upwards and outwards. No clavicles. No third 
trochanter to the femur. Ungual phalanges bifid at their ter- 
minations. Caudal vertebre with very long, strong transverse 
processes and numerous chevron bones. Tongue long, vermiform, 
flattened towards the tip; its retractor or sterno-glossal muscles 
arising from the hinder extremity of the immensely prolonged 
ensiform cartilage of the sternum. Stomach with thick lining 
membrane and muscular walls, and a special gland near the 
middle of the great curvature, consisting of a mass of complex 


1 Linn. Syst. Nat. 12th ed. vol. i. p. 52 (1766). 


MANIDA 205 


secreting follicles, the ducts of which terminate in a common 
orifice. No cecum. A gall-bladder. Head small, depressed, 
narrow, pointed in front, with a very small mouth-opening. 
Eyes and pinna of ear very small. Body elongated, narrow. 
Tail more or less elongated, convex above, flat underneath. The 
whole of the upper surface of the head, the upper surface and sides 
of the body, the whole of the tail, and the outer sides of the ex- 
tremities covered with large, overlapping, horny scales, but usually 
with a few stiff hairs growing between and projecting beyond 
them. The sides and under surface of the head, the under surface 
of the body, and the inner sides of the limbs without scales, but with 
a rather scanty covering of hair. Limbs short. In walking the 
dorsal surface and outer sides of the phalanges of the two outer 
digits of the front feet alone rest on the ground, the points of the 
nails turning upwards and inwards. The third toe the longest, 
with a powerful compressed curved claw; the second and fourth 
with similar but smaller claws, that of the pollex often almost 
rudimentary. Hind feet plantigrade, with the hallux very short, 
and the four other toes subequal, with moderate, curved, subcom- 
pressed nails. 

The reproductive organs of Munis are of a totally different 
type from those of the families already noticed. The testes lie 
in the inguinal canal; and the penis is external and well developed. 
The uterus is truly bicornuate, the vagina not divided, and the 
placenta diffused and non-deciduate. All the organs and fetal 
membranes are, indeed, formed very much on the plan of those 
of the Ungulates, without any trace of the special peculiarities 
obtaining in the typical American Edentates. 

The animals of this genus, which includes all the existing forms, 
are called Pangolins or Scaly Anteaters, and are all of small or 
moderate size, terrestial and burrowing, and feed mainly on termites. 
Several of them can climb trees. Their length varies from 1 to 5 
feet. They can roll themselves up in a ball when in danger. Their 
peculiar elongated form, short limbs, long, gradually-tapering tail, 
and scaly covering give them on a superficial inspection more the 
appearance of reptiles than of mammals. The species are not 
numerous, and may be divided into two groups distinguished by a 
few not very important external characters ; these groups also coin- 
ciding with the present geographical distribution of the genus. 
These two groups, according to Mr. O. Thomas, may be distinguished 
as follows. 

The Asiatic pangolins are characterised by having the central 
series of body-scales continued quite to the extreme end of the tail, 
by having many isolated hairs growing up between the scales of the 
back, and by their small external ears. They all have a small 
naked spot beneath the tip of the tail, which is said to be of service 


206 EDENTATA 


as an organ of touch. There are three species, viz. Alunis javanica, 
ranging from Burma, through Malacca and Java, to Borneo; Jf. 
auritu, found in China, Formosa, and Nipal; and the common Indian 
Pangolin, WZ. pentadactyla, distributed over the whole of India and 
Ceylon. The African species have the central series of scales 
suddenly interrupted and breaking into two at a point about 2 or 3 
inches from the tip of the tail; they have no hair between the 
scales, and no external ear-conch. The following are the four species 
belonging to this 
group: — the 
Long-tailed Pan- 
golin (J. mae- 
rura), which has 
a tail nearly twice 
as long as_ its 
body, and con- 
taining as many 
as forty-nine 
caudal vertebra, 
being the largest 
number known 
among mammals ; 
the White-bellied 
Pangolin (MZ. ft7- 
cuspis), Fig. 70, 


hats Ae Nh closely allied to 
oes z SS , ‘3 
ASG = 4 the last, but with 
oan € OS longer and _ tri- 


cuspid scales, and 
white belly hairs. 
These tio, like 
the Indian species, have a naked spot beneath the tail tip, a char- 
acter probably correlated with the power of climbing, and they 
are, moreover, peculiar in having the outer sides of their fore legs 
clothed with hair, all the other species being scaly there as else- 
where. Lastly, the Short-tailed and the Giant Pangolins (1. 
femmincki and gigantea), both of which have their tails covered 
entirely with scales, and evidently never take to arboreal habits. 
All the four species of the second group are found in the West 
African region, one only, AL temmincki, extending also into south 
and eastern equatorial Africa. 

According to Professor W. K. Parker,! who remarks upon the 
peculiarly aberrant nature of the group, the horny seales of the 
Pangolins really consist of cemented hairs. This writer states that 
“in the early embryo lozenge-shaped tracts of skin are seen all over 

* Mammalian Descent, p. 95, 


Fic. 70.—The White-bellied Pangolin (Manis tricuspis). 


MANIDA 207 


its body, with lines of thinner cuticle between. Under the micro- 
scope, sections of these thicker tracts show that they are composed 
of fine hairs, cemented together by a copious growth of epidermic 
cells; here and there larger hairs are seen, but these fail to reach 
the surface, turning again towards the inside, like nails driven into 
wood that is too hard for their points.” 

The same author also observes! that there are occasional in- 
stances of the presence of eight cervical vertebre in the Pangolins 
—a feature which has been considered to indicate some former 
genetic connection between this family and the Sloths.. 

The following account of the habits of Manis tricuspis is given by 
Mr. L. Fraser in his Zoologia Typica -— 

“During my short residence at Fernando Po I succeeded in 
procuring two living specimens of this animal. The individuals, 
judging from the bones, were evidently not adult; the largest 
measured 30 inches in length, of which the head and body were 
12 inches and the tail 18 inches. I kept them alive for about a 
week at Fernando Po, and allowed them the range of a room, where 
they fed upon a small black ant, which is very abundant and trouble- 
some in the houses and elsewhere. Even when first procured they 
displayed little or no fear, but continued to climb about the room 
without noticing my occasional entrance. They would climb up 
the somewhat roughly-hewn square posts which supported the 
building with great facility, and upon reaching the ceiling would 
return head foremost; sometimes they would roll themselves up 
into a ball and throw themselves down, and apparently without 
experiencing any inconvenience from the fall, which was in a 
measure broken upon reaching the ground by the semi-yielding 
scales, which were thrown into an erect position by the curve of 
the body of the animal. In climbing, the tail, with its strongly 
pointed scales beneath, was used to assist the feet; and the grasp 
of the hind feet, assisted by the tail, was so powerful that the 
animal would throw the body back (when on the post) into a 
horizontal position, and sway itself to and fro, apparently taking 
pleasure in this kind of exercise. It always slept with the body 
rolled up; and when in this position in a corner of the building, 
owing to the position and strength of the scales, and the power of 
the limbs combined, I found it impossible to remove the animal 
against its will, the points of the scales being inserted into every 
little notch and hollow of the surrounding objects. The eyes are 
very dark hazel, and very prominent. The colonial name for this 
species of Afanis is ‘Attadillo,” and it is called by the Boobies, 
the natives of the island, ‘Gahlah.’ The flesh is said to be 
excedingly good eating, and is-in great request among the 
natives.” 

1 Mammalian Descent, p. 99. 


208 EDENTATA 


The Indian species is said to live in pairs, and to give birth to 
one or two young at a time in the spring. Their burrow reaches a 
depth of some twelve feet, and terminates in a large chamber, which 
may be as much as six feet in diameter. A faint hiss appears to be 
the only sound emitted by these animals. 

Remains of a large species of Manis, which are indistinguishable 
from the corresponding bones of the existing West African I. 
gigantea, are found fossil in cave-deposits in the Karnul district of 
Madras. This is one among several instances of the close connection 
between the Pleistocene and Pliocene mammalian fauna of India with 
the existing African fauna. 

Paleomanis.'—The lower Pliocene deposits of the Isle of 
Samos, in the Turkish Archipelago, have yielded remains of a 
Pangolin fully three times the dimensions of JL. gigantea, upon the 
evidence of which the genus Paleomanis has been established. 


Family ORYCTEROPODID.£ 


External surface scantily covered with bristle-like hairs. Teeth 
numerous, apparently heterodont, diphyodont, and of peculiar and 
complex structure, being traversed by a number of parallel vertical 
pulp-canals. Lumbar vertebra with no accessory zy gapophyses. 
Femur with a third trochanter. Fore feet without pollex, but all 
the other digits well developed, with strong moderate-sized nails, 
suited to digging, the plantar surfaces of which rest on the ground 
in walking. Hind feet with five subequal toes. Mouth elongated 
and tubular. Tongue subvermiform. Uterus bicornuate. Placenta 
broadly zonular. Feeding on animal substances. Terrestrial and 
fossorial in habits. Now mainly limited to the Ethiopian region. 

Orycteropus.’—The total number of permanent teeth appears to 
be from eight to ten in each side of the upper, and eight in the 
lower jaw; but they are never all in place at one time, as the 
small anterior teeth are shed before the series is completed behind. 
In the adult they number usually five on each side above and below, 
of which the first two are simple and compressed, the next two 
larger and longitudinally grooved at the sides, the most posterior 
simple and cylindrical. The last three in either jaw having no 
milk-predecessors, may be regarded as true molars. The structure 
of all these teeth is quite peculiar among mammals, though 
resembling that of some fishes. Their summits are rounded before 
they are worn; their bases do not taper to a root, but are evenly 
truncated and continually growing. Each tooth is made up of an 
aggregation of parallel dental systems, having a slender pulp-cavity 


1 Forsyth-Major, Comptes Rendus, vol. evii. p. 1180 (1888), 
* Geoffroy, Décade Philosophique, 1795 (teste Agassiz) 


ORYCTEROPODIDE 209 


in the centre, from which the dentinal tubes radiate outwards, and 

- being closely packed together each system assumes a polygonal 
outline as seen in transverse section. The small anterior teeth have 
milk-predecessors which are fully noticed below. Skull moderately 
elongated. The facial portion subcylindrical and slightly tapering. 
The zygoma complete and slender. The palate ends posteriorly in 
the thickened transverse border of the palatines, and is not 
continued back by the pterygoids. The tympanic is annular, and 
not ankylosed to the surrounding bones. The mandible is slender 
anteriorly, but rises high posteriorly, with a slender recurved 
coronoid, and an ascending pointed process on the hinder edge 
below the condyle, which is small, oval, and looks as much forwards 
as upwards. Vertebre: C7, D13, L8, 86, C27. The large 
number of lumbar vertebre is peculiar among Edentates. Tongue 
less vermiform than in Alyrimecophaga, being thick and fleshy at the 
base, and gradually tapering to the apex. The salivary apparatus 
is developed much in the same manner as in that genus, but the 
duct of the submaxillary gland has no reservoir. The stomach 
consists of a large subglobular cardiac portion, with a very thick, 
soft, and corrugated lining membrane, and a smaller muscular, 
pyloric part, with a comparatively thin and smooth lining. There 
is a very distinct ileo-cecal valve, and a considerable-sized cecum ; 
also a gall-bladder. Head elongated, with a tubular snout, terminal 
nostrils, and small mouth-opening. Ears large, pointed, erect. 
Tail nearly as long as the body, cylindrical, very thick at the base, 
tapering to the extremity. 

The reproductive organs and placentation of Orycteropus are 
formed upon a principle unknown in the more typical Edentates, 
or, in combination, in any other mammals. Thus the testes, in the 
one described example, were inguinal, but appeared to descend, at 
all events temporarily, into a scrotum; but the penis is scarcely 
larger than that of the Great Anteater. The uterus is still more 
fully bicornuate than in J/unis, with its two lateral chambers 
opening separately into the vagina, as in certain Rodents. The 
placenta is broadly zonary, but it is not known whether it is 
deciduate or not. It might readily be derived from the diffused 
placenta of A/unis by the abortion of the foetal villi at the two poles 
of the ovum. 

The Orycteropodide have long been regarded as widely different 
from other Edentates, their presumed affinity with the J/anide 
being more or less problematical ; but the discovery recently made 
by Mr. O. Thomas? that they have a milk-dentition still further 
emphasises their aberrant nature. According to this observer, it 
appears that there are normally no less than seven milk-teeth in the 
upper jaw, the hindmost of which is far larger than the others, 

1 Proceedings of the Royal Society, vol. xlvii. p. 246 (1890). 
14 


210 EDENTATA 


having a rudimentary crown, and a distinct anterior and posterior 
root. The other milk-teeth are styliform, the four anterior ones 
being very minute, and separated from one another by equal 
intervals; the foremost of all is situated immediately behind the 
premaxillo-maxillary suture. In the mandible only four milk-teeth 
have hitherto been detected, of which the hindmost has the 
comparatively complex form found in the corresponding upper tooth. 
None of these milk-teeth appear, however, to cut the gum, so that 
the whole set is entirely functionless. Under the microscope these 
milk-teeth show signs of possessing a commencement of the 
remarkable histological structure found in the permanent teeth. 

Mr. Thomas remarks that since “the three large posterior teeth 
of Orycteropus, already distinguished by their more molariform shape, 
do not have milk-predecessors, while all the small teeth anterior to 
them do, and in addition the last milk-tooth is markedly different 
from those in front of it, we ought apparently no longer to look 
upon this animal as an homodont, but instead to consider it as an 
originally heterodont form in which the incisors and canines have 
been suppressed to allow free play to the mobile vermiform tongue. 

“But important as a knowledge of the presence of a milk- 
dentition in Orycteropus is, it does not at present render any easier 
the difficult questions as to the phylogeny and systematic position 
of that animal. Although called an Edentate, it has always been 
recognised as possessing many characters exceedingly different from 
those of the typical American members of the order. It has in fact 
been placed with them rather on account of the inconvenience of 
forming a special order for its reception than because of its real 
relationship to them. Now, as they are either altogether toothless, 
or else homodont and monophyodont (apart from the remarkable 
exception of Tatusta), it seems more than ever incorrect to unite 
with them the solitary member of the Tubulidentata, toothed, 
heterodont, and diphyodont, and differing from them in addition by 
its placentation, the anatomy of its reproductive organs, the minute 
structure of its teeth, and the general characters of its skeleton. 

“But if Orycteropus is not genetically a near relation of the 
Edentates, we are wholly in the dark as to what other mammals it 
is allied to, and I think it would be premature to hazard a guess on 
the subject. Whether even it has any special connection with 
Manis is a point about which there is the greatest doubt, and unfor- 
tunately we are as yet absolutely without any paleontological 
knowledge of the extinct allies of either. Afacrotherium even, 
usually supposed from the structure of its phalangeal bones, to be 
related to Manis, has lately proved to have the teeth and vertebrx 
of a perissodactyle Ungulate, and one could not dare to suggest 
that anceStors of Aunis, or Orycteropus were to be sought in that 
direction. Lastly, as the numerous fossil American Edentates do 


ORYCTEROPODIDE 211 


not show the slightest tendency to an approximation towards the 
Old World forms, we are furnished with an additional reason for 
insisting on the radical distinctness of the latter, whose phylogeny 
must therefore for the present remain one of the many unsolved 
zoological problems.” 

The Aard-Varks (Earth-Pigs) as these creatures are commonly 
termed, from the name bestowed on them by the Dutch Boers of 
the Cape, are of nocturnal habits, sleeping during the day in their 
burrows, which are usually found in the neighbourhood of the tall 
hills or mounds made by termites. Indeed, wherever these hills are 
abundant it is stated there is a good chance of finding an Aard-Vark, 
the food of these animals consisting almost exclusively of termites 
and ants. 

Two existing species are recognised, namely the Cape Aard-Vark 
(0. afra) from South Africa, and another (0. ethiopicus) from the 
north-eastern parts of Africa, ranging into Egypt. An extinct 
species has been described from the Lower Pliocene of the Isle 
of Samos, in the Turkish Archipelago, differmg from the exist- 
ing forms by the larger proportionate size of the lateral meta- 
tarsals. 


Bibliography of Edentata.—No general work on the order has been published 
since that of Rapp (Anat. Untersuchungen iiber die Edentaten, 2d ed. 1852). 
Among numerous memoirs on special groups the following may be cited :— 
Myrmecophagide :—R. Owen, ‘‘ Anatomy of Great Anteater,” Trans. Zool. Soc. 
vol. iv.; G. Pouchet, MWém. sur le Grand Fourmilier, 1874; W. A. Forbes, 
‘‘Anat. of Great Anteater,” Proc. Zool. Soc. 1882, p. 287. Megatheriide :—R. 
Owen, Extinct Gigantic Sloth (Mylodon Robustus), 1842; Id., ‘‘On the Mega- 
therium,” Phil. Trans. 1851-56; J. Leidy, ‘‘Extinct Sloth-tribe of North 
America,” Smithsonian Contrib. to Knowledge, vii. 1855; H. Burmeister, 
Description de la République Argentine, t. iii. Mammiféres, 1879,—which contains 
full references to various memoirs by Owen, Gervais, Reinhardt, and others. 
Glyptodontide :—Owen, Catalogue of Fossil Mammals, Mus. Roy. Coll. Surgeons, 
1845 ; T. H. Huxley, ‘‘ Osteol. of Glyptodon,” Phil. Trans. 1865 ; H. Burmeister, 
Annales del Museo Publico de Buenos Aires, and Descript. de la République 
Argentine, 1879; H. Gervais and F. Ameghino, Les Mammiferes Fossiles de 
2 Amérique Méridionale, Paris, 1880,—which also contains a list of all the 
S. American Edentates described at that date. Dasypodide :—J. Murie, ‘‘Ana- 
tomy of Tolypeutes,” Trans. Linn. Soc. vol. xxx. 1874; A. H. Garrod, Proc. 
Zool. Soc. 1878. For Placentation of Edentates see W. Turner, Trans. Roy. Soc. 
Edin, xxvii. (1873) p. 72, and Journ. Anat. and Physiol. vols. viii. and x.; A. 
Milne-Edwards, Ann. Sciences Nat. [6] viii. p. 1; and for brain, P. Gervais, 
“«Formes cérébrales des Edentés,” Nowv. Arch. du Muséum, tom. v. ; W. Turner, 
Jour. Anatomy, i. 313 (1867). For the dentition of Orycteropus see O. Thomas, 
“A Milk Dentition in Orycteropus,” Proc. Roy. Soc. vol. xlvii. p. 246 (1890). 
Fuller observations on the mutual relations of the various families are given by 
W. H. Flower, “‘ On the Mutual Affinities of the Animals composing the Order 
Edentata,” Proc. Zool. Soc. 1882, p. 358. 


» CHAPTER VIE 
THE ORDERS SIRENIA AND CETACEA 


Order STRENIA. 


THE purely aquatic habits and fish-like form of the animals of this 
order caused them to be formerly confounded with the Cetacea, 
but a more intimate knowledge of their structure has shown that 
they really belong to a widely different type of the mammalian 
class. 

The head is rounded and not disproportionate in size as com- 
pared with the trunk, from which it is scarcely separated by any 
externally visible constriction or neck. Nostrils valvular, separate, 
and placed above the fore part of the obtuse truncated muzzle. 
Eyes very small, with imperfectly formed eyelids, capable, however, 
of contraction, and with a well-developed nictitating membrane. 
Ear without any pinna. Mouth of small or moderate size, with 
tumid lips beset with stiff bristles. General form of the body 
depressed, fusiform. No dorsal fin. Tail flattened and horizontally 
expanded. Fore limbs paddle-shaped, the digits being enveloped 
in a common cutaneous covering, on which rudiments of nails are 
sometimes present. No trace of hind limbs in existing forms. Ex- 
ternal surface covered with a tough, finely wrinkled, or verr 
rugose skin, naked, or with fine hairs sparsely scattered over it. _ 

The skeleton is remarkable for the massiveness and density of 
most of the bones of which it is composed, especially the skull and 
ribs, which must add to the specific gravity of these slow-moving 
animals, and aid in keeping them to the bottom of the shallow 
waters in which they dwell, while feeding on aquatic vegetables. 
The skull presents many peculiarities, among which may be indicated 
the large size and backward position of the anterior narial aperture, 
a further modification of that met with in the Tapirs among Ungu- 
lates, and presenting some approach to that so characteristic of the 
Cetacea. The nasal bones are generally absent in the recent forms, 


STRENTA 213 


or are only found in a most rudimentary condition, attached to the 
edge of the frontals, far away from the middle line; but in some at 
least of the extinct species these bones, though small in size, are 
normal in situation and relations. In very few other respects does the 
skull present any resemblance to that of the Cetacea. In the spinal 
column of existing forms none of the vertebrae are united together 
to form a sacrum, and the flat ends of the bodies do not ossify 
separately, so as to form disc-like epiphyses in the young state, as 
in nearly all other mammals; traces of epiphyses have, however, 
been recently detected in Afanatus, and they were fully developed in 
Halitherium and other fossil forms. The anterior caudal vertebre 
have well-developed chevron bones. In one genus (J/anatus) there 
are only six cervical vertebree. There are noclavicles. The humerus 
has a small but distinct trochlear articulation at the elbow-joint. 
The two bones of the forearm are about equally developed, and 
generally ankylosed together at both extremities. The carpus is 
short and broad, and the digits five in number, with moderately 
elongated and flattened phalanges, which are never increased in 
number beyond the limit usual in the Mammalia. The pelvis is 
extremely rudimentary, consisting of a pair of bones suspended at 
some distance from the vertebral column. In no existing species 
is there any trace of a hind limb, but in the extinct Halitherium 
an acetabular depression and rudimentary femur have been dis- 
covered. 

Two kinds of teeth, incisors and molars, separated by a wide 
interval, are generally present. The former may be developed into 
tusks in the upper jaw, or may be quite rudimentary. The molars 
vary much in character. In one genus (Lhytina) no teeth of any 
kind are present, at least in the adult. Some fossil forms show a 
more decidedly heterodont dentition, while Halitherium has milk- 
teeth, of which no traces have been observed in the recent genera. 
In all recent types the anterior part of the palate, and a corre- 
sponding surface on the prolonged symphysis of the lower jaw, are 
covered with rough horny plates of peculiar structure, which doubt- 
less assist in mastication. The tongue is small and fixed in position, 
with a surface resembling that of the plates just spoken of. The 
salivary glands are largely developed. The stomach is compound, 
being divided by a valvular constriction into two principal cavities, 
the first of which is provided with a singular glandular pouch near 
the cardiac end, and the second usually with a pair of elongated, 
conical, cecal sacs or diverticula. The intestinal canal is long, and 
has very muscular walls. There is a cecum, either simple, conical, 
and with extremely thick walls, as in Hualicore, or bifid, as in Aanatus. 
The heart is broad and flat, with its apex deeply cleft between the 
ventricles. The principal arteries form very extensive and complex 
retia mirabilia. The lungs are remarkably long and narrow, as, 


2145 SIRENTA 


owing to the very oblique position of the diaphragm, the thoracic 
cavity extends far back over the abdomen. The epiglottis and 
arytenoid cartilages of the larynx do not form a tubular prolong- 
ation as in the Cetacea, so that the epiglottis is not intranarial. 
The brain is of comparatively small size, and the convolutions on 
the surface of the cerebrum are few and shallow. The kidners are 
simple. The testes abdominal. The uterus is bicornuate. The 
placenta (in the Dugong) is non-deciduate and zonary. The um- 
bilical vesicle disappears early. The mamme are two, and pectoral, 
or rather post-axillary in position. 

The Sirenia pass their whole life in the water, being denizens of 
shallow bays, estuaries, lagoons, and large rivers, but, unlike the 
Cetacea, are not met with in the high seas, far away from the shore. 
Their food consists entirely of aquatic plants, either marine alze or 
freshwater grasses, upon which they browse beneath the surface, as 
the terrestrial herbivorous mammals do upon the green pastures on 
shore. They are generally gregarious, slow and inactive in their 
movements, mild, inoffensive, and apparently unintelligent in dis- 
position. Though occasionally found stranded by the tide or waves, 
there is no satisfactory evidence that they voluntarily leave the water 
to bask or feed on the shore. The habit of the Dugong of raising 
its round head out of the water, and carrying its young under the 
fore fin, seems to have given rise, among the imaginative early 
voyagers in the Indian Ocean, to the legendary beings, half human 
and half fish, in allusion to which the name Sirenia was bestowed by 
Illiger on the order, though certainly the face of a Dugong, when 
closely inspected, does not bear the slightest resemblance to that of 
the mermaid of romance. The specie: now existing are very few, 
and there is reason to believe that the time is not far distant when 
they will all become extinct. One species, Rhytina st-lleri, of the 
North Pacific, was totally exterminated through the azency of man 
during the last century; and the others, being valuable for their 
flesh as food, for their hides, and especially for the oil obtained from 
the thick layer of fat which lies immediately beneath their skin, 
rapidly diminish in numbers as civilised populations occupy the 
regions forming their natural habitat. The surviving species are 
confined to the tropical regions of the shores of both sides of the 
Atlantic and the great rivers which empty themselves into that 
ocean, and to the coasts of the Indian Ocean from the Red Sea to 
North Australia. In the Miocene and early Pliocene epoch 
Sirenians abounded in the seas of Europe, and their remains 
have been found in deposits of corresponding periods in North 
America. Evidence has also been discovered of the existence 
of an animal of this group in the seas at the bottom of which 
the Eocene nummulitic limestone mountain ranges of Egypt were 
deposited. * 


MANATIDA 215 


The existing genera present such well-marked distinguishing 
characters that it is on the whole convenient to place them in 
separate families, although, as in so many similar cases, our know- 
ledge of the extinct forms, imperfect as it is, goes far to bridge over 
the distinction between them. 


Family MANATIDA. 


The characters of this and the two following families may be 
conveniently included under the heading of the single genus by which 
they are respectively represented. 

Manatus.\—Incisors 2, rudimentary, concealed beneath the horny 
oral plates, and disappearing before maturity. Molars 44, but 
rarely more than £ present at one time, the anterior teeth falling 
before the posterior come into use; similar in characters from 
beginning to end of the series ; with square, enamelled crowns, the 
grinding surface raised into tuberculated transverse ridges. The 
upper teeth with two ridges and three roots, the lower teeth with 
an additional (posterior) ridge, or talon, and two roots. The cer- 
vical vertebra present the remarkable anomaly of being reduced to 
six in number, the usual vertebral formula being C6, D 17, L 2, 
and C 23-25. Rostrum of the skull, formed by the union of the 
premaxille in front of the anterior narial aperture, shorter than the 
length of the aperture and scarcely deflected from the basi-cranial 
axis ; premaxillz and mandibular symphysis not markedly deflected 
(Fig. 72). Tail entire, rounded, or shovel-shaped. Rudimentary 
nails on the fore limbs. Czcum bifid. Habitat the shores of, 
and the great rivers which empty themselves into, the Atlantic 
within the tropics. These animals are rather fluviatile than marine, 
ascending large rivers almost to their sources. 

The Manatee may be selected for a somewhat full description, 
as being one of the best known representatives of this very remark- 
able order. 

The name Muanati was apparently first applied to this animal hy 
the early Spanish colonists of the West Indies, in allusion to the 
hand-like use which it frequently makes of its fore limbs ; by English 
writers from the time of Dampier (who gives a good account of its 
habits) downwards it has been generally spelt “Manatee.” It was 
placed by Linneeus in his heterogeneous genus Trichechus, but Storr’s 
name Manatus is now generally accepted for it by zoologists. The 
question of the specific distinction of the African and American 
Manatees will be treated of further on, but it will be chiefly to the 
latter and better known form that the following description applies. 

The size of the Manatee has been much exaggerated, but 


1 Storr, Prodromus Meth. Mamm. p. 41 (1780). 


216 SIRENIA 


there is no trustworthy evidence of its attaining a greater length 
than 8 feet. Its general external form may be seen in Fig. 71, 
taken from a living example in the Brighton Aquarium. The 
body is somewhat fish-like, but depressed and ending posteriorly in 
a broad, flat, shovel-like, horizontal tail, with rounded edges. The 
head is of moderate size, oblong, with a blunt, truncated muzzle, 
and divided from the body by a very slight constriction or neck. 
The fore limbs are flattened oval paddles, placed rather low on the 
sides of the body, and showing externally no signs of division into 
fingers, but with a tolerably free motion at the shoulder, elbow, 
and wrist joints, and with three diminutive flat nails near their 
extremities. No traces of hind limbs are discernible either exter- 
nally or internally ; and there is no dorsal fin. The mouth is very 
peculiar, the tumid upper lip being cleft in the middle line into two 
lobes, each of which is separately movable, as will be described in 
speaking of its manner of feeding. The nostrils are two semilunar 


Fic, 71.—American Manatee (Manatus americanus), from life. Proc. Zool. Soc. 1581, p. 457. 


valve-like slits, at the apex of the muzzle. The eves are very 
minute, placed at the sides of the head, and with a nearly circular 
aperture with wrinkled margins. The external ear is a minute 
orifice situated behind the eye, without any trace of pinna. The 
skin generally is of a dark grayish colour, not smooth and glistening 
like that of the Cetacea, but finely wrinkled. At a little distance 
it appears naked, but a close inspection, at all events in young 
animals, shows a scanty covering of very delicate hairs, and both 
upper and under lips are well supplied with short stiff bristles. 

The general form of the skull is seen in Fig. 72. The cerebral 
cavity is rather small as compared with the size of the animal 
and of oblong form; its roof is formed of the parietal bones as 
in ordinary mammals. The squamosal has an extremely large 
and massive zygomatic process, which joins the largely developed 
jugal bone in front. The orbit is small, but prominent and 
nearly surrounded by bone. The anterior nares taken together 
form a lozenge-shaped aperture, which looks upwards and extends 


MANATIDE 217 


backwards considerably behind the orbits. Their sides are formed 
by the ascending processes of the premaxille below, and by the 
supraorbital processes of the frontals above, no traces of nasals 
being found in most skulls, though these bones are occasionally 
present in a most rudimentary condition, attached to the edges 
of the frontals, far away from the middle line, in a position 
quite unique among the Mammalia. In front of the narial aper- 
ture the face is prolonged into a narrow rostrum, formed by 
the premaxille, supported below and at the sides by the maxille. 
The under surface of this is very rugose, and in life covered by a 
horny plate. The rami of the mandible are firmly united together 
at the symphysis, which is compressed laterally, slightly deflected, 
and has a rugose upper surface; to this another horny plate is 
attached, which, with that of the upper jaw, functionally supplies the 


Fic, 72.—Skull of African Manatee (Manatus senegalensis). 4 natural size. 
From Mus. Roy. Coll. Surgeons. 


place of teeth in the anterior part of the mouth. In the young 
state there are rudimentary teeth concealed beneath these horny 
plates, which never penetrate through them, and must therefore be 
quite functionless, and altogether disappear before the animal is full- 
grown. There is besides on each side of the hinder part of both 
upper and lower jaws, a parallel row of molar teeth, similar in 
characters from the beginning to the end of the series, with square 
enamelled crowns raised into tuberculated transverse ridges, some- 
thing like those of the Tapir and Kangaroo. The upper teeth have 
two ridges and three roots; the lower teeth have an additional 
posterior small ridge or talon, and but two roots. These teeth 
succeed each other from before backwards, as in the Proboscidea, 
those at the front of the mouth being worn out and shed before 
those at the back are fully developed. There are altogether about 
eleven on either side of each jaw, but rarely more than six are 


218 SITRENIA 


present at one time. The brain is remarkably simple in structure, 
its hemispheres exhibiting none of the richness of convolution so 
characteristic of the Cetacea. The mammary glands of the female 
are situated just behind and to the inner side of the origin of 
the pectoral limb. The red corpuscles of the blood are among 
the largest of those of any members of the class, averaging in 
diameter, according to Gulliver, zJyy of an inch. 

Manatees pass the whole of their life in the water, inhabiting 
bays, lagoons, estuaries, and large rivers ; but the open sea, so con- 
genial to the Cetacea, is quite unsuited to their peculiar mode of 
life. Asa general rule they prefer shallow water, in which, when 
not feeding, they lie near the bottom, supporting themselves on the 
extremity of the tail, or slowly moving about by the assistance of 
the fore limbs, the tips of which are just allowed to touch the 
ground, and only raising the top of the head above the surface for 
the purpose of breathing at intervals of two or three minutes. In 
deeper water they often float, with the body much arched, the 
rounded back close to the surface, and the head, limbs, and tail 
hanging downwards. The air in the lungs obviously assists them 
to maintain this position, acting in the same manner as that in the 
air-sac of fishes. Their food consists exclusively of aquatic plants, 
on which they browse beneath the water. They are extremely 
slow and inactive in their movements, and perfectly harmless and 
inoffensive. Frequent attempts have been made to keep specimens 
alive in captivity, and sometimes with considerable success, one 
having lived in the Brighton Aquarium for upwards of sixteen 
months. It was fed chiefly on lettuce and endive, but would also 
eat leaves of the dandelion, sow-thistle, cabbage, turnip, and carrot. 
From this and other captive specimens some interesting observations 
upon the mode of life of the animal have been made. One of these 
is the free use it makes of its fore limbs. From the shoulder-joint 
they can be moved in all directions, and the elbow and wrist permit 
of free extension and flexion. In feeding these creatures push the 
food towards their mouths by means of one of the hands, or both 
used simultaneously, and any one who has seen these members thus 
employed can readily believe the stories of their carrying their 
young about under their arms. Still more interesting and quite 
unique among mammals is the action of the peculiar lateral pads 
formed by the divided upper lip, thus described by the late Pro- 
fessor Garrod: “These pads have the power of transversely 
approaching towards and receding from one another simultaneously 
(see Fig. 73, Aand B). When the animal is on the point of seizing 
(say) a leaf of lettuce, the pads are diverged transversely in such a 
way as to make a median gap of considerable breadth. Directly 
the leaf is within grasp the lip-pads are approximated, the leaf is 
firmly seized between their contiguous bristly surfaces, and then 


MANATIDZ 219 


drawn inwards by a backward movement of the lower margin of 
the lip as a whole.” The animal is thus enabled by the unaided 
means of the upper lip to introduce food placed before it without 
the assistance of the comparatively insignificant lower lip, the action 
greatly recalling to the observer that of the mouth of the silkworm 
and other caterpillars, in which the mandibles diverge and converge 
laterally during mastication. When out of water the Manatee is 
an extremely helpless animal; and, although statements are fre- 
quently met with in books of its voluntarily leaving the water for 
the purpose of basking or feeding on shore, all trustworthy ob- 
-servations of those acquainted with it, either in a state of nature 


Fic. 73.—Front view of head of American Manatee, showing the eyes, nostrils, and mouth. 
A, With the lobes of the upper lip divaricated; B, with the lip contracted. From Muzie, 
Trans. Zool. Soc. vol. xi. 


or in captivity, indicate that it has not the power of doing so. 
None of the specimens in confinement have been observed to emit 
any sound. 

Manatees, though much less numerous than formerly, are still 
occasionally found in creeks, lagoons, and estuaries in some of the 
West India Islands, and at various spots on the Atlantic coast of 
America from Florida as far south as about 20° S. lat., and in the 
great rivers of Brazil, almost as high as their sources. They are 
also met with in similar situations on the opposite African coast, 
from about 16° N. to 10°S. lat., and as far into the interior as 
Lake Tchad. Their range may even extend, if native reports 
obtained by Schweinfurth are correctly interpreted, to the river 
Keebaly, 27° E. long. 

A considerable number of specific names have been applied to 
the existing Manatees, but according to the researches of Dr. 
Hartlaub! they may be reduced to three species, distinguished from 
one another, among other features, by the characters of the skull, 
and more especially the relations of the nasals to the adjacent 


1 Zool. Jahrbuch, vol. i. p. 1 (1886). 


225, STRENTA 


hones. Of these the American Manatee may be known as J 
americanus, although it has been described under the names ci 
AL latirestris. and VW. australis. The African Manatee (IL senegalensis) 
ditfers from the American species in the following cranial characters : 
the anterior part of the rostrum is shorter. shallower. and altozether 
smaller; the orbit is smaller; the zvzomatic process is more deep 
and massive; the jugal bone is deeper from above downwards: the 
upper margin of the anterior nares is narrower and with a smooth 
and rounded, instead of a thin and serrated, edze: the upper surface 
oi the irontal is fat. instead of concave; the foramen magnum and 
occipital condyles are narrower from side to side, and the srmphvsis 
of the mandible is smaller and shallower. 

Finally, If inunguiz is a fluviatile species confined to the 
Amazon and Orinoco, which has been but recently fully brouzht 
under the notice of zoolozisis. 


Family Hariconm =. 


Haliesr:.*—In the upper jaw a pair of large, nearly straight, tusk- 
like incisors, directed downwards and iorwards, partially coated 
with enamel. In the male they have persistent pulps, and bevelled 
cutting edges, which project a short distance from the mouth, but 
in the female, though they remain through life in the alveolar 
cavity, they are not exserted, and, the pulp-cavity being filled with 
osteodentine, they soon cease to grow (as in the female Narwhal). 
In the young there is also a second small deciduous incisor on 
each side above. At this aze there are also beneath the horny plate 
which covers the anterior portion of the mandible four pairs of 
slender conical teeth lodged in wide alveolar depressions : these 
become absorbed before the animal reaches maturity. The molars 
are usually 3, sometimes 3, altogether, but not all in place at once. 
as the first falls before the last rises above the eum: they are more 
or less nearly cylindrical in section (except the last, which is com. 
pressed and grooved laterally). without distinction into crown and 
root, increasing in size from before backwards, with persistent pulps 
and noenamel. The summits of the crowns are tuberculated before 
Wearing, afterwards flattened or slizhtly concave. Skull with rostrum 
formed by the union of the premaxille in front of the narial 
aperture, longer than the aperture itself. bending downwards at a 
right angle with the basi-cranial axis, and enclosing the sockets 
of the large incisor tusks. Anterior part of che lower jaw bent 
down in a corresponding manner. Vertebre: C 7, D 1$-19, L and 
C30. Tail broadly notched in the middle line, and with two 
pointed lateral lobes. No nails on the fore limbs. Cxeum sinele. 


+ Titger. Prodromus Sust. Mocnim. et Avinm, p. 120 (1221) 


RHYVTINIDA 221 


The Dugongs are more distinctly marine in their habits than the 
Manatees, feeding chiefly on sea-water alge. They inhabit the 
shallow bays and creeks of the Red Sea, east coast of Africa, 
Ceylon, islands of the Bay of Bengal and the Indo-Malayan 
Archipelago. (including the Philippines), and the north coast of 
Australia, ranging from Barrow Reefs on the west to Moreton Bay 
on the east. Although the distinctive characters are not very 
obvious, they have been divided into three species, according 
to the localities which they respectively inhabit :—Z. tubernacult 
from the Red Sea, H. dugong from the Indian seas, and A. australis 
from Australia. The last-named has lately been the object of a 
regular “fishery,” chiefly on account of its oil, which is peculiarly 
clear, limpid, and free from disagreeable smell, and is said to have 
the same medicinal properties as cod-liver oil. Although often stated 
in books to attain the length of 20 feet when adult, there does 
not appear to be any evidence from actual specimens in museums 
that Dugongs ever reach half that size, 8 feet being the common 
length of adult animals. 

The placentation of this genus has been recently described by 
Sir W. Turner, who first indicated its zonary form. 


Family RHYTINIDA. 


Ehytina.tNo teeth, their place being supplied functionally by 
the dense, strongly-ridged, horny oral plates. Premaxillary rostrum 
about as long as the anterior narial aperture, and moderately 
deflected. Vertebree: C7,D19, Land C 34-37. Head very small 
in proportion to the body. Tail with two lateral pointed lobes. 
Pectoral limbs small and truncated. Skin naked and covered with 
a very thick, hard, rugged, bark-like epidermis. Stomach without 
cecal appendages to the pyloric cavity. Czcum simple. 

Only one species of this genus is known, J. stelleri, the Northern 
Sea-cow, by far the largest animal of the order, attaining the length 
of 20 to 25 feet. It was formerly an inhabitant of the shores of 
two small islands in the North Pacific, Behring and the adjacent 
Copper Island, on the former of which it was discovered by the 
illfated navigator whose name the island bears, when, with his 
accomplished companion, the German naturalist Steller, he was 
wrecked upon it in 1741. Twenty-seven years afterwards (1768), 
as is commonly supposed, the last of the race was killed,” and its 


1 [lliger, Prodromus Syst. Mamm. et Avium, p. 141 (1811).—Amended from 
Rytina. 

2 Nordenskiéld, during his voyage in the Vega, obtained some information 
from the natives of Behring Island which led him to believe that a few individ- 
uals may have survived to a much later date, even to 1854; but this conclusion 
is disputed by later writers. 


a3 STRENTA 


very existence would have been unknown to science but for the 
interesting account of its anatomy and habits left by Steller, and 
the few more or less imperfect skeletons which have recently re- 
warded the researches carried on in the frozen soil of the islands 
around which it dwelt. There is no evidence at present of its 
having inhabited any other coasts than those of the islands just 
named, although it can hardly be supposed that its range was 
always so restricted. When first discovered it was extremely 
numerous in the shallow bays round Behring Island, finding 
abundant nutriment in the large laminarie growing in the sea. 
Its extirpation is entirely due to the Russian hunters and traders 
who followed upon the track of the explorers, and, upon Steller’s 
suggestion, lived upon the flesh of the great Sea-cows. Its 
restricted distribution, large size, inactive habits, fearlessness of 
man, and even its affectionate disposition towards its own kind 
when wounded or in distress, all contributed to accelerate its final 
extinction. 

According to Steller’s account, the Rhytina had a skin of a dark 
brown colour, sometimes spotted or streaked with white. The fore 
limb was covered with short brush-like hairs. 


EXTINCT SIRENLANS. 


Halitherium.—The Miocene and early Pliocene seas of Europe 
abounded in Sirenians, to which the generic name of Halitherium 
was given by Kaup, but which have also received other names. 
They had large tusk-like incisors in the upper jaw, as in the 
existing Dugongs, though not so greatly developed. Their molar 
teeth were 3 or $, anteriorly simple and single-rooted, posteriorly 
those above with three and those below with two roots, and with 
enamelled and tuberculated or ridged crowns, in all which respects 
they more resemble those of the Manatee than of the Dugong. 
The anterior molars were deciduous ; and there is evidence of the 
presence of milk-teeth. Germs of inferior incisors were also 
present. Some species at least had nasal bones, short, broad, 
but normal in position, whereas in all the existing genera these 
bones are quite rudimentary. Another and still more important 
evidence of conformity to the general mammalian type is the 
better development of the pelvic bone, and the presence of a small 
styliform femur articulated to the acetabulum, although no traces 
of any other part of the limb have been discovered. These ancient 
Sirenians, which may be regarded as representing a distinct family 
—Halitheriide—were thus, in dental, cranial, and other osteological 
characters, less specialised than are either of the existing species, 


1 Kaup, Veues Jahrbuch, 1838, pp. $19 and 536. 


HALITHERIHDA 223 


and if the intermediate links could be discovered might well be 
looked upon as the ancestral forms from which the latter have been 
derived, but at present the transitional conditions have not been 
detected. So far as is yet known, when changes in the physical 
conditions of the European seas rendered them unfitted to be the 
habitation of Sirenians, the Halitherium type still prevailed. If the 
existing Dugongs and Manatees are descended from it, their evolu- 
tion must have taken place during the Pliocene and Pleistocene 
epochs, the one in seas to the east, the other to the west of the 
African continent, which has long formed a barrier to their inter- 
communication. Halithertum remains have been found in many 
parts of Germany, especially near Darmstadt, also in France, Italy 
Belgium, Malta, ete. 
Until a few years ago 
none were known from 
England, probably owing 
to the absence of beds 
of an age corresponding 
to those in which they 
are found on the Eu- 


ropean continent; but Fic. 74.—The penultimate and last right lower molars 
a skull and several of Halitherium fossile ; fron the Miocene of the Continent. 


teeth have been detected ‘**" Pe Bamvile) 

among the rolled debris of which the Red Crag of Suffolk is partially 
composed. The species are not yet satisfactorily characterised. 
Some of them appear to have attained a larger size than the existing 
Manatee or Dugong. One of these, from the Pliocene of Italy and 
France, having but 4 molar teeth, has been separated generically 
under the name of Felsinotherium by Capellini, by whom it has been 
fully described; but the difference in the number of the teeth 
does not afford sufficient grounds for separation from Halitherium. 
Miosiren of the Belgian Miocene, differs in that the last upper 
molar is the smallest, in place of the largest of the whole series 
of teeth. 

Other forms.—Remains from the Pliocene of France described as 
Prohalicore are regarded as indicating a Sirenian closely allied to 
Halicore ; while a molar from the Tertiary of California has been 
made the type of Desmotylus, which is likewise referred to the 
Halicoride. Dioplotherium, from the Phosphorites of South Carolina, 
has been considered to connect Halicore with Halitherium, but even 
its ordinal position is uncertain. 

A portion of a skull found in the Pliocene of Belgium has been 
described as Crassitherium by Van Beneden ; and some compressed 
teeth, somewhat similar to but larger than those of the Dugong, 
discovered in the Miocene of the department of Lot-et-Garonne, 
France, gave origin to the genus Fytiodus of E. Lartet. Of this 


224 STRENIA 


genus, which may be identical with TYrachytheriwm of the French 
Miocene, better preserved remains have subsequently been described 
by Delfortrie. These show that the rostrum is more elongated 
than in Hulitheriwm, but the skull is otherwise very similar, as are 
the molar teeth. The incisors are very large, exserted, strongly 
compressed, almost sabre-like, rounded on the upper or anterior 
surface, sharp below, concave on the external and convex on the 
inner side, and transversely striated. 

Pachyacanthus from the Miocene of the Vienna basin is also, ac- 
cording to Van Beneden, another form of Sirenian, of which, however, 
the skull is not known. In various Miocene marine formations of 
the United States of America other remains of Sirenians have 
been found, but mostly in such a fragmentary condition that they 
afford at present little evidence of the early history of the group 
in that country. A more satisfactory discovery is that of a 
nearly complete skull and some bones from a Tertiary limestone 
formation in Jamaica. It is of smaller size than the Manatee, 
and, so far as the teeth are concerned, of a still more generalised 
character than Halitherium, the dentition being apparently i 3, ¢ 4, 


ptm ( rs) =48. The incisors are small, not developed into tusks ; 


the canines (wanting in all existing Sirenians) are rather larger 
than the incisors, judging by the sockets; and the molars are 
bilophodont, and covered with enamel. It has been described 
by Sir R. Owen under the name of Prorastomus sirenoides. Some 
writers regard this genus as the type of a distinct family—the 
Prorastomatide. Unfortunately we have no knowledge of the geo- 
logical antiquity of the formation in which it was embedded. Lastly 
must be mentioned the Hotherium egyptiacum, Owen, founded on the 
cast of a brain, with a small quantity of surrounding bone, discovered 
in the nummulitic limestone of Eocene age in the Mokattam Hills, 
near Cairo. The brain is narrower than in Muanatus, and resembles 
that of Haktherium. This is of interest as the most ancient known 
evidence of any Sirenian whose age has been geologically deter- 
mined. Teeth from the same deposits referred to JJanatus not 
improbably belong really to Eotherium. 

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, so much favoured by modern zoologists, 
receives no very material support from them. 


Bibliography of Sirenia.—J. F. Brandt, Symbol Sirenologicee, St. Petersburg, 
3 fasciculi, 1846-61-68—an exhaustive account of the anatomy, aftinities, and 
literature of the group, with copious illustrations of the osteology of Rhytina. 


CETACEA 225 


Anatomy of Dugong :—Everard Home, Phil. Trans. 1820, p. 315; Owen, Proc. 
Zool. Soc. 1838, p. 29. Placenta of do.:—W. Turner, Trans. Roy. Soc. Edin. 
vol. xxxv. (1889). Janatee:—W. Vrolik, Bijdragen tot de Dierkunde, 1851 ; 
J. Murie, ‘‘On the Form and Structure of the Manatee,” Z'rans. Zool. Soc. Lond. 
vol. viii. p. 127, 1872, and ‘‘ Further Observations on the Manatee,” Jbid. vol. 
xi. p. 19, 1880; A. H. Garrod, ‘‘ Notes on the Manatee recently living in the 
Zoological Society’s Gardens,” Ibid. vol. x. p. 137, 1875; H. C. Chapman, 
‘‘Observations on the Structure of the Manatee,” Proc. Acad. Nat. Sciences of 
Philadelphia, 1875, p. 452; A. Crane, ‘‘ Notes on the Habits of the Manatees in 
Captivity in the Brighton Aquarium,” Proc. Zool. Soc. Lond. 1881, p. 456. 
Extinct Sirenia :—Gervais, Journal de Zoologie, tom. i. p. 832, 1872. R. Lydek- 
ker, Catalogue of Fossil Mammalia in the British Museum, pt. v. 


Order CETACEA. 


This is perhaps the most distinctly circumscribed and natural 
of all the larger groups into which the class is divided. 

The external form is fish-like, the body being fusiform, passing 
anteriorly into the head without any distinct constriction or neck, 
and posteriorly tapering off gradually towards the extremity of the 
tail, which is provided with a pair of lateral, pointed expansions of 
skin supported by dense fibrous tissue, called “flukes,” forming 
together a horizontally-placed triangular propelling organ, notched 
in the middle line behind. 

The head is generally large, in some species attaining to even 
more than one-third of the entire length of the animal, and the 
aperture of the mouth is always wide, and bounded by stiff 
immobile lips. The fore limbs are reduced to the condition of 
flattened ovoid paddles, encased in a continuous integument, show- 
ing no external sign of division into arm, fore arm, and manus, or of 
separate digits, and without any trace of nails. There are no traces 
of hind limbs visible externally. The general surface of the skin is 
smooth and glistening, and devoid of hair, although in many species 
there are a few fine bristles in the neighbourhood of the mouth, 
which may either persist through life, or be present only in the 
young state. Immediately beneath the skin, and intimately 
connected with it, is a thick layer of fat, held together by a dense 
mesh of areolar tissue, constituting the “ blubber,” which serves the 
purpose of the hairy covering of other mammals in retaining the 
heat of the body. In nearly all species a compressed median dorsal 
tegumentary fin is present. The eye is small, and is not provided 
with a nictitating membrane or true lachrymal apparatus. The 
external auditory meatus is a very minute aperture in the skin 
situated at a short distance behind the eye, and there is no vestige 
of a pinna. The nostrils open either separately or by a single 
crescentic valvular aperture, not at the extremity of the snout, but 
near the vertex of the head. 

15 


226 CETACEA 


The bones generally are spongy in texture, the cavities being 
filled with oil. In the vertebral column the cervical region is 
remarkably short and immobile, and the vertebre, originally 
always seven in number, are in many species more or less fused 
together into a solid mass. The odontoid process of the axis, when 
that bone is free, is usually very obtuse, or even obsolete. None 
of the vertebr are united together to form a sacrum. The lumbar 
and caudal vertebre are numerous and large, and, as their arches 
are not connected by any articular processes (zygapophyses), they 
are capable of a very free motion in all directions. The epiphyses 
at the ends of the vertebral bodies are very distinct flattened disks, 
not uniting until after the animal has attained its full dimensions.* 
There are largely developed chevron bones, the presence of which 
indicates the distinction between the caudal and lumbar vertebre. 

The skull (Fig. 75) is modified in a very peculiar manner. The 
brain-case is short, broad, and high, in fact almost spherical. The 
supraoccipital bone rises upwards and forwards from the foramen 
magnum, to meet the frontals at the vertex, thus completely 
excluding the parietals from the upper region of the cranium. The 
frontals are expanded laterally to form the roof of the orbits. The 
anterior narial aperture opens upwards, and has in front of it a 
more or less horizontally prolonged rostrum, formed of the maxillz, 
premaxille, vomer, and mesethmoid cartilage, extending forwards 
to form the upper jaw or roof of the mouth. 

There are no clavicles. The humerus is freely movable on the 
scapula at the shoulder-joint, but beyond this the articulations of 
the limb are imperfect, the flattened ends of the bones coming in 
contact with each other, with fibrous tissue interposed, allowing of 
scarcely any motion. The radius and ulna are distinct, about 
equally developed, and much flattened, as are also all the bones 
of the manus. There are four, or more commonly five digits, and 
the number of the phalanges of the second and third digits always 
exceeds the normal number in mammals, sometimes very con- 
siderably (hyperphalangism) ; they present the exceptional character 
of having epiphyses at both ends.2_ The pelvis is represented by a 
pair of small styliform bones placed longitudinally, suspended below 
and at some distance from the vertebral column at the commence- 
ment of the caudal region. These appear to represent the ischia, 
as the crura of the corpora cavernosa are attached to them. In 
some species, to the outer surface of these are fixed other small 
bones or cartilages, the rudiments of the hind limb. 


1 This is an important distinction from the Sirenia, but a character common 
to nearly all other mammals. It is doubtful whether there is any foundation 
for the statement that these epiphyses remain ununited for an exceptionally long 
period in the Cetacea. 

2 A character repeated in some of the Seals, 


GENERAL CHARACTERS 227 


Teeth are generally present, but exceedingly variable in number. 
In the existing species they are of simple, uniform character, all 
having conical or compressed crowns and single roots, and are never 
preceded by milk-teeth. They are therefore homodont and 
monophyodont. In one group, the Mystacocetes, the teeth are 
absent (except in the foetal condition), and the palate is provided 
with numerous transversely placed horny lamine or “baleen.” 
The salivary glands are rudimentary or absent. The stomach is 
multilocular, its structure being fully noticed under the genus 


wa IP Fp 


Fic, 75.—A section of the skull of a young Dolphin (Globicephalus melas). x}. PMzx, Pre- 
maxilla; Mz, maxilla; ME, ossified portion of the mesethmoid; an, anterior nares; Na, 
nasal; IP, inter-parietal; Fr, frontal; Pa, parietal; SO, supraoccipital; Hx0, exoccipital. 
BO, basioccipital; Sq, squamosal; Per, periotic; AS, alisphenoid; Ps, presphenoid ; Pt 
pterygoid ; pn, posterior nares; Pl, palatine; Vo, vomer; s, symphysis of mandible; id, 


inferior dental canal ; ep, coronoid process of mandible ; cd, condyle ; a, angle; sh, stylo-hyal ; 
bh, basi-hyal ; th, thyro-hyal. (From Flower’s Osteology of Mammalia.) 


Phocena. The intestinal canal is simple, and only in some species 
provided with a small cecum. The liver is very little fissured, and 
there is no gall-bladder. The vascular system is greatly complicated 
by arterial and venous plexuses, or retia mirabiliv. The larynx is of 
peculiar shape, the arytenoid cartilages and the epiglottis being 
much elongated, and together forming a tubular prolongation, which 
projects into the posterior nares, and when embraced by the soft 
palate produces a continuous passage between the nostrils and the 
trachea, as in Ungulates, but in a more perfect manner. The 


228 CETACEA 


brain is large relatively to the size of the animal, very round in 
form, and with its surface divided by sulci into very numerous and 
complex convolutions. The kidneys are deeply lobulated. The 
testes are abdominal. There are no vesicule seminales, nor os 
penis. The uterus is bicornuate, and the placenta nondeciduate 
and diffuse. The mammz are two in number, and the nipples 
placed in depressions on each side of the vulva. The principal 
ducts of the gland are dilated during lactation into large reservoirs, 
into which the milk collects, and from which it is injected by the 
action of a compressor muscle into the mouth of the young animal, 
by which means the process of sucking under water is greatly 
facilitated and expedited. 

The animals of the order Cetacea abound in all known seas, 
and some species are inhabitants of the larger rivers of South 
America and Asia. Their organisation necessitates passing their life 
entirely in the water, as on land they are absolutely helpless. 
They have, however, to rise very frequently to the surface for the 
purpose of respiration ; and, in relation to the constant upward and 
downward movement in the water thus necessitated, their principal 
instrument of motion, the tail, is expanded horizontally, quite 
unlike that of a fish, whose movements are mainly in straight- 
forward or lateral directions. The position of the respiratory orifice 
or nostril on the highest part of the head is very important for 
this mode of life, since it is the only part of the body of which 
the exposure above the surface is absolutely necessary. Of the 
numerous erroneous ideas connected with natural history, few are 
so wide spread and still so firmly believed, notwithstanding repeated 
expositions of its falsity, as that the Cetacea spout out through 
their blowholes water taken in at the mouth. The fact is, the 
“spouting,” or more properly “blowing,” of the Whale is nothing 
more than the ordinary act of expiration, which, taking place at 
longer intervals than in land animals, is performed with a greater 
amount of emphasis. The moment the animal rises to the surface 
it forcibly expels from its lungs the air taken in at the last inspira- 
tion, which of course is highly charged with watery vapour in 
consequence of the natural respiratory changes. This, rapidly 
condensing in the cold atmosphere in which the phenomenon is 
generally observed, forms a column of steam or spray, which has 
been erroneously taken for water. It also often happens, especially 
when the surface of the ocean is agitated into waves, that the 
animal commences its expiratory puff before the orifice has quite 
cleared the top of the water, some of which may thus be driven 
upwards with the blast, tending to complete the illusion. In 
hunting Whales the harpoon often pierces the lungs or air passage’s 
of the unfortunate victim, and then fountains of blood may be 
forced high in the air through the blowholes, as commonly depicted 


GENERAL CHARACTERS 229 


in scenes of Arctic adventure ; but this is nothing more (allowance 
being made for the Whale’s peculiar mode of breathing) than what 
always follows severe wounds of the respiratory organs of other 
mammals. 

All the Cetacea are predaceous, subsisting on living animal food 
of some kind. One genus alone (rca) eats other warm-blooded 
animals, as Seals, and even members of its own order, both large 
and small. Some feed on fish, others on small floating crustaceans, 
pteropods, and medusz, while the principal staple of the food of 
many is constituted by the various species of cephalopods, Loligo 
and other Teuthide, which must abound in certain seas in vast 
numbers, as they form almost the entire support of some of the 
largest members of the order. In size the Cetacea vary much, some 
of the smaller Dolphins scarcely exceeding 4 feet in length, while 
others are the most colossal of all animals. It is true that most 
statements of their bulk found in general and even zoological 
literature are greatly exaggerated, but even when reduced to 
their actual dimensions (which will be stated under the respective 
genera) some of the existing Whales exceed in size any animal 
living either at present or in former times of which we have any 
certain evidence. With some exceptions, the Cetacea generally are 
timid inoffensive animals, active in their movements, and very 
affectionate in their disposition towards one another, especially the 
mother towards the young, of which there is usually but one, or 
at most two ata time. They are generally gregarious, swimming 
in herds or “schools” (so termed by the whalers) sometimes 
amounting to many thousands in number; though some species 
have hitherto only been met with either singly or in pairs. 

Although by their mode of life so far removed from close ob- 
servation that it is impossible to become as familiar with them in 
their natural condition as with many other animals, Whales are in 
many respects the most interesting and wonderful of all creatures ; 
and there is much in their structure and habits well worthy of 
study, much that is difficult to understand, and much that leads to 
great generalisations and throws light upon far-reaching philosophical 
speculations. One of the first lessons which a study of these 
animals affords is that, in the endeavour to discover what a creature 
really is, from what others it is descended, and to what it is related, 
the general outward appearance affords little clue, and we must go 
deep below the surface to find out the essential characteristics of its 
nature. There was once, and may be still in many places, a 
common idea that a Whale is a fish. To realise the fallacy of this 
notion we have only to consider what a fish really is, what under 
all the diversities of form, size, and colour known among fishes 
there is common to them all, and we see that in everything which 
characterises a true fish and separates it from other classes, as 


230 CETACEA 


reptiles, birds, and mammals, the Whale resembles the last-named 
and differs from the fish. It is as essentially a mammal as a Cow 
or a Horse, and simply resembles a fish externally because it is 
adapted to inhabit the same element; but it is no more on that 
account a fish than is a bat, because adapted to pass a great part of 
its existence on the wing in the air, nearly related to a bird. The 
whole structure of a whale is a most instructive instance of a type 
of organisation which is common to and characteristic of the class 
Mammalia, but specially modified or adapted to a peculiar mode of 
life. We see in every part the result of two great principles acting 
and reacting upon each other—on the one hand, adherence to type, 
or rather to fundamental inherited structural conditions, and, on 
the other, adaptation to the peculiar circumstances under which it 
lives, and to which in all probability it has become gradually more 
and more fitted. The external fish-like form is perfectly suited for 
swimming through the water; the tail, however, is not placed 
vertically as in fishes, but horizontally, a position which accords 
better with the constant necessity for rising to the surface for the 
purpose of breathing. The hairy covering characteristic of all 
mammals, which if present might interfere with rapidity of move- 
ment through the water, is reduced to the merest rudiments—a 
few short bristles about the chin or upper lip—which are often 
only present in very young animals; and the function of keeping 
the body warm is supplied by the “blubber.” The fore-limbs, 
though functionally reduced to mere paddles, with no power of 
motion except at the shoulder-joint, have beneath their smooth and 
continuous external covering all the bones, joints, and even most of 
the muscles, nerves, and arteries of the human arm and hand; and 
the rudiments of hind legs found buried deep in the interior of the 
animal apparently subserve no useful purpose, but point an in- 
structive lesson to those who are able to read it. 

As before said, the Cetacea form a perfectly well-defined group, 
sharply separated from all other mammals, and with no outlying or 
doubtful forms at present known. 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 which are reckoned 
as orders. The extinct Zewglodon, so far as its characters are known, 
does not fall into either of these groups, but is in some respects an 
annectant form, and therefore must be placed, provisionally at least, 
in a third group by itself. 

The Mystacocetes appear at first sight to be the most specialised 

+and aberrant of the existing Cetacea, as indicated by the absence of 
teeth, the presence of baleen, and the form and size of the mouth ‘ 
but, as we see in other groups, dental characters, and all such as 


GENERAL CHARACTERS 231 


relate to the prehension of food generally, are essentially adaptive 
and consequently plastic or prone to variation, and hence can- 
not well be relied upon as tests of affinity. In another character, 
also adaptive, the laxity of the connection of the ribs with the 
vertebral column and with the sternum, and the reduction of that 
bone in size, allowing great freedom of expansion of the thoracic 
cavity for prolonged immersion beneath the water, the Mystacocetes 
have passed beyond the Odontocetes in specialisation. On the other 
hand, the greater symmetry of the skull, the more anterior position 
of the external nostrils and their double external orifice, the form 
of the nasal bones, the presence of a distinctly developed olfactory 
organ, the mode of attachment of the periotic bone to the cranium, 
the presence of a cecum and the regular arrangement of the 
alimentary canal, the more normal characters of the manus and the 
better development of the muscles attached to it, and the presence, 
in many species at least, of parts representing not only the bones 
but also the ligaments and muscles of a hind limb,! all show less 
deviation from the ordinary mammalian type than is presented by 
the Odontocetes. Taking all these characters into consideration, it 
does not appear reasonable to suppose that either type has been 
derived from the other, at all events in the form in which we see 
it now, but rather that they are parallel groups, both modified in 
different fashions from common ancestors. 

Among the Mystacocetes, in the especially distinguishing 
characters of the division, the Rorquals are less specialised than the 
Right Whales, which in the greater size of the head, the length and 
compression of the rostrum, the development of the baleen, and 
shortness of the cervical region, are exaggerated forms of the type, 
and yet they retain more fully some primitive characters, as the 
better development of the hind limb, the pentadactylous manus, 
and the absence of a dorsal fin. Both types are found distinct in 
a fossil state at least as far back as the early Pliocene age, but: 
generally represented by smaller species than those now existing. 
Some of the Pliocene Rorquals (Cetotherium) were, in the elongated 
flattened form of the nasal bones, the greater distance between the 
occipital and frontal bone at the top of the head, and the greater 
length of the cervical vertebrae, more generalised than those now 
existing. In the shape of the mandible also, Van Beneden, to 
whose researches we are much indebted for a knowledge of these 
forms, discerns some approximation to the Odontocetes. 

Among the last-named group there are several distinct types, of 
which that represented by Platanista, although in some respects 
singularly modified, has been considered to present on the whole 
approximations towards the more normal and general type of 


1 These have been described in detail by Professor Struthers in the Journal of 
Anatomy and Physiology, 1881. 


232 ; CETACEA 


mammalian structure. It is therefore interesting to find an 
apparently allied form well represented among the earliest fossil 
remains of Cetaceans in Europe. Almost all the other members of 
the suborder range themselves under the two principal heads of 
Ziphioids (or Physeteroids) and Delphinoids. The former is an 
ancient and once abounding type, of which the Sperm Whale 
(Physeter) is a highly specialised form. Among the latter, Globi- 
cephalus is a modified form as regards the structure of its anterior 
extremity, and Monodon as regards its dentition, while Delphinus, 
with the various allied genera, may be regarded as the domi- 
nating type of Cetaceans at the present day, abundant in slightly 
differentiated species and also in individuals. They are in this 
respect to the rest of the order much as the hollow-horned 
Ruminants are to the other Ungulates. 

The earliest Cetaceans of whose organisation we have anything 
like complete evidence are the Zeuglodonts of the Eocene period,! 
which approach in the structure of the skull and teeth to a much more 
generalised mammalian type than either of the existing suborders. 
The smallness of the cerebral cavity compared with the jaws and the 
rest of the skull they share with the primitive forms of many other 
types. The forward position of the narial aperture and the length 
and flatness of the nasal bones, which distinguish them from all 
existing forms, we must also suppose to be a character at one time 
common to all Cetaceans, though now retained (but to a less degree) 
only by the Mystacocetes. Even Sgualodon, which in its heterodont 
dentition so much resembles Zeuglodon as to have been placed by 
some zoologists in the same genus, entirely differs from it, and 
conforms with the ordinary Dolphins in its essential cranial 
characters. 

The origin of the Cetacea is at: present involved in much ob- 
scurity. They present no signs of closer affinity to any of the 
lower classes of vertebrates than do many other members of their 
own class. Indeed in all that essentially distinguishes a mammal 
from the oviparous vertebrates, whether in the osseous, nervous, 
reproductive, or any other system, they are as truly mammalian as 
any other group. Any supposed marks of inferiority, as absence 
of limb structure, of hairy covering, of lachrymal apparatus, etc., are 
obviously modifications (or degradations, as they may be termed) 
in adaptation to their special mode of life. The characters of the 
teeth of Zeuglodon and other extinct forms, and also of the foetal 
Mystacocetes, clearly indicate that they have been derived from 
mammals in which the heterodont type of dentition was fully 


1 The ankylosed mass of cervical vertebr, on which the genus Paleocetus was 
established, was regarded by its describer as having probably come from the 
Kimeridge Clay, but the mineral condition of the specimen points to the Red 
Crag as the place of origin. 


GENERAL CHARACTERS 233 


established. The steps by which a land mammal may have been 
modified into a purely aquatic one are indicated by the stages 
which still survive among the Carnivora in the Otariide and in 
‘the true Seals. A further change in the same direction would pro- 
duce an animal somewhat resembling a Dolphin; and it has been 
thought that this may have been the route by which the Cetacean 
form has been developed. There are, however, great difficulties in 
the way of this view. Thus if the hind limbs had ever been 
developed into the very efficient aquatic propelling organs they 
present in the Seals, it is not easy to imagine how they could have 
become completely atrophied and their function transferred to the 
tail. So that from this point of view it is more likely that Whales 
were derived from animals with long tails, which were used in 
swimming, eventually with such effect that the hind limbs became 
no longer necessary. The powerful tail, with its lateral cutaneous 
flanges, of an American species of Otter (Lutra brasiliensis) may give 
an idea of this member in the primitive Cetaceans. But the struc- 
ture of the Cetacea is, in so many essential characters, so unlike 
that of the Carnivora that the probabilities are against these orders 
being nearly related. Even in the skull of the Zeuglodon, which 
has been cited as presenting a great resemblance to that of a Seal, 
quite as many likenesses may be traced to one of the primitive Pig- 
like Ungulates (except in the purely adaptive character of the form 
of the teeth), while the elongated larynx,! complex stomach, simple 
liver, reproductive organs both male and female, and foetal mem- 
branes of the existing Cetacea are far more like those of that group 
than of the Carnivora. Indeed it appears probable that the old 
popular idea which affixed the name of “Sea-Hog”? to the Porpoise 
contains a larger element of truth than the speculations of many 
accomplished zoologists of modern times. The fact that Platanisia, 
which, as mentioned above, appears to retain more of the primitive 
characteristics of the group than any other existing form, and also 
the somewhat related Inia from South America, are both at the 
present day exclusively fluviatile, may point to the fresh-water origin 
of the whole group, in which case their otherwise rather inexplic- 
able absence from the seas of the Cretaceous period would be 
accounted for. 

On the other hand, it should be observed that the teeth of the 
Zeuglodonts approximate more to a carnivorous than to an ungulate 
type. It is scarcely necessary to allude to the hypothesis started 
by some Continental writers to the effect that the Whales are the 
most primitive type of mammals with which we are acquainted, 


1 There is much resemblance in the larynx of the Hippopotamus, but none 
in that of the Seal, to the same organ in the Cetacea. 

2 German Meerschwein, whence the French Marsowin. ‘‘ Porpoise” is said 
to be derived from ‘‘ Porc-poisson.”’ 


234 CETACEA 


and that they are the descendants of the Mesozoic reptilian order 
Ichthyopterygia, from which their hyperphalangism is a direct 
inheritance. The Ichthyopterygia have been shown, on very strong 
evidence, to have been derived from land reptiles, and to have 
gradually acquired their hyperphalangism as an adaptive character 
suitable to their peculiar mode of life, and there can be but little 
doubt that a similar adaptation has taken place in the case of the 
Whales. 


Suborder MYSTACOCETI, 
the BALENOIDEA, Whalebone, or True IV hates. 


Family BALZENIDA. 


Teeth never functionally developed, but always disappearing 
before the close of intra-uterine life. Palate provided with plates 
of baleen or “whalebone.” Skull symmetrical. Nasal bones form- 
ing a roof to the anterior nasal passages, which are directed upwards 
and forwards. Maxilla produced in front of, but not over, the 
orbital process of the frontal. Lachrymal bones small and distinct 
from the jugal. Tympanic bone involuted (Fig. 76), and ankylosed 
with the periotic, which is attached to the base of the cranium by 
two strong diverging processes. Olfactory organ distinctly de- 
veloped. Rami of mandible arched outwards, their anterior ends 
meeting at an angle, and connected by fibrous tissue without any 
true symphysis. All the ribs at their upper extremities articulating 
only with the transverse processes of the vertebra; their capitular 
processes, when present, not articulating directly with the bodies of 
the vertebr. Sternum composed of a single piece, and articulating 
only with a single pair of ribs. No ossified sternal ribs. External 
openings of nostrils distinct from each other, longitudinal. A short 
conical cecum. 

These animals have, when in the foetal state, numerous minute 
calcified teeth lying in the dental groove of both upper and lower 
jaws. They are best developed about the middle of foetal life, after 
which period they are absorbed, and no trace of them remains at the 
time of birth.? The baleen or whalebone does not make its appear- 
ance until after birth. It consists of a series of flattened horny 
plates, between three and four hundred in number, on each side of 


1Tcel. Avalr; Dan. and Swed. heal; Anglo-Saxon hwel; Germ. wal, 
walfisch, The meaning apparently is ‘‘roller,” the word being closely allied to 
“wheel” (Skeat). 

° These were discovered in the Greenland Whale by Geoffroy St. Hilaire, 
whose observations were confirmed and extended to other genera by Eschricht. 
They have been very fully described in Budwnoptera rostrata by Julin (Arechivs 
de Biologic, i. 1880). 


BALENIDE 235 


the palate, with a bare interval along the middle line. These plates 
are placed transversely to the long axis of the palate, with very short 
intervals between them. Lach plate or blade is somewhat triangular 
in form, with the base attached to the palate and the apex hanging 
downwards. The outer edge of the blade is hard and smooth; but 
the inner edge and apex fray out into long bristly fibres, so that the 
roof of the Whale’s mouth looks as if covered with hair, as described 
by Aristotle. At the inner edge of each principal blade are two 
or three much smaller or subsidiary blades. The principal blades 
are longest near the middle of the series, and gradually diminish 
towards the front and back of the mouth. The horny plates grow 
from a dense fibrous and highly vascular matrix, covering the 
- palatal surface of the maxille, and sending out lamellar processes, 
one of which penetrates the base of each blade. Moreover, the 
free edge of these processes is covered with very long vascular 
thread-like papille, one of which forms the central axis of each of 
the hair-like epidermic fibres of which the blade is mainly composed. 
A transverse section of fresh whalebone shows that it is made up of 
numbers of these soft vascular papill, circular in outline, each. 
surrounded by concentrically arranged epidermic cells, and the 
whole bound together by other epidermic cells, that constitute the 
smooth cortical (so-called “ enamel”) surface of. the blade, which, 
disintegrating at the free edge, allows the individual fibres to 
become loose and assume the hair-like appearance before spoken of. 
These fibres differ from hairs in not being formed in depressed 
follicles in the enderon, but rather resemble the fibres composing the 
horn of the Rhinoceros. The whalebone in fact consists of nothing 
more than modified papille of the buccal mucous membrane, with 
an excessive and cornified epithelial development. The blades are 
supported and bound together for a certain distance from their 
base by a mass of less hardened epithelium, secreted by the surface 
of the palatal membrane or matrix of the whalebone in the intervals 
of the lamellar processes. This is the “intermediate substance” of 
Hunter, the “gum” of the whalers. Baleen varies much in colour 
in different species. In some it is almost jet black, in others slate- 
colour, horn-colour, yellow, or even creamy-white. In some the 
blades are variegated with longitudinal strips of different hues. 
Baleen differs also greatly in other respects, being short, thick, 
coarse, and stiff in some, and greatly elongated and highly elastic 
in those species in which it has attained its fullest development. 
Its function is to strain the water from the small marine molluscs, 
crustaceans, or fish upon which the Whales subsist. In feeding the 
immense mouth is filled with water containing shoals of these small 
creatures, and then, on the Whale closing the jaws and raising the 
tongue, so as to diminish the cavity of the mouth, the water streams 
out through the narrow intervals between the hairy fringe of the 


236 CETACEA 


whalebone blades, and escapes through the lips, leaving the living 
prey to be swallowed.? 

Our knowledge of the different structural modifications attained 
by members of this important group of mammals, though largely 
increased of late years, is still imperfect. Formerly they were all 
divided into Right Whales (Balena) and Rorquals or Fin-Whales 
(Balenoptera), the latter distinguished by their smaller heads, 
elongated and slender form, free cervical vertebre, tetradactylous 
manus, and the presence of very conspicuous longitudinal furrows or 
folds in the skin of the throat and chest, and of a small adipose 
dorsal fin. Recent discoveries have, however, brought to light 
several forms holding a somewhat intermediate position, and pre- 
senting combinations of characters not found in either of the longer 
known sections. According to our present knowledge the group is 
naturally divided into five very distinct genera, of which the leading 
characters are given below. 

Balena.2—Skin of throat smooth, not furrowed. No dorsal fin. 
Cervical vertebra united into a single mass. Pectoral limb short, 
broad, and pentadactylous. Head very large. Baleen very long 
and narrow, highly elastic, and black. Scapula high, with a distinct 
coracoid and acromion process. Tympanic (Fig. 78) deep and angular, 
its inflation comparatively slight, and the involuted portion not fig- 
shaped, and frequently without a well-marked depression at the 


anterior extremity of the superior border of the inner surface for 
the Eustachian canal. 


Fic. 76.—Greenland or Arctic Right Whale (Balcena mysticetus). 


The Greenland, or more properly Arctic, Right Whale (Balena 
mysticetus) attains, when full grown, a length of from 45 to 50 
feet. Its usual vertebral formula is C 7, D 12, L 14, © 29. 
The external form is shown in Fig. 76 from a careful drawing by 


1 For the structure of whalebone see Hunter, ‘‘ Observations on the Structure 
and Economy of Whales,” Phil. Trans. 1787 ; Eschricht and Reinhardt, On the 
Greenland Right Whale, English translation by the Ray Society, 1866, pp. 67-78 ; 
and Sir W. Turner, in Trans. Roy. Soc. Edin. 1870. 

2 Linn. Syst. Nat. 12th ed. vol. i. p. 105 (1766). 


BALENIDE 237 


Mr. Robert Gray. In this species all the peculiarities which 
distinguish the head and mouth of the Whales from those of other 
mammals have attained their greatest development. The head is 
of enormous size, exceeding one-third of the whole length’ of the 
creature. The cavity of the mouth is actually larger than that of 
the body, thorax and abdomen together. The upper jaw is very 
narrow, but greatly arched from before backwards, to increase the 
height of the cavity and allow for the great length of the baleen 
blades ; the rami of the mandible are widely separated posteriorly, 
and have a still further outward sweep before they meet at 
the symphysis in front, giving the floor of the mouth the shape 
of an immense spoon. The baleen blades attain the number 
of 380 or more on each side, those in the middle of the series 
having a length of 10 or sometimes 12 feet. They are black in 
colour, fine and highly elastic in texture, and fray out at the inner - 
edge and ends into long, delicate, soft, almost silky, but very tough, 
hairs. The remarkable development of the mouth and the structures 
in connection with it, which distinguishes the Right Whale among 
all its allies, is entirely in relation to the nature of its food. It 
is by this apparatus that the animal is enabled to avail itself of 
the minute but highly nutritious crustaceans and pteropods which 
swarm in immense shoals in the seas it frequents. The large mouth 
enables it to take in at one time a sufficient quantity of water filled 
with these small organisms, and the length and delicate structure 
of the baleen provide an efficient strainer or hair-sieve by which the 
water can be drained off. If the baleen were rigid, and only as 
long as is the aperture between the upper and lower jaws when the 
mouth is shut, a space would be left beneath it when the jaws were 
separated, through which the water and the minute particles of food 
would escape together. But instead of this the long, slender, 
brush-like, elastic ends of the whalebone blades fold back when 
the mouth is closed, the front ones passing below the hinder 
ones in a channel lying between the tongue and the lower jaw. 
When the mouth is opened, their elasticity causes them to 
straighten out like a bow unbent, so that at whatever distance 
the jaws are separated the strainer remains in perfect action, 
filling the whole of the interval. The mechanical perfection of 
the arrangement is completed by the great development of the 
lower lip, which rises stiffly above the jaw-bone and prevents the 
long, slender, flexible ends of the baleen from being carried 
outwards by the rush of water from the mouth, when its cavity 
is being diminished by the closure of the jaws and raising of the 
tongue. 

If, as appears highly probable, the “bowhead” of the Okhotsk 
Sea and Behring Strait belongs to this species, its range is cireum- 
polar. Though found in the seas on both sides of Greenland, and 


238 CETACEA 


passing freely from one to the other, it is never seen so far south 
as Cape Farewell ; but on the Labrador coast, where a cold stream 
sets down from the north, its range is somewhat farther. In the 
Behring Sea, according to Scammon, “it is seldom seen south of 
the fifty-fifth parallel, which is about the farthest southern extent 
of the winter ice, while on the Sea of Okhotsk its southern limit is 
about the latitude of 54°.” As has been abundantly shown by 
Esehricht and Reinhardt in the case of the Greenland seas, “ every- 
thing tends to prove,” Scammon says, “that the Bulan inysticetus 
is truly an ‘ice whale,’ for among the scattered floes, or about the 
borders of the ice-fields or barriers, is its home and feeding-ground. 
It is true that these animals are pursued in the open water during 
the summer months; but in no instance have we learned of their 
being captured south of where winter ice-fields are occasionally met 
with.” The occurrence of this species, therefore, on the British or 
any European coast is exceedingly unlikely, as when alive and in 
health the southern limit of its range in the North Sea has been 
ascertained to be from the east coast of Greenland at 64° N. lat. 
along the north of Iceland towards Spitzbergen, and a glance at a 
physical chart will show that there are no currents setting south- 
wards which could bear a disabled animal or a floating carcase to 
British shores. To this & priori improbability may be added the 
fact that no authentic instance has been recorded of the capture or 
stranding of this species upon any European coast; for the cases 
in which it has been reported as seen in British waters may be ex- 
plained by the supposition of one of the other species of the genus 
being mistaken for it. Still, as two other essentially Arctic 
Cetaceans, the Narwhal and the Beluga, have in a few undoubted 
instances found their way to British shores, it would be rash 
absolutely to deny the possibility of the Greenland Right Whale 
doing the same. 


Mia. 77.—Southern Right Whale (Balirni corstralis). 


i : Fling “ ix, - is ipa tee: 

The southern Right Whale (B. australis, Fig. 77) resembles the 
last in the absence of dorsal fin and of longitudinal fwrows in the 
skin of the throat and chest, but differs in that it possesses a smaller 
head in proportion to its body, shorter baleen, a different shaped 
contour of the upper margin of the lower lip, and a greater number 


BALA NIDA 239 


(fifteen) of ribs and dorsal vertebre. This form inhabits the tem- 
perate seas of both northern and southern hemispheres, and is 
divided into several so-called species, according to their geographical 
distribution :—B. biscayensis of the North Ailantic, B. japonica of 
the North Pacific, B. australis of the South Atlantic, and B. anti- 
poderum and B. novee-zealandic of the South Pacific. The differential 
characters by which they have been separated, external as well as 
anatomical, are, however, slight and subject to individual variation ; 
and the number of specimens available for comparison in museums 
is not yet sufficient to afford the necessary data to determine 
whether these characters can be regarded as specific or not. 
The most interesting of these is the Atlantic Right Whale, 
which was formerly abundant in the North Atlantic, but is 
now so scarce as to appear verging on extinction. This was 
the Whale the pursuit of which gave occupation to a numerous 
population on the shores of the Basque provinces of France and 
Spain in the Middle Ages. From the tenth to the sixteenth centuries 
Bayonne, Biarritz, St. Jean de Luz, and San Sebastian, as well as 
numerous other towns on the north coast of Spain, were the centres 
of an active Whale “fishery,” which supplied Europe with oil and 
whalebone. In later times these Whales were pursued as far as the 
coast of Newfoundland. They were, however, already getting scarce 
when the voyages undertaken towards the close of the sixteenth 
century for the discovery of the north-eastern route to China and 
the East Indies opened out the seas around Spitzbergen ; then for 
the first time the existence of the Greenland Whale became known, 
and henceforth the energies of the European whale-fishers were 
concentrated upon that animal. It is a singular fact that the 
existence of the Atlantic Right Whale was quite overlooked by 
naturalists till lately, all accounts referring to it being attributed to 
the Greenland Whale, supposed once to have had a wider distribu- 
tion than now, and to have been driven by the persecution of man 
to its present circumpolar haunts. To the two Danish cetologists 
Eschricht and Reinhardt is due the credit of having proved its 
existence as a distinct species, from a careful collation of numerous 
historical notices of its structure, distribution, and habits; and their 
restoration of the animal, founded upon these documents, has been 
abundantly confirmed by the capture of various specimens in recent 
times, showing that it still lingers in some of the localities where it 
formerly was so abundant. The only known instances of its 
occurrence on the coasts of Europe in modern times are in the 
harbour of San Sebastian in January 1854, in the Gulf of Taranto, 
in the Mediterranean, in February 1877, and on the Spanish coast 
between Guetaria and Zarauz (Guipuzcoa) in February 1878. The 
skeletons of these three whales are preserved in the museums of 
Copenhagen, Naples, and San Sebastian respectively. On the coast 


240 CETACEA 


of the United States several Whales of this species have been taken 
within the last few years. In the North Pacific a very similar if 
not identical species is regularly hunted by the Japanese, who tow 
the carcases ashore for the purposes of flensing and extracting 
the whalebone. In the tropical seas, however, according to Captain 
Maury’s whale charts, Right Whales are never or rarely seen; but 
the southern temperate ocean, especially the neighbourhood of the 
Cape of Good Hope, Kerguelen’s Island, Australia, and New Zea- 
land, is inhabited by “Black Whales,” once abundant, but now 
nearly exterminated through the wanton destruction of the females 
as they visit the bays and inlets round the coast, their constant 
habit in the breeding time. The range of these Whales southward 
has not been accurately determined ; but no species corresponding 
with the Arctic Right Whale has as yet been met with in the 
Antarctic icy seas. 


Fic. v8.—The right tympanic bone of an immature individual of the Greenland Whale 
(Balena mysticetus), from the inner (4) and outer (B) aspects. 3 natural size. (From the 
Proce. Zool. Soc.) 


_ Remains of Right Whales are of not uncommon occurrence in the 
Pliocene Crag deposits of England and Belgium. The tympanics 
of B. afinis from these deposits appear to indicate a species closely 
allied to B. mysticctus, in which this bone is long and angulated 
anteriorly (Fig. 78); while the tympanics from the same deposits 
described as B. primigenia are shorter and more rounded at the 
antero-inferior angle, thus resembling those of B. australis. A 
smaller species, having an estimated length of about 20 feet, has 
been described as Balenula balenopsis, the generic distinction being 
made on account of the free condition of the atlas and seventh 
cervical vertebre ; but it seems scarcely advisable to regard such a 
feature as indicating more than a less specialised species. Balena 
(Balenotus) insignis is a whale of somewhat larger dimensions, in 
which the atlas is generally, and the seventh cervical vertebra, 


BALA NIDA 241 


always, free, while in young individuals the axis vertebra may 
likewise be separate. 

Neobalena..—Head about one-fourth the total length. Skin of 
the throat not plicated. A small falcate dorsal fin. Vertebre, 
C7,D17,L3,C16=43 The cervical vertebre are united. The 
manus small, narrow, and tetradactylous, wanting the pollex. The 
ribs remarkably expanded and flattened. The scapula very low 
and broad, with completely developed acromion and coracoid pro- 
cesses. Tympanic approximating to that of Balena, but with certain 
very characteristic peculiarities of shape. Baleen very long, slender, 
elastic, and white. A single species, at present very rare, VV. mar- 
ginata, from the Australian and New Zealand seas is the smallest 
of the Whalebone Whales, being not more than 20 feet in length. 

Rhachianectes.2—This combines the small head, elongated form, 
and narrow pectoral fin of Balcenoptera with the smooth skin of the 
throat and absence of the dorsal fin of Balena. The baleen is the 
shortest and coarsest of any of the group. Its osteology is im- 
perfectly known. One species, 2. glaucus, the Gray Whale of the 
North Pacific. 

Mlegaptera.2—Head of moderate size. Baleen plates short and 
broad. Vertebre, C 7, D 14, L 11, C 21=53. Cervical vertebrae 
free. Scapula with acromion and coracoid process absent or rudi- 
mentary. Skin of throat plicated. Dorsal fin low. Pectoral limb 
tetradactylous, very long and narrow, attaining about one-fourth of 
the length of the entire animal, the metacarpus and phalanges 
being greatly developed, and the latter very numerous. Tympanic 
still more inflated than in Balenoptera, with the involuted portion 
more distinctly pyriform, the Eustachian part of the aperture well 
defined, and two well-marked longitudinal ridges on the lower 
surface of adult specimens. 

The Whale commonly called “ Humpback” (Megaptera boops) by 
whalers, perhaps on account of the low hump-like form of the 
dorsal fin, is very distinctly characterised from all others of the 
group, especially by the immense length of the pectoral fins or 
flippers, which are indented or scalloped along their margins, and 
are, except at their base, of a white colour, nearly all the rest of 
the body being black. The baleen plates are. of a deep black 
colour. Though common in the North Atlantic between Norway 
and Greenland, this Whale does not frequently appear on the coasts 
of the British Isles. One came ashore at Newcastle in 1839; 
another, a young one, was taken in the estuary of the Dee in 1863, 
and its skeleton is preserved in the Liverpool museum; and a 
nearly full-grown animal was captured in the mouth of the Tay in 

1 Gray, Suppl. Cat. Seals and Whales in Brit. Mus. p. 39 (1871). 
2 Cope, Proc. Ac. Nat. Sei. Philad. 1869, p. 15. 
2 Gray, Zoology of Ercbus and Terror, p. 16 (1846). 
16 


BALA NIDZ 243 


have the plicated skin of the throat like that of Meguptera, the 
furrows being more numerous and close set; but the pectoral 
fin is comparatively 
small, the dorsal fin 
distinct and falcate, 
and the tail very 
much compressed 
before it expands 
into the “flukes.” 
The Rorquals are 
perhaps the most 
abundant and widely 
distributed of all the 
whales, being found 
in some of their 
modifications in all 
seas, except the ex- 
treme Arctic, and 
probably Antarctic 
regions. Owing to 
the small quantity 
and inferior quality 
of their whalebone, 
the comparatively 
limited amount of 
blubber, and their 
great activity and 


the difficulty of cap- B 
turing them by the Fic. 81.—The right tympanic of Balenoptera musculus from 


the inner (A) and outer (B) aspects. natural size. (F th 
old methods, these 5,,, Lah cae ere a ar 


Whales were not 

until recently an object of pursuit by whale-fishers; but, since the in- 
troduction of steam-vessels, and especially of explosive harpoons fired 
from guns in the place of those hurled by the human hand, a regular 
fishery has been established on the coast of Finmark, There are four 
distinct species of this genus in British seas. (1) Balwnoptera sib- 
baldi, the “Blue Whale,” the largest of all known animals, attains a 
length of 80 or even sometimes 85 feet. Its colour is dark bluish 
gray, with small whitish spots on the breast ; the baleen is black ; 
the flippers are larger proportionally than in other Rorquals, 
measuring one-seventh of the total length of the body; and the dorsal 
fin is small and placed very far back. This Whale has usually 64 
vertebree, of which 16 bear ribs. Like the others of the genus, this 
species seems to pass the winter in the open seas, and approaches the 
coast of Norway at the end of April or beginning of May. At this 
time its sole food is a small crustacean (Zuphwusia inermis) which 


My 2 ea < " 
i Se * \ 
nen - 


Uy 


a 5, aM 
s Mi 
FMA P \) 
pu 
Med LU Loki 


\N . KAN \\ 
, WW oe . 


\ 

\ 
NAY 
ANN 
AK 

\ ‘ \ 
AR 
AN 


244 CETACEA 


swarms in the fjords. Several specimens have been taken on the 
British coasts, two fine skeletons from the Firth of Forth being pre- 
served in the Edinburgh museums. (2) Balenoptera musculus, the 
Common Rorqual, has a length of 65 to 70 feet, is of a grayish slate 
colour above and white underneath, and the baleen is slate colour 
variegated with yellow or brown. It has usually 62 vertebre, 
of which 15 bear ribs. This is the commonest of all the large 
Whales on the British coasts, scarcely a winter passing with- 
out the body of one being somewhere washed ashore, usually 
after stormy weather, and more frequently on the south coast, 
as this species has a more southern range than the last, and 
frequently enters the Mediterranean. It feeds largely on fish, 
and is frequently seen feasting among shoals of herring. (3) 
Balenoptera borealis, often called Rudolphi’s Whale from its first 
describer, is a smaller species, scarcely attaining a length of 50 feet. 
It is bluish-black above, with oblong, light-coloured spots, whilst 
the under parts are more or less white; the whole of the tail and 
both sides of the flippers are black; the baleen is black, and the 
bristly ends fine, curling, and white; the flippers are very small, 
measuring one-eleventh of the total length of the body. There are 
56 vertebra, with 14 pairs of ribs. This species, according to 
Collett, feeds chiefly on minute crustaceans, mainly Calanus finmar- 
chicus and Euphausia inermis, and not on fish. Until lately it was 
considered the rarest of the Whales of European seas, and was only 
known to science from a few individuals stranded on the coasts of 
northern Europe at long intervals, the skeletons of which have been 
preserved in museums. The most southern point at which it has 
been met with hitherto is Biarritz in France. Since the establish- 
ment of the whaling station near the North Cape it has been shown 
to be a regular summer visitor, and in 1885, 771 individuals were 
captured on the coast of Finmark. (4) Balenoptera rostrata, the 
lesser Fin-Whale or Rorqual, is the smallest species found in the 
northern seas, rarely exceeding 30 feet in length. Its colour is 
grayish-black above, whilst the under side is white, including the 
whole of the lower side of the tail; the inner side of the flippers 
is white ; and there is a broad white band across the outer side, 
which is a very characteristic mark of the species; the baleen is 
yellowish-white. The dorsal fin in this and the last species is 
comparatively high, and placed far forwards on the body. This 
Whale has usually 48 vertebra, 11 of which bear ribs. It is common 
in summer in the fjords of Norway, and is often seen around the 
British Isles. It has been taken, though rarely, in the Mediterranean ‘ 
and ranges as far north as Davis’s Straits. 

Rorquals are met with in almost all seas throughout the world, 
but further and more accurate observations are required before 
their specific characters and geographical distribution can be made 


BALENIDE 245 


out. Nearly all the individuals hitherto examined with any care, 
whether from the North Pacific, the Australian seas, or the Indian 
Ocean, come very near in structure to one or the other of the 
Atlantic forms described above, so much so that some zoologists 
have been induced to believe that there are but four species, each 
of which has a wide, almost cosmopolitan range, while others have 
described and named almost every individual specimen captured as 
belonging to a different species.+ 

Tympanics, vertebre, and other bones of Rorquals are among 
the commonest cetacean remains found in the Pliocene Crags of 
England and Belgium. Several species, varying in dimensions, are 
known from these deposits, B. definita (sibbaldina) being apparently 
nearly related to the existing B. sibbaldi. A caudal vertebra from 
the Upper Eocene of Hampshire has been referred to Balenoptera, but 
does not afford sufficient evidence to prove the existence of the 
genus at that date. 

Extinct Genera.—The extinct genus Cetotherium of the European 
Pliocene may be taken to include a number of fossil Whalebone 
Whales allied to the Balenopterine group, several of which have 
been described under other names, such as Plesiocetus, Heterocetus, 
and Amphicetus. They are readily characterised by the form of 
the tympanic bone, which is much narrower in front than behind, 
the roughened inferior surface being in the shape of an isosceles 
triangle, and the notch for the Eustachian canal being smaller, and 
descending nearer to the inferior border of the inner wall than in 
Balenoptera. The skull is longer than the latter, with a greater 
interval between the occiput and the frontal, and with longer and 
more flattened nasals. The relative thickness of the cervical 
vertebre is also greater. In the typical forms (eg. C. brialmonti 
and C. dubiwm) the mandibular condyle is simple; but in C. 
(Heterocetus) brevifrons it is furnished with a projecting posterior 
talon, as in the Sperm Whale. 

Herpetocetus is known by a comparatively small species from the 
Belgian and English Crags, characterised by the extreme inflation 
of the egg-shaped tympanic bone, which approximates to that of 
Meguptera, but has the greater part of the cavity filled by bone. 
There is a talon to the condyle of the mandible. 

Paleocetus, as-already mentioned (p. 232), is founded upon the 
ankylosed cervical vertebre of a small Whale originally considered as 
having been derived from the Kimeridge Clay, but which doubtless 
came from the Suffolk Crag; if it belongs to the Balenidw it indi- 
cates a Right Whale. 


1 See P. J. Van Beneden, ‘‘ Histoire Naturelles des Balénoptéres,” A/ém. Acad. 
Belgique, xli. 1887, 


246 CETACEA 


Suborder’ ARCHAOCETI. 


Family ZEUGLODONTIDA. 


This group is formed to include certain extinct Cetacean-like 
animals at present only known by more or less fragmentary por- 
tions of their skeleton and teeth, and whose position and affinities 
are, therefore, still subject to doubt.1 

In the anterior part of both jaws the teeth are simple, conical, 
or slightly compressed, and sharp pointed. The first three in the 
upper jaw are distinctly implanted in the premaxillary bone, and 
so may be reckoned as incisors. The tooth which succeeds, or the 
canine, is also simple and conical, but it does not exceed the others 
in size. This is followed by five teeth having two distinct roots 
and compressed pointed crowns, with denticulated cutting-edges. 
The dentition is therefore ¢ 3, ¢ 4, p and m & = 36, resembling that 
of some Seals.? General form of the skull elongated and much 
depressed. Brain-cavity very small, and the skull between it and 
the orbits elongated and narrow. Temporal fosse very large. A 
strong sagittal crest. Rostrum long and narrow, differing from 
that of other Cetaceans in the large extent to which the premaxille 
form the sides of the anterior extremity. Nasal bones elongated, 
flat, and narrow, the opening of the anterior nares being over the 
middle of the elongated compressed rostrum. All the cervical 
vertebree free. The characters of the dorsal vertebree and mode of 
articulation of the ribs appear to have resembled those of Platanista 
rather than Balena, Physeter, or Delphinus. Lumbar vertebrae 
with elongated bodies, low neural spines, and the transverse pro- 
cesses placed low down on the bodies. Characters of the limbs 
not known with certainty.? 

All the known fossil remains belonging to the animals of this 
group may be referred, provisionally at least, to the genus Zeuglodon, 
so named because the first section of a molar tooth examined was 
taken from the base of the crown, where it was beginning to divide 
into the two roots, and looked like two single teeth “linked or 


1 In a recent memoir Professor D’Arcy Thompson has brought forward some 
arguments to show that the Zeuglodonts have no direct affinities with the Cetacea, 
but have on the other hand the strongest possible relation with the Pinnipede 
Carnivora. ‘‘On the Systematic position of Zeuglodon,” Studies from the Museum 
of Zoology, Dundee, vol. i. No. 9, 1890. 

2 An appearance in one specimen has been described by C. G. Carus as in- 
dicating a vertical succession of the teeth, but the evidence upon which this rests 
is by no means satisfactory, and appears to admit of another explanation. 

* A mutilated humerus of Zeuglodon cctoides has given rise to many con- 
jectures, appearing to some anatomists to indicate seal-like freedom of motion 
at the elbow-joint, while to others its characters appear to be truly Cetacean. 


PHYSETERIDE 247 


yoked together.” This name was substituted by Owen for the 
earlier one Basilosaurus of Harlan, with the consent of that author, 
on the mammalian nature of the animal being demonstrated! The 
latter name is, however, still generally retained by American 
zoologists. The remains have hitherto been found chiefly in the 
Eocene formations of the States of Alabama, Louisiana, Mississippi, 
and Arkansas, and have been assigned to several species. A portion 
of a skull is recorded from the Barton Clay (Eocene) of Hampshire, 
England. 


Suborder ODONTOCETI, 
the DELPHINOIDEA, or Toothed Whales. 


Calcified teeth always present after birth; generally numerous, 
but sometimes a very limited number (in a few cases none) are 
functionally developed. No baleen. Upper surface of the skull 
more or less asymmetrical. Nasal bones in the form of nodules or 
flattened plates, applied closely to the frontals, and not forming 
any part of the roof to the narial passage, which is directed upwards 
and backwards. Olfactory organ rudimentary or absent. Hinder 
end of the maxilla expanded and covering the greater part of the 
orbital plate of the frontal bone. Lachrymal bone either inseparable 
from the jugal, or, when distinct, very large, and forming part of 
the roof of the orbit. Tympanic bone not ankylosed with the 
periotic, which is usually only attached to the rest of the skull by 
ligament. Rami of mandible nearly straight, much expanded in 
height posteriorly, with a wide funnel-shaped aperture to the dental 
canal, and coming in contact in front by a flat surface of variable 
length, but always constituting a true symphysis. Several of the 
anterior ribs with well-developed capitular processes, articulating 
with the bodies of the vertebre. Sternum almost always composed 
of several pieces, placed one behind the other, with which several 
pairs of ribs are always connected by the intervention of well- 
developed cartilaginous or ossified sternal ribs. External respiratory 
aperture single, the two nostrils uniting before they reach the 
surface, usually in the form of a transverse subcrescentic valvular 
aperture, situated on the top of the head. Manus always penta- 
dactylous, though the first and fifth digits are usually very little 
developed. No caecum, except in Platanista. 


Family PHYSETERIDA. 


No functional teeth in the upper jaw. Mandibular teeth various, 
often much reduced in number. Bones of the cranium raised so as 


1 See Trans. Geol. Soc. ser. 2, vol. vi. p. 67. 


248 CETACEA 


to form an elevated prominence or crest behind the nares. Pterygoid 
bones thick, produced backwards, meeting in the middle line, and 
not involuted to form the outer wall of the post-palatine air-sinuses, 
but simply hollowed on their outer side. Anterior facet of periotic 
bone (Fig. 87) for articulation with the tympanic quite smooth ; 
and the posterior tympanic surface of the former broad, with a 
median longitudinal ridge. Transverse processes of the arches of 
the dorsal vertebra, to which the tubercles of the ribs are attached, 
ceasing abruptly near the end of the series, and replaced by 
processes on the body at a much lower level, and not on a line or 
serially homologous with them, but serially homologous anteriorly 
with the heads of the ribs, and posteriorly with the transverse 
processes of the lumbar vertebra. (In some genera, as Physeter, 
the two processes, upper and lower on each side, are both present 
and well developed in the same vertebra in the region of transition. 
In others, as Ziphius and Berardius, they are not both developed on 
any single vertebra.) Costal cartilages not ossified. 

Subfamily Physeterine.—Numerous teeth in the mandible, 
which are not set in distinct bony alveoli, but in a long groove 
imperfectly divided by partial septa, and held in place by the 
strong, fibrous gum surrounding them. No distinct lachrymal bone. 
Cranium strikingly asymmetrical in the region of the narial 
apertures, in consequence of the left opening greatly exceeding the 
right in size. 

Physeter.—Upper teeth apparently of uncertain number, rudi- 
mentary, and functionless, being embedded in the gum. Lower jaw 


Fia. $2.—Skull of Sperm Whale (Physeter macrocephalus), 


with from 20 to 25 teeth on each side, stout, conical, recurved, and 
pointed at the apex until they are worn, without enamel. Upper 
surface of the cranium concave; its posterior and lateral edoes 
raised into a very high and greatly compressed semicircular crest 
or wall, Zygomatic processes of jugal bones thick and massive. 
Rostrum greatly elongated, broad at the base, and gradually tapering 


? Linn. Syst, Nat, 12th ed. vol. i. p. 107 (1766). 


PHYVSETERID.A£ 249 


to the apex. Upper edge of the mesethmoid forming a roughened 
irregular projection between the narial apertures, inclining to 
the left side. Mandible exceedingly long and narrow, the 
symphysis being more than half the length of the ramus. Vertebree: 
C7,D 11, L 8, C 24; total 50. Atlas free; all the other cervical 
vertebre united by their bodies and spines into a single mass. 
Eleventh pair of ribs rudimentary. Head about one-third the 
length of the body; very massive, high and truncated, and rather 
compressed in front; owing its huge size and remarkable form 
mainly to the accumulation of an oily substance secreted by 
the lining membranes of great cells surrounding the narial passage 
and filling the large hollow on the upper surface of the cranium 
and overlying the rostrum. The single blowhole is longitudinal, 
slightly sigmoid, and placed at the upper and anterior extremity 
of the head to the left side of the middle line. The opening 
of the mouth is on the under side of the head, considerably behind 
the end of the snout. Pectoral fin short, broad, and obliquely 
truncated. Dorsal fin a mere low protuberance. 

The only representative of this genus is the Cachalot or Sperm 


Fic. 83.—The Sperm Whale (Physeter macrocephalus). 


Whale (P. macrocephalus, Fig. 83), one of the most colossal of 
animals, quite equalling, if not exceeding, the Greenland Whale 
in bulk. The length of the full-grown male is from 55 to 
60 feet, but the female is stated not to reach more than half 
that size. The general colour of the surface is black above and 
gray below, the colours gradually shading into each other. — The 
Sperm Whale is one of the most widely distributed of animals, 
being met with usually in herds or “schools ” in almost all 
tropical and subtropical seas, but not occurring, except accident- 
ally, in the Polar regions. Not unfrequently specimens appear 
on the coasts of the British Isles, but only as solitary stragglers, 
or as dead carcases, floated northwards by the Gulf Stream. It 
is remarkable that every one of these of which we have an accurate 
record has been an old male. The food of this Whale consists 
mainly of various species of cephalopods (squid and cuttle-fish), 
but fish of considerable size are also eaten. The substance called 
“ambergris,” formerly used in medicine and now in perfumery, 
is a concretion formed in the intestine of this Whale, and is found 


250 CETACEA 


floating on the surface of the seas it inhabits. Its genuineness is 
proved by the presence of the horny beaks of the cephalopods on 
which the Whale feeds. 

The oil contained in the great cavity above the skull, when re- 
fined, yields “spermaceti,” and the thick covering of blubber which 
everywhere envelops the body produces the valuable “sperm-oil ” 
of commerce; hence this animal has long been the subject of a 
regular chase, by which its numbers have been greatly diminished. 

Cogia.1—Teeth of the upper jaw absent, or reduced to a rudiment- 
ary pair in front; in the lower jaw 9 to 12 on each side, rather long, 
slender, pointed, and curved, with a coating of enamel. Upper 
surface of the cranium concave, with thick, raised posterior and 
lateral margins, massive and rounded at their anterior terminations 
above the orbits. Upper edge of the mesethmoid forming a pro- 
minent sinuous ridge, constituting a kind of longitudinal septum 
to the base of the great supra-cranial cavity. Rostrum not longer 
than the cranial portion of the skull, broad at the base, and rapidly 
tapering to the apex. Zygomatic process of the jugal styliform. 
Mandible with the symphysis less than half the length of the entire 
ramus. Vertebre: C 7, D 13 or 14, L and C 30; total 50 or 51. 
All the cervical vertebre united by their bodies and arches. Ex- 
ternal characters not well known, but, judging by the somewhat 
conflicting accounts of those that have had an opportunity of ob- 
serving them, the head is about one-sixth of the length of the body, 
and obtusely pointed in front; the mouth small, and placed far 
below the apex of the snout; the spiracle crescentic, and placed 
obliquely on the top of the head anteriorly to the eyes, and to the 
left of the middle line ; the pectoral fins are obtusely falcate ; and 
there is a triangular dorsal fin. 

The history of this genus is a good illustration of the difficulties 
in which the study of the Cetacea has been involved by the super- 
ficial manner in which it has been investigated. The first known 
example, a skull from the Cape of Good Hope in the Paris Museum, 
was described by De Blainville under the name of Physeter breviceps. 
This was afterwards with good reason generically separated by Gray. 
Until within a very few years ago only five other individuals had 
been met with, each of which had been described under a different 
specific name (viz. grayi, macleayi, simus, floweri, and potsit), and 
which are arranged by Gray in two distinct genera. The most 
careful examination of the description given of these specimens, or 
of the now numerous osteological remains available, fails to detect 
any differences beyond those which may be attributed to age or sex, 
and hence, according to our present knowledge, these six supposed 
species must all be included under one name, C. lericeps. This 
animal appears to attain the length of 10 feet when adult, and has 


? Gray, Zoology of Erebus and Terror, p. 22 (1846). Usually spelt Kogia. 


PHYSETERIDAZA 251 


been met with at various distant localities in the Southern Ocean, 
and also off the coast of Madras and in the North Pacific. 

Extinct Physeteroids.—Teeth of Physeteroids are of very common 
occurrence in the Belgian and English Crags, and evidently indicate 
the former existence of Whales more or less closely allied to the 
Sperm Whale, but often distinguished by the presence of an enamel- 
cap on the crowns of the teeth. The generic determination of these 
teeth is, however, exceedingly difficult, owing to the water-worn 
condition in which they are frequently found, and also on account 
of the impossibility of knowing whether small and large teeth may 
not be referable to different parts of the jaws of the same species 
or to individuals of different ages. Moreover, in the cases of 
isolated teeth it is impossible to know how many were contained 
in the jaws, and therefore to distinguish Physeteroid from Ziphioid 
teeth. Physeterula is a small form about one-third the dimensions 
of the Sperm Whale, and distinguished by the length of the mandib- 
ular symphysis being only about one-third that of the entire ramus ; 
it is identified by Professor Cope with Cogia. Eucetus (Dinoziphius) is 
founded on teeth which are regarded as closely resembling those of 
Physeter, but distinguished by their subcylindrical form and the 
small size of the aperture of the pulp-cavity. It does not appear, 
however, to be certain that these teeth are not worn specimens of 
those described as Scaldicetus. Physetodon, from the Pliocene of 
Australia, is founded upon the evidence of similar teeth. The teeth 
from the Belgian Crag described as Scaldicetus are somewhat smaller 
than those of the Sperm Whale, and are readily characterised by 
their cap of grooved enamel. Other teeth with enamel-caps have 
been described as Physodon and Hoplocetus. The genus Balcenodon 
is founded upon a very imperfect large tooth from the English Crag, 
which is not sufficiently well preserved to admit of exact comparison 
with the other types. 

Subfamily Ziphiinee.—Teeth of the mandible (at least in existing 
forms) quite rudimentary and concealed in the gum, except one, or 
very rarely two, pairs which may be largely developed, especially 
in the male sex. A distinct lachrymal bone. Externally the mouth 
is produced into a slender rostrum or beak, from above which the 
rounded eminence formed by a cushion of fat resting on the cranium 
in front of the blowhole rises somewhat abruptly. Spiracle or 
blowhole single, crescentic, median, as in the Delphinidw. Pectoral 
fin small, ovate, the five digits all moderately well developed. A 
small obtusely falcate dorsal fin situated considerably behind the 
middle of the back. Longitudinal grooves on each side of the skin 
of the throat, diverging posteriorly, and nearly meeting in front. 
In external characters and habits the animals of this group closely 
resemble each other. They appear to be almost exclusively feeders 
on various species of cephalopods, and occur either singly, in pairs, 


252 CETACEA 


or in small herds. By their dental and osteological characters they 
are easily separated into four distinct genera. 

Hyperotdon.1—A_ small conical pointed tooth at the apex of each 
ramus of the mandible, concealed by the gum during life. Skull 
with the upper ends of the premaxille rising suddenly behind the 
nares to the vertex and expanded laterally, their outer edges 
curving backwards and their anterior surfaces arching forwards and 
overhanging the nares; the right larger than the left. Nasal bones 
lying in the hollow between the upper extremities of the premaxille, 
strongly concave in the middle line and in front; their outer edges, 
especially on the right side, expanded over the front of the inner 
border of the maxilla) Very high longitudinal crests on the 
maxille at the base of the rostrum, extending backwards almost to 
the nares, approaching each other in the middle line above; some- 
times so massive that their inner edges come almost in contact. 
Anteorbital notch distinct. Mesethmoid but slightly ossified. 
Vertebre: C 7, D 9, L 10, C 19; total 45. All the cervical 
vertebree united. Upper surface of the head in front of the blow- 


Fic. 84.—Hyperoddon rostratus. From a female specimen taken off the coast of Scotland, 1882. 


hole very prominent and rounded, rising abruptly from above the 
small, distinct snout. 

The genus is known typically by H. rostratus (Fig. 84), but an 
imperfect skull has been made the type of H. planifrons—a species 
differing considerably in cranial characters from the typical one. 
The females and young males of the first-named species have the 
contour of the head of the same general form as in Fig. 84; the 
premaxillary crests of the cranium being widely separated from 
one another, and terminating in comparatively sharp edges. In the 
males, however, as age advances the summits of these crests become 
gradually expanded and flattened, till they are almost or quite in 
contact in the middle line. This development of the maxillary 
crests produces a corresponding elevation and flattening of the front 
of the head, so that in very old males this aspect presents a flattened 
disc -like surface rising abruptly from the beak (which thus 
becomes almost buried) and situated in a plane nearly at right angles 
to the line of the back.” So different, indeed, is the appearance of 
the skull of an old male from that of a female individual that 


1 Lacépéede, ‘Table des Ordres,” Hist. Nut. des Cetacés, p. xliv. (1804). 
* See the figures in the Proc. Zool. Soc. 1882, pp. 728, 729. 


PHYSETERID.E 253 


it was long considered that they belonged to different species— 
the male form having been described as H. lutifrons. The length 
of an adult male reaches 30 feet, while that of the female does not 
exceed 24 feet. 

The Hyperoéddon, sometimes called “Bottlenose,” a name also 
vaguely given to several species of Dolphin, is a regular inhabitant 
of the North Atlantic, passing the summer in the Spitzbergen seas 
and going farther south in winter. It resembles the Sperm Whale 
in possessing a large store of oil in the upper part of the head, 
which yields spermaceti when refined; on this account, and also 
for the sake of the blubber, which supplies an oil almost indis- 
tinguishable from sperm-oil, this Whale has been the object of a 
regular chase in recent years. 

The following account of its habits is taken from a paper 
by Captain D. Gray, published in the Zoological Suciety’s Proceedings 
for 1882 :— 

“These Whales are occasionally met with immediately after 
leaving the Shetland Isles in March, and north across the ocean 
until the ice is reached, near the margin of which they are found 
in the greatest numbers; but they are seldom seen amongst it. 
Although it is not in their nature to keep in amongst the ice, they 
like to frequent the open bays for the shelter it gives them from 
the sea. Sometimes a point of ice overlaps them; it is then only 
that they are seen going out again towards the ocean. They are 
also to be met with from the entrance of Hudson’s Straits and up 
Davis’s Straits, as far as 70° N. lat., and down the east side 
round Cape Farewell, all round Iceland, north along the Greenland 
ice to 77° N. lat.; also along the west coast of Spitzbergen, 
and east to Cherry Island in lat. 72° N. and long. 19° E. Beyond 
these limits I have never seen them; but doubtless they are to be 
found as far as the Straits of Belle Isle on the west, and east to 
Nova Zembla. From the fact that they are not seen in summer 
farther south than a day’s sail from the ice, it would appear that 
they migrate south in the autumn, and north again in the spring. 
They are gregarious in their habits, going in herds of from four 
to ten. It is rare to see more than the latter number together, 
although many different herds are frequently in sight at the same 
time. The adult males very often go by themselves; but young 
bulls, cows, and calves, with an old male as a leader, are sometimes 
seen together. They are very unsuspicious, coming close alongside 
the ship, round about underneath the boats, until their curiosity 
is satisfied. . . . They vary in colour from black in the young to 
light brown in the older animals. The very old turn almost yellow, 
the beak and front of the head being quite white, with a white 
band round their necks ; all of them are grayish-white on the belly. 
They can leap many feet out of the water, even having time while 


234 CETACEA 


in the air to turn round their heads and look about them, taking 
the water head first, and not falling helplessly into it sideways like 
the larger whales. The full-grown whale is 30 feet long by 20 
feet in circumference, and vields two tons of oil besides two hundred- 
weight of spermaceti. . . . Their ordinary food consists of a bluish- 
white cuttle-fish, six inches long by three inches in circumference, 
and pointed towards the tail... . They evidently have a great 
depth to go to find them, judging from the length of time that 
they remain away, and from the long heavy blasts they make on 
coming to the surface again.” 

Periotic bones of Hyp roédon are found in the Red Crag of 
Suffolk, presenting no character by which they can be specifically 
distinguished from those of the common existing species. 

Ziphius.i—A single conical tooth of moderate size on each side 
of the mandible close to the anterior extremity, and directed 
forwards and upwards. Skull with the premaxille immediately in 
front of, and at the sides of the nares expanded, hollowed, and with 
elevated lateral margins, the posterior ends rising to the vertex and 
curving forwards, the right being considerably more developed than 
the left; the conjoint nasals forming a strongly pronounced sym- 
metrical eminence at the top of the cranium, projecting forwards 
over the nares, flat above, most prominent and rounded in the 
middle line in front, and separated by a notch on each side from 
the premaxille. Anteorbital notch not distinct. Rostrum (seen 
from above) triangular, gradually tapering from the base to the 
apex; upper and outer edges of maxille at base of rostrum raised 
into low roughened tuberosities. Mesethmoid cartilage densely 
ossified in adult age, and coalescing with the surrounding bones of 
the rostrum. Vertebre: C7,D10,L10, C22; total 49. The 
three anterior cervical vertebrxe united, the rest free. 

The type of this genus is Z. cavirestris of Cuvier, founded upon 
an imperfect skull picked up in 1804 on the Mediterranean coast 
of France, and described and figured in the Ossemens Fossiles under 
the impression that it was that of an extinct species. Many other 
individuals have, however, been subsequently met with in various 
parts of the world, from the Shetland Islands to New Zealand, all 
referable to the same genus, if not to the same species ; although, 
as is usual in such cases, they have mostly been described under 
different names, the so-called genera Petrorhynchus and E'piodon 
being probably referable to the type species. 

It is quite probable that some of the Physcteroid teeth from the 
Crag deposits mentioned on p. 251 may be referable to Ziphius, 

Mesoplodon.-—A much compressed and pointed tooth in each 


Cuvier, Ossemens Fossiles, 2d ed. vol. v. p, 352 (1828). 
* Gervais, «fru. Sei. Nad. sér, 3, vol. xiv, p. 16 (1850). For the very com- 
plicated synonymy of this genus, see Trans. Zool. Sov. vol. viii. p. 208, 


PHYSETERIDE 255 


ramus of the mandible, variously situated, but generally at some 
distance behind the apex (Fig. 86); its point directed upwards, and 
often somewhat backwards, “occasionally developed to a great size. 


Fic, 85.—Mesoplodon didens. From Reinharat. 


Skull with the region around the nares as in Hyperoddon, except 
that the nasals are narrow and more sunk between the upper ends 
of the premaxille; like those of Hyperoddon, they are concave in 
the middle line in front and above. No maxillary tuberosities. 
Anteorbital notch not very distinct. Rostrum long, narrow, and 
solid throughout. Mesethmoid in adult age ossified in its entire 


Fic. $6.—Left lateral view of skull of Mesoplodon densirostris. 


length, coalescing with the surrounding bones, and showing as a 
narrow band on the upper surface of the rostrum. V ertebree : 
C7, D10,L10 or 11, C19 or 20; total 46 to 48. Two or three 
anterior cervicals united, the rest usually free. 

Though varying in form, the mandibular teeth of the different 
members “of this genus agree in their essential structure, having a 
small and pointed enamel- covered crown, composed of true dentine, 
which, instead of surmounting a root of the ordinary character, is 
raised upon a solid mass of osteodentine. The continuous growth of 
this greatly alters the form and general appearance of the organ 
as age advances, as seen most strikingly i in the case of JI. layardi, 
where the long, narrow, flat, strap-like teeth, curving inwards at 
their extremities, actually meet over the rostrum, and must greatly 
interfere with the movements of the jaw. In one species (1/. vvuyi) 
a row of minute, conical, pointed teeth, like those of ordinary 
Dolphins, 17 to 19 in number, are present eyen in the adults, on 


256 CETACEA 


each side of the middle part of the upper jaw, but embedded by 
their roots only in the gum, and not in bony alveoli. This fact, 
with the frequent presence of rudimentary teeth in other species 
of this and the last genus in both upper and lower jaws, 
suggests the idea that the Ziphioids are derived from ancestral forms 
which had teeth of normal character in both jaws; the dentition 
of the living forms having become greatly specialised. The existing 
species of this genus are widely distributed in both northern and 
southern hemispheres, but most frequent in the latter. The best 
established are Jf. bidens, M. ewropeus, M. densirostris, M. layardi, 
M. gray, and M. hectori ; but there is still much to be learned with 
regard to their distinctive characters and geographical distribution. 
They were abundant in the Pliocene age, as attested by the fre- 
quency with which the most im- 
perishable and easily recognised 
portion of their structure, the 
long, cylindrical rostrum of the 
skull, of more than ivory dense- 
ness, is found among the rolled 
and water-worn fragments of 
animal remains which compose 
Fig. 87.—The left periotic bone of Meso. the well-known ‘bone-bed” at 


plodon; from the Red Crag of Suffolk. The 
smooth concave surface in the right upper the base of the Red Crag of Suf- 


corner of the figure forms the anterior ar- folk. Several species have been 
ticulation with the tympanic. (From the foynded upon the evidence of 
Cat. Foss. Mamm. Brit. Mus. pt. v. p. 70.) these rostra. Periotic bones of 
this genus (Fig. 87) are of less common occurrence in the Crag ; 
the figure is given to illustrate the characteristic features of this 
bone in the present family. 

Berardius.|—Two moderate-sized, compressed, pointed teeth on 
each side of the symphysis of the mandible, with their apices directed 
forwards, the anterior being the larger of the two and close to the 
apex. Upper ends of the premaxille nearly symmetrical, moder- 
ately elevated, very slightly expanded, and not curved forward over 
the nares. Nasals broad, massive, and rounded, of nearly equal 
size, forming the vertex of the skull, flattened in front, most 
prominent in the middle line. Anteorbital notch distinct. Rostrum 
long and narrow. Mesethmoid only partially ossified. Small 
rugous eminences on the outer edge of the upper surface of the 
maxillz at base of rostrum. Vertebrate: C 7, D 10,L 12,019; 
total 48. The three anterior cervicals ankylosed, the rest free and 
well developed. 

The only known species, B. arnua?, attains the length of 30 
feet, and has hitherto only been met with in the seas around New 
Zealand. 


1 Duvernoy, Ann. Sct. Nat.-Zoologic, sér. 3, vol. xv. p. 41 (1851). 


SQUALODONTID.£E 257 


Choneziphius4—The rostral portions of crania from the Antwerp 
and Sutfolk Crags, on the evidence of which this genus has been 
established, agree with those of JJesoplodon in having the premaxille 
in contact with the intervening bones throughout the length of 
their inner surfaces, and also in showing only a very small portion 
of the vomer on the inferior surface ; they differ, however, in that 
the mesethmoid cartilage remains unossified, whereby a fistular 
vacuity remains. In some species the soldering of the inner 
surfaces of the premaxille is incomplete. The interorbital region 
of the skull is flat ; and there are two pits in the nasal region, of 
which the right is the larger. 


Fumily SQUALODONTID-E. 


Numerous extinct forms, chiefly known by teeth and fragments 
of cramia, may be provisionally placed here, until more of ‘their 
osteological characters shall be brought to light. They differ from 
all existing Cetaceans in having the teeth distinctly differentiated 
into groups, as in the Archzoceti, the posterior molars being two- 
rooted. The cranium has, however, none of the distinguishing 
characteristics of the Zeuglodonts, but essentially resembles that of 
the Odontoceti, especially in the position of the anterior nares and 
form of the nasal bones. 

Syuetlodon?—All the forms may be included in this genus, the 
so-called Lizoprion not being distinct. Dentition: 7 3, ¢ 7, simple 
teeth of the molar series (premolars ?) 4, two-rooted molars #=14 ; 
total 60. The double-rooted molars differ from those of Zenglodon 
in having the denticulations of the crown confined to the posterior 
border, or at all events much less developed on the front edge. 
Very little is known of the structure of these animals beyond the 
skull and teeth, fragments of which have been found widely 
distributed throughout the marine Miocene and early Pliocene 
formations of Europe, especially in the Vienna basin, many parts 
of France, and the Antwerp and Suffolk Crags. They have also 
been found in formations of corresponding age in North America 
and South Australia. A few isolated teeth have been met with in 
the cave-deposits of Italy, which, if contemporancous with the beds 
in which they occur, indicate the survival of the genus into the 
Pleistocene period. 


Family PLATANISTID-£. 
Under this heading may be placed three very singular genera, 
which, though differing considerably from each other, have several 


1 Duvernoy, op. cif. p. 61. 
2 Grateloup, lef. fe. 2. Sef. Bordeana, 1840, p. 208. 
17 


258 CETACEA 


points in common, and do not altogether come under the definition 
either of the Physeteride or the Delphinide, especially in the 
important character of the mode of articulation of the ribs with 
the dorsal vertebra, the tubercular and capitular articulations, 
distinct at the commencement of the series, gradually blending 
together, as they do in most ordinary mammals. The cervical 
vertebree are all free. The lachrymal bone is not distinct from the 
jugal. The jaws are long and narrow, with numerous teeth in 
both. The symphysis of the mandible exceeds half the length of 
the whole ramus. Externally the head is divided from the body 
by a slightly constricted neck. Pectoral limbs broad and truncated. 
Dorsal fin small or obsolete. Fluviatile or estuarine in habits. 
There are three distinct genera, which might almost be made the 
types of families, but it is probably more convenient to keep them 
together, only regarding them as representing three subfamilies. 

Platanista..—Teeth about $9 on each side, set near together, 
rather large, cylindrical, and sharp-pointed in the young; in old 
animals acquiring a large laterally compressed base, which in the 
posterior part of the series becomes irregularly divided into roots. 
As the conical enamel-covered crown wears away, the teeth of the 
young and old animals have a totally different appearance. The 
rostrum and dentigerous portion of the mandible are so narrow 
that the teeth of the two sides are almost in contact. Maxille sup- 
porting very large, incurved, compressed bony crests, which over- 
arch the nares and base of the rostrum, and almost meet in the 
middle line above. Orbits very small and eyes rudimentary, without 
crystalline lens, External respiratory aperture longitudinal, linear. 
Vertebre: C 7, D 10, L 9,C 26; total 52. A small cecum. No 
pelvic bones. Dorsal fin represented by a low ridge. 

One species, P. gangetica, entirely fluviatile, being extensively 
distributed throughout nearly the whole of the river systems, not 


Fic. 88.—Platanista gangetica, (From Anderson.) 


only of the Ganges, but of the Brahmaputra and Indus, ascending 
as high as there is water enough to swim in, but never passing out 
to sea, It is quite blind, and feeds on small fish and crustaceans, 
groping for them with its long snout in the muddy water at the 
bottom of the rivers. It attains the length of 8 feet.2 

1 Wagler, Syst. iphib. ete., p. 85 (1830). 

* The anatomy of Platanista is fully described by J. Anderson, Zoological 
Results of Two Expeditions to Western Yunnan, 1878. 


PLATANISTIDE 259 


Inia.\—Teeth variable, from 26 to 33 on either side of each jaw; 
those at the posterior part with a distinct tubercle at the inner side 
of the base of the crown. Vertebre: C 7, D 13, L 3, C 18; total 
41. Transverse processes of lumbar vertebra very broad. Sternum 
short and broad, and consisting of a single segment only. Dorsal 
fina mere ridge. The long cylindrical rostrum externally furnished 
with scattered, stout, and crisp hairs. One species only is known, 
L. geoffroyensis, about 7 feet in length, inhabiting the upper Amazon 
and its tributary streams. 

Pontoporia.2—Teeth 50 to 60 on either side of each jaw, with a 
cingulum at the base of the crown. Jaws very long and slender. 
Vertebre: C 7,D 10, L 5, C19; total 41. Transverse processes 
of the lumbar vertebrae extremely broad. Sternum elongated, 
composed of two segments, with four sternal ribs attached. Dorsal 
fin rather small, triangular, pointed. External respiratory aperture 


Fia. 89.—Pontoporia blainvillei. (From Burmeister.) 


transverse, crescentic. This genus connects the last two forms with 
the true Delphinide. The only species, P. blainvillei, is one of the 
smallest members of the whole order, not exceeding 5 feet in length. 
It has only been met with at the mouth of the Rio de la Plata, near 
Buenos Ayres, and there is at present no evidence that it ascends 
into the fresh waters of the river. 

Fossil forms.—Remains of a Cetacean from the Pleistocene of 
South America were referred by Bravard to Pontoporia, but they 
have been regarded by other writers as indicating a distinct genus, 
for which the names Palwopontoporia and Pontistes have been pro- 
posed. The Upper Tertiary European genera Champsodelphis and 
Schizodelphis are generally referred to the present family. The 
former has wide transverse processes to the lumbar vertebrae, as in 
Inia, while the teeth also resemble those of that genus. In Schizo- 
delphis the form of the rostrum presents a great resemblance to that 
of the Delphinoid genus Steno, but the symphysis of the mandible 
is relatively longer. A number of fossil Cetaceans from the 
Miocene of the United States, such as Priscodelphinus, Lophocetus, 
Txacunthus, Rhabdosteus, etc., are referred by Professor E. D. Cope to 


1 D'Orbigny, Nouv. Ann. Mus. Paris, vol. iii, p. 31 (1834). 
2 Gray, Zoology of Erebus and Terror, p. 46 (1846). 


260 CETACEA 


this family. Agabdelus, from the same deposits, is an apparently 
allied, but toothless type. 


Family DELPHINID.£. 


Teeth usually numerous in both jaws. Pterygoid bones short, 
thin, each involuted to form with a process of the palate bone the 
outer wall of the post-palatine air-sinus. Symphysis of mandible 
short, or moderate, never exceeding one-third of the length of the 
ramus. Lachrymal bone not distinct from the jugal. The anterior 
facet on the periotic (Fig. 96) for articulation with the tympanic 
deeply grooved ; and the posterior tympanic surface of the same 
bone comparatively narrow, with its ridge for articulation with the 
free border of the tympanic ill-defined, and situated close to one 
edge. Transverse processes of the dorsal vertebre gradually trans- 
ferred from the arches to the bodies of the vertebre without any 
sudden break, and becoming posteriorly continuous serially with the 
transverse processes of the lumbar vertebra. Anterior ribs attached 
to the transverse process by the tubercle, and to the body of the 
vertebra by the head; the latter attachment lost in the posterior 
ribs. Sternal ribs firmly ossified. External respiratory aperture 
transverse, crescentic, with the horns of the crescent pointing» 
forwards. 

A very large group, closely united in essential characters but 
presenting great modifications in details. The different types are 
mostly so connected by intermediate or osculant forms that there 
are great difficulties in grouping them into natural subfamilies. 
Even the formation of well-defined genera is by no means satis- 
factory in all cases. They may, however, be divided, perhaps 
artificially, into two groups. 

Group A.—Head rounded, without distinct rostrum or beak. 
Rostrum of skull about as long as cranial portion. 

Monodon.'— Besides some irregular rudimentary teeth, the entire 
dentition is reduced to a single pair of teeth which lie horizontally 
in the maxilla, and in the female remain permanently concealed 
within the alveolus, so that this sex is practically toothless, while 
in the male (see Fig. 90) the right tooth usually remains similarly 
concealed and abortive, and the left is immensely developed, attaining 
a length equal to more than half that of the entire animal, projecting 
horizontally from the head in the form of a cylindrical, or slightly 
tapering, pointed tusk, without enamel, and with the surface 
marked by spiral grooves and ridges, running in a sinistral direction, 
(When, as occasionally happens, both tusks are developed, the 
spiral grooves have the same direction in each.) Pterygoids very 


1 Linn. Syst. Nat. 12th ed. vol. i. p. 105 (1766). 


DELPHINIDA 


261 


small, not meeting in the middle line, but approxi- 
mating posteriorly. Vertebre: C7, D 11, L6, 
C 26; total 50. Cervical region comparatively 
long, and all the vertebre distinct, or with ir- 
regular unions towards the middle of the series, 
the atlas and axis being usually free. Manus 
small, short, and broad; second and third digits 
nearly equal, fourth slightly shorter. No dorsal 
fin. 

This genus is now represented only by the 
well-known Narwhal (Jf. monoceros), in which the 
horn-like tusk of the male often grows to a 
length of 7 or 8 feet. In very young animals 
several small additional teeth, irregular in number 
and position, are present, but these usually dis- 
appear soon after birth. 

The head is rather short and rounded ; the 
fore limbs or paddles are small and broad com- 
pared with those of most Dolphins ; and (as in the 
Beluga) the median dorsal fin, found in nearly 
all other members of the group, is wanting or 
replaced by a low ridge. The general colour of 
the surface is dark gray above and white below, 
but variously marbled and spotted with different 
shades of gray. In the general contour of the 
body the Narwhal resembles the White Whale 
or Beluga. - 

The Narwhal is essentially an Arctic animal, 
frequenting the icy cireumpolar seas, and but 
rarely seen south of 65° N. lat. Three instances 
have, however, been recorded of its occurrence 
on the British coasts, one in the Firth of Forth 
in 1648, one near Boston in Lincolnshire in 1800, 
while a third, which entangled itself among 
the rocks in the Sound of Weesdale, Shetland, 
in September 1808, is described by Fleming 
in the Memoirs of the IWernerian Society, vol. i. 
Like most other Cetaceans, it is gregarious in 
its habits, being usually met with in “ schools ” 
or herds of fifteen or twenty individuals. Its 
food appears to be various species of cephalo- 
pods, small fishes, and crustaceans. The pur- 
pose served in the animal’s economy by the 
wonderfully developed asymmetrical tusk—or 
“horn,” as it is commonly but erroneously 
called —is not known. As it is present only 


Fra, 90.—Upper surface of the skull of male Narwhal (Monodon monoceros), with the whole of both teeth exposed 


by remoyal of the upper wall of their alveolar cavities. 


262 CETACEA 


in the male sex, no function essential to the well-being of the 
individual, such as the procuring of sustenance, can be assigned 
to it, but it must be looked upon as belonging to the same cate- 
gory of organs as the antlers of deer, and perhaps may be 
applied to similar purposes. Very little is, however, known of the 
habits of Narwhals. Scoresby describes them as “extremely 
playful, frequently elevating their horns and crossing them with 
each other as in fencing.” They have never been known to charge 
and pierce the bottom of ships with their weapons, as the swordfish 
often does. The name “Sea Unicorn,” sometimes applied to the 
Narwhal, refers to the resemblance of its tusk to the horn 
represented as projecting from the forehead of the fabled unicorn. 
The ivory of which the tusk is composed is of very good quality, 
but, owing to the central cavity, which extends the greater part of 
its length, is only fitted for the manufacture of objects of small 
size. The entire tusks are sometimes used for decorative purposes, 
and are of considerable, though very fluctuating, commercial value. 

Delphinapterus.A—This genus is closely allied to the last in ex- 
ternal form, as well as anatomical structure, differing mainly in the 
very distinct character of the dentition. Teeth from § to +9, 
occupying the anterior three-fourths of the rostrum and correspond- 
ing portion of the mandible, rather small, conical, and pointed 
-when unworn, but usually becoming obliquely truncated, separated 
by intervals considerably wider than the diameter of the tooth, and 
implanted obliquely, the crowns inclining forwards, especially in 
the upper jaw. Skull rather narrow and elongated, depressed. 
Premaxille convex in front of the nares. Rostrum about equal in 
length to the cranial portion of the skull, triangular, broad at the 
base, and gradually contracting towards the apex, where it is some- 
what curved downwards. Vertebre: C7,D 11, L 9, C 23; total 50. 
Cervical vertebra free. Manus broad, short, and rounded, all the 
digits being tolerably well developed, except the first. No dorsal 
fin, but a low ridge in its place. 


Fic. 91.—Beluga or White Whale (Delphinapterus leucas). From a specimen taken in the river 
St. Lawrence, and exhibited in London, 1877. 


One existing species, D. lewcas (Fig. 91), the Beluga or White 
Whale, so called from its pure white colour, about 12 feet long, 
abundant in the Arctic seas, and extending as far south on the 


1 Lacépéde, Hist. Nat. des Cétacés, p. xli. (1804). 


DELPHINIDA 263 


American coast as the river St. Lawrence, which it ascends for a 
considerable distance. On rare occasions it has been seen on the 
coast of Scotland. 

Remains of a Cetacean from the Lower Pliocene of Tuscany have 
been referred by Brandt to this genus under the name D. brocchit. 

In all the remaining genera of Delphinide the cervical region of 
the vertebral column is very short, and the first two, and usually 
more, of the vertebree are firmly united. 

Phocena,$—Teeth 25, small, occupying nearly the whole length 
of the rostrum, with compressed, spade-shaped crowns, separated 
from the root by a constricted 
neck (Fig. 92). Rostrum rather 
shorter than the cranium 
proper, broad at the base and 
tapering towards the apex. 
Premaxille raised into tuber- 
osities in front of the nares. 
The frontal bones forming a 
somewhat square, elevated pro- 
tuberance in the middle line of the skull behind the nares, rising 
altogether above the flattened nasals. Pterygoids very small, 
and widely separated in the middle line. Symphysis of mandible 
very short. Vertebre: C7, D13, L14,C31; total 65 (subject 
to slight individual variations). First to sixth cervical vertebra, 
and sometimes the seventh also, coalesced. Manus of moderate 
size, oval, slightly falcate ; second and third digits nearly equal in 
length ; fourth and fifth well developed, but shorter. Dorsal fin 
near the middle of the back, triangular ; its height considerably less 
than the length of the base ; its anterior edge frequently furnished 
with one or more rows of conical horny tubercles. 

The common Porpoise (Fig. 93), P. communis, is the best known 
of British Cetaceans. The word Porpoise (sometimes spelled Porpus 
and Porpesse) is apparently derived from the French pore and 
poisson, or the Italian porco and pesce, and thus corresponds with 
some of the English vernacular appellations, “ hog-fish,” “ sea-hog,” 
“herring-hog,” and the German Meerschwein, whence the usual modern 
French name of the animal, marsowin. ‘“ Porpoise” is commonly used 
by sailors to designate all the smaller Cetaceans, especially those 
numerous species which naturalists call “Dolphins ”; but in scientific 
language it is restricted to the genus Phocwna of Cuvier, of which the 
Porpoise of the British seas, Phocwna communis, Cuvier (Delphinus 
phocena, Linnus), is the type. 

The Common Porpoise, when full grown, attains a length of 5 
feet or a little more. The dimensions of an adult female specimen 
from the English Channel were as follows :—length in straight line 


1 Cuvier, Regne Animal, vol. i. p. 279 (1817). 


Fic, 92.—Teeth of Porpoise. Twice natural size. 


264 CETACEA 


from nose to median notch between the flukes of the tail, 624 
inches ; from the nose to the anterior edge of the dorsal fin, 29 
inches ; height of dorsal fin, 44 inches ; length of base of dorsal fin, 
8 inches ; length of pectoral fin, 9} inches ; breadth of pectoral fin, 
34 inches; breadth of tail flukes, 13 inches. The under jaw 
projects about half an inch beyond the upper one. The aperture — 
of the mouth is tolerably wide, and is bounded by stiff immobile 
lips, and curves slightly upwards at the hinder end. The eye is 
small, and the external ear represented by a minute aperture in the 
skin, scarcely larger than would be made by the puncture of a pin, 
situated about 2 inches behind the eye. The pectoral fins are of 


Fic. 93.—The Common Porpoise (Phocena communis). 


moderate size, and slightly faleate. The upper parts are dark gray, 
or nearly black, according to the light in which they are viewed, 
and the state of moisture or otherwise of the skin; the under parts 
are pure white. The line of demarcation between these colours is 
not distinct (washes or splashes of gray encroaching upon the 
white on the sides), and varies somewhat in different individuals, 
Usually it passes from the throat (the anterior part of which, with 
the whole of the under jaw, is dark) above the origin of the 
pectoral fin, along the middle of the flank, and descends again to 
the middle line before reaching the tail. Both sides of the pectoral 
and caudal fins are black. 

The Porpoise is sociable and gregarious in its habits, being usu- 
ally seen in small herds, and frequenting coasts, bays, and estuaries 
rather than the open ocean. It is the commonest Cetacean in the 
seas around the British Isles, and not unfrequently ascends the 


DELPAINIDA 265 


river Thames, having been seen as high up as Richmond; it has 
also been observed in the Seine at Neuilly, near Paris. It frequents 
the Scandinavian coasts, entering the Baltic in the summer ; and 
is found as far north as Baffin’s Bay, and as far west as the coasts 
of the United States. Southward its range is more limited than 
that of the Common Dolphin, as, though very common on the 
Atlantic coasts of France, it rarely enters the Mediterranean. 

It feeds on fish, such as mackerel, pilchards, and herrings, of 
which it devours large quantities, and, following the shoals, is often 
caught by fishermen in the nets along with its prey. In former 
times it was a common and esteemed article of food in England and 
in France, but is now rarely if ever eaten, being commercially 
valuable when caught only for the oil obtained from its blubber. 
Its skin is sometimes used for leather and boot-thongs, but 
the so-called “porpoise hides” are generally obtained from the 
Beluga. 

A closely similar if not identical species from the American 
coast of the North Pacific has been described under the name of 
Phocena vomerina, and another from the mouth of the Rio de la 
Plata as P. spinipennis. 

The stomach of the Porpoise (Fig. 94) may be taken as a typical 
example of this 
organ in the Ceta- 
cea. The first and 
by far the largest 
compartment ()) 
may be regarded 
as a kind of crop, 
or dilatation of 
the large ceso- 
phagus (a). It is 
lined by a thick 
white epithelium, 
which ceases 
abruptly at the 
entrance into the 
next cavity. It 
corresponds to 
the cardiac com- 
partment of the 
stomach in the 


Fic. 94.—Diagrammatic section of the stomach of the Porpoise. 
a, Esophagus ; 8, left, or cardiac, compartment ; ¢, middle compart- 
Ungulates and ment; d and e, the two divisions of the right, or pyloric, compart- 
certain Rodents 3 ment; f, pylorus; g, duodenum, dilated at its commencement; h, 


but, although its evel 


walls do not appear to contain peptic glands, its contents undergo 
partial digestion—probably caused by the regurgitation into it 


266 CETACEA 


of the secretions of the second, or true digestive compartment 
(c). This, which is much smaller than the first, has very thick 
walls, the mucous membrane being filled with numerous tubular 
glands. The surface of this membrane is smooth and soft, 
being thrown into numerous folds, which in this genus are arranged 
in a very peculiar and characteristic manner, so as to form a 
series of prominent longitudinal ridges, each of which sends off 
short lateral ridges at right angles to itself, which interdigitate 
with those proceeding from the next longitudinal ridge. The 
remainder of the stomach (d to f) may be compared to the pyloric 
antrum of the stomach of ordinary mammals. It is elongated, 
cylindrical, and intestiniform, with a smooth lining membrane, 
sharply bent upon itself, and terminating in a very small cir- 
cular pyloric aperture (f). In the Porpoise the commence- 
ment of this cavity is constricted off from the remainder, so as to 
form a small globular sac. In most Dolphins (as Tursiops, Globi- 
cephalus, and Grampus) there are two such small sacs of very similar 
size and form, communicating by circular pylorus-like apertures ; 
and in Hyperoddon the whole compartment is divided by a series of 
constrictions into as many as seven separate cavities, which have 
been regarded as distinct stomachs. Immediately beyond the 
pylorus the duodenum has a globular dilatation, as in the camels 
and some other Ungulates, into the lower end of which the biliary 
duct (h) enters. 

An allied species, differing mainly in the absence of dorsal fin, 
and in the teeth (with the same form of crown) being fewer in 
number and of larger size, called Delphinus phocenoides by Cuvier, 
D. melas by Schlegel, forms the type of Gray’s genus Neomeris.+ 
It is rather smaller than the Common Porpoise, and almost entirely 
black in colour. Common off the coast of Bombay, it has been 
met with in other parts of the Indian Ocean, and near Japan. 
The British Museum recently received a specimen taken in the 
Chinese river Yang-tse-kiang nearly a thousand miles from the 
sea, which only differs from others from India in wanting a patch 
of small horny tubercles on the back. As such tubercles are 
present or absent in otherwise similar individuals of P. communis, it 
is doubtful whether they can be regarded as constituting a specific 
character. 

Cephalorhynchus.’— Rostrum: as long and sometimes slightly 
longer than the cranial portion of the skull. Pterygoids widely 
separated from one another. Teeth small (less than 3 mm. in 
diameter), $3 to $3. Vertebre: C 7, D 13, L 15, C 30; total 65. 
Dorsal fin low, obtusely triangular or rounded. Pectoral fins rather 

1 Zoology of Erebus and Terror, p. 30 (1846). The name is preoccupied by 
Lamarck for a genus of Polyzoa (1816). 

2 Gray, Cat. Cetacea Brit. Mus. p. 106 (1850). 


DELPHINIDAE 267 


small, narrow, and ovate. Typified by C. heavisilvi, from the 
southern seas. C. eutropia is a very distinct form from the same 
seas, known only by the skull, and referred provisionally to this 
genus. 

Orcella..—Teeth 12 to 44, small, conical, pointed, rather closely 
set, and occupying nearly the whole length of the rostrum. Skull 
subglobular, high. Rostrum nearly equal in length to the cranial 
portion of the skull, tapering. Pterygoids widely separated from 
one another. Manus of moderate size, not elongated, but some- 
what pointed. All the bones of the digits broader than long, 
except the proximal phalanges of the index and third fingers. 
Dorsal fin rather small, placed behind the middle of the body. 
Two species, both of small size—0O. brevirostris, from the Bay of 
Bengal, and O. fluminalts, from the Irawadi river, from 300 to 
900 miles from the sea. Our present knowledge of the anatomy, 
geographical distribution, and habits of these interesting Cetaceans 
is almost entirely due to the researches of Dr. J. Anderson.” 

Orca2—Teeth about 12, occupying nearly the whole length of 
the rostrum, very large and stout, with conical recurved crowns, 
and large roots, expanded laterally and flattened, or rather hollowed, 
on the anterior and posterior surfaces. Rostrum about equal in 
length to the cranial part of the skull, broad and flattened above, 
rounded in front; premaxille broad and rather concave in front of 
the nares, contracted at the middle of the rostrum, and expanding 
again towards the apex. Pterygoids of normal form, but not quite 
meeting in the middle line. Vertebre: C 7, D 11-12, L 10, 
C 23; total 51 or 52. Bodies of the first and second and some- 
times the third cervical vertebre united; the rest free. Pectoral 
fin very large, ovate, nearly as broad as long. All the phalanges 
and metacarpals broader than long. General form of body robust. 
Dorsal fin near the middle of the back, very high and pointed. 
Anterior part of the head broad and depressed. 

The animals composing this genus are met with in almost all 
seas from Greenland to Tasmania, but the number of species is still 
uncertain, and possibly they may be all reduced to one. They are 
readily known, when swimming in the water, by the high, erect, 
faleate dorsal fin, whence their common German name of Schwert- 
fisch (Sword-fish). By English sailors they are generally known as 
“Grampuses” or “Killers.” They are distinguished from all their 
allies by their great strength and ferocity, being the only Cetaceans 
which habitually prey on warm-blooded animals, for, though fish 
form part of their food, they also attack and devour Seals, and 


1 Gray, Cat. Seals and Whales in Brit. Mus. p. 285 (1866). 

2 Anatomical and Zoological Researches, comprising an Account of the Zoological 
Results of the two Expeditions to Western Yunnan, tn 1868 and 1875 (1878). 

3 Gray, Zoology of Erebus and Terror, p. 33 (1846). 


268 CETACEA 


various species of their own order, not only the smaller Porpoises 
and Dolphins, but even full-sized Whales, which last they combine 
in packs to hunt down and destroy, as Wolves do the larger 
Ruminants. 


Fic, 95.--The Killer Whale, or Grampus (Orca gladiator). From Hunter. 


Orea citoniensis, of the Italian Pliocene, was of smaller size than 
the existing Killer. Teeth and periotic bones from the Suffolk Crag 
not improbably belong to the same species. 

Psewlorca.1—Teeth about +2. Cranial and dental characters 
generally like those of Orca, except that the roots of the teeth are 
cylindrical. Vertebre: C 7, D 10, L 9, C 24; total 50. First 
to sixth or seventh cervical vertebree united. Bodies of the lumbar 
vertebre distinguished from those of the preceding genera by being 
more elongated, the length being to the width as 3 to 2. Pectoral 
fin of moderate size, narrow, and pointed. Dorsal fin situated near 
the middle of the back, of moderate size, faleate. Head in front of 
the blowhole high, and compressed anteriorly, the snout truncated. 

This genus was first known by the discovery of a skull in a 
sub-fossil state in a fen in Lincolnshire, named by Sir R. Owen 
Phocena crassidens. Animals of apparently the same species were 
afterwards met with in small herds on the Danish coast, and fully 
described by Reinhardt. Others subsequently received from Tas- 
mania were supposed at first to indicate a different species, but 
comparison of a larger series of specimens from these extremely 
distant localities fails to establish any characteristic difference, and 
indicates an immense range of distribution for a species appar- 
ently so rare. The length of this Cetacean is about 14 feet, and 
its colour entirely black. 


Globicephalus.~—Teeth —, confined to the anterior half of the 


rostrum and corresponding part of the mandible, small, conical, 
curved, sharp-pointed when unworn, sometimes deciduous in old 
age. Skull broad and depressed. Rostrum and cranial portion 
about equal in length. Upper surface of rostrum broad and flat. 


' Reinhardt, Overs. Dan. Sezsk, Forh, 1862, p. 151, 
2 Lesson, V. Zab. d. Regne Animal—Mamm. p. 200 (1842). 


DELPHINIDE 269 


Premanillie strongly concave in front of the nares, as wide at the 
middle of the rostrum as at the base, or wider, and very nearly or 
completely concealing the maxille in the anterior half of this 
region. Pterygoids of normal form, meeting, or very nearly so, 
in the middle line. Vertebre: C 7, D 11, L 12-14, C 28-29; 
total 58 or 59. Bodies of the anterior five or six cervical vertebrae 
united. Length of the bodies of the lumbar and anterior caudal 
vertebrs about equal to their width. Pectoral limb very long and 
narrow, the second digit the longest, and having as many as 12 
or 13 phalanges, the third shorter (with 9 phalanges), the first, 
fourth, and fifth very short. Fore part of the head very round, in 
consequence of the great development of a cushion of fat, placed 
on the rostrum of the skull in front of the blowhole. Dorsal fin 
low and triangular, the length of its base considerably exceeding its 
vertical height. 

The type of this well-marked genus is G. melas, the Pilot 
Whale, Caing Whale, or Grindhval of the Faroe islanders, which 
attains the length of 20 feet, and is of nearly uniform black colour, 
except the middle of the under surface, which is lighter. These 
animals are extremely gregarious, and, unlike the Killers, are mild 
and inoffensive in disposition, feeding principally on cephalopods. 
Their eminently sociable character constantly leads to their destruc- 
tion, since when attacked they instinctively rush together and 
blindly follow the leaders of the herd. When they are seen in 
the neighbourhood of land, the fishermen endeavour to get to sea- 
ward of them in their boats, and with shouting and firing of guns 
to drive them into a bay or fjord, pursuing them until they run 
themselres on shore in their alarm. In this way many hundreds 
at a time are frequently driven ashore 
and killed, when a herd enters one of 
the bays or fjords of the Faroe Islands 
or north of Scotland. Animals of this 
well-marked genus are found in nearly 
all seas, and their specific distinctions 
are not yet made out. Specimens from 
the Australian coasts, where they are 


generally called “Blackfish,” are quite 
indistinguishable, either by external or 
osteological characters, from those of the 
North Atlantic. 

Teeth, periotic (Fig. 96) and tym- 
panic bones from the Suffolk Crag, 
described as G. uncidens, indicate a form 
apparently closely allied to the existing 
species. 


Fia. 96.—The left periotic bone 
of Globicephalus uncidens ; from the 
Suffolk Crag. Natural size. The 
grooved surface on the right is the 
anterior facet for articulation with 
the tympanic; the posterior tym- 
panic articulation being on the op- 
posite side of the figure. (From the 
Cat. Foss. Mumm. Brit, Mus. pt. v.) 


The periotic is figured in order to illustrate the dis- 


tinctive characters of that bone in the Delphinide. 


270 CETACEA 


Grampus..—Teeth none in the upper jaw; in the mandible few 
(3 to 7 on each side), and confined to the region of the symphysis. 
Vertebre: C7, D12, L19, C30; total 68. General external 
characters much as in Globicephalus, but the fore part of the head 
less rounded, and the pectoral fin less elongated. 

But one species, G. griseus, is certainly known, about 13 feet 
long, and remarkable for its great variability of colour. It has 
been found, though rarely, in the North Atlantic and Mediterranean. 
A skull from the Cape of Good Hope, which differs slightly from 
that of the above, has been described under the name of G. richard- 
soni. 

Feresia.2—This genus, known at present only by two skulls, 
may be provisionally placed here. These appear to indicate a form 
connecting Globicephalus, Grampus, and Lagenorhynchus. From the 
latter they differ chiefly in the smaller number (about 12) and much 
larger size (6-7 mm. in diameter at base of crown) of the teeth. 

Lagenorhynchus.2—Rostrum scarcely exceeding the length of the 
cranium, broad at the base and gradually tapering towards the 
apex, depressed. Pterygoids normal, meeting in the middle line. 
Teeth small (not exceeding 4 mm. in diameter), 23 to 33. Vertebre 
very numerous, 80 to 90. Spines and transverse processes of the 
lumbar vertebrae very long and slender; centra short. Externally, 
head with a short but not very distinct beak. Two species, 
’ L. albirostris and L. acutus, are occasionally captured on the British 
coasts. Other species occur elsewhere. 

Group B.—Head with distinctly elongated rostrum, or beak, 
generally marked off from the prenarial adipose elevation by a V- 
shaped groove. Rostrum of skull considerably longer than the 
cranial portion. Atlas and axis firmly united ; all the other cervical 
vertebree free. 

Tf we add to it the above-mentioned genus, Lagenorhynchus, this 
group will include all the true Dolphins, Bottle-noses, or, as they 
are more commonly called by seafaring people, “Porpoises,” which 
are found in considerable abundance in all seas, some species being 
habitually inhabitants of large rivers, as the Amazon. They are all 
among the smaller members of the order, none exceeding 10 feet in 
length. Their food is chiefly fish, for the capture of which their 
long narrow beaks, armed with numerous sharp-pointed teeth, are 
well adapted, but some appear also to devour crustaceans and 
molluscs. They are mostly gregarious, and the agility and grace 
of their movements in the water are constant themes of admiration 
to the spectators of the scene when a “school of Porpoises” is 
observed playing round the bows of a vessel at sea. 

1 Gray, Zoology of Erebus and Terror, p. 30 (1846), 
* Gray, Proc. Zool. Soc. 1870, Pp. 7% 
® Gray, Zoology of Erebus and Terror, p. 35 (1846), 


DELPHINIDAE oo 


Delphinus Tooth very numerous in both jaws, {8 to 89, 
occupying nearly the whole length of the rostrum, small, close-set, 
conical, pointed, slightly curved. Rostrum elongated, usually about 
double the length of the cranial portion of the skull. Pterygoids of 
normal form, meeting in the middlo line throughout their length, 
Palate with deep lateral grooves. Vertebras 73 to 75. Pectoral fin 
of moderate size, narrow, pointed, somewhat faleate. Second and 
third digits well developed ; the rest rudimental. 

The type of the genus is the Common Dolphin of the Mediter- 
ranean (2). delphis, Fig. 97), also found in’ the Atlantic, and of 


Fra, 87. The Common Dolphin (Delphinus delphis) From Reinhardt. 


which a closely allied if not identical form is met with in the 
Australian seas (2), forsteri) and in the North Pacitie (D. bairdi). 
Other species ave 1), janira, D. major, ete. 

Tursiops’°—Rostrum tapering moderately from base to apex : 
palate not grooved ; symphysis of mandible short; other cranial 
characters as in Delphinus. Teeth 31 to 33, stout (6 to 7 mm. in 
antero-posterior diameter). Vertebras: C7, D138, L17, C27; total 
64, Limbs as in Delphinus. Represented by the widely distributed 
T. tursio s Ty catalania Weing a second form. Fossil remains of this 
genus from the Italian PHocene have been recently deseribed. 

Prodelphinus%—Rostrm somewhat variable; mandibular sym- 
physis short (less than one-fifth the length of the ramus): other 
eramal characters as ine the preceding genus. Teeth §§ to §9, 
small, not exceeding 3 mm. in diameter, Vertebre 73 to 78. 
Limbs as in Delphinus. Four leading types of this genus are 
recognised (all of which have numerous synonyms) viz. 2. cbscurie, 
Po cuphrosne, Py davis, and 2. longirestris. 

Peron’s Dolphin (Delphinus lateorhamphus, Peron, or Leuco- 
rhamphus peroni, Lilljeborg) resembles some forms of Prodelphinars in 
its cranial characters; but having no dorsal fin, it has heen separated 
generically by some writers. It is not improbable that J/phinus 
borealis, Peale, from the North Pacitic. in which there is likewise no 
dorsal tin, may he an allied form. 

Sfevo.4—Rostrum long, narrow, and compressed, very distinet 
from the cranium; mandibular symphysis as long as, or longer than 
L Linn. Sve’. Vat. 12th ed. vol. ip. 10s (1700), 

2 Gervais, Hist, Vat. des Manmireres, vol. ii, p. 828 (1850). 
¥ Gervais, Osfcographic des Cetaces. po GOA (1S80%, 
+ Gray. Zoology of Hrebus and Perror, po 43 S46, 


273 CETACEA 


one-fourth the length of the ramus; other cranial characters as in 
the preceding genus. Teeth 31 to 34, of comparatively large size 
(5-6 mm. in diameter); surface of their crowns finely grooved. 
Vertebre: C7, D12, L15, C32; total 66. Represented by 
S. rostratus, from which the forms which have received other names 
are probably not specifically separable. 

Sotalia.—Pterygoids narrow, not meeting in the middle line, 
and in their inner borders diverging posteriorly, instead of being 
parallel as in the preceding genera; other cranial characters much 
as in Steno. Teeth tolerably large (4-5 mm. in diameter), 3% to 35, 
with smooth enamelled surface. Vertebre: C7, D12, L 10-14, 

22; total 51-55. Pectoral fin broad at base, the breadth being 
caused by the considerable development and position of the two 
outer digits. Six species are provisionally recognised as distinct, 
including the Chinese White Dolphin (8. sinensis) and 8. pallidus 
from the river Amazon. 


Bibliography of Cetacea—D. ¥. Eschricht, Untersuchungen iiber die Nordischen 
Wallthierc, 1849, contains a copious bibliography of the group up to the date of 
publication. Since that time numerous monographs on special families and 
genera have been published, and a large illustrated general work, Ostéographie des 
Cétacés, by P. J. Van Beneden and P. Gervais, 1869-80. Besides those already 
referred to in the footnotes, the following may be mentioned ; viz. J. F. Brandt, 
“Untersuchungen itber die Fossilen und Subfossilen Cetaceen Europa’s,” in 
Mém. de UV Acad. Imp, de St. Pétersbourg, 7®me sér. vol. xx. 1873 ; C. M. Scammon, 
Marine Maminals of the N. W. Coast of North America, 1874; W. H. Flower, 
“On the characters and Divisions of the Families of the Delphinidw,” Proc. Zool. 
Soc. 1888, p. 466, and List of the Specimens of Cetacea in the British Museum, 
1885 ; F. W. True, ‘ Review of the Family Delphinide,” Bu//. U.S. Nat. Museum, 
No. 36, 1889; P. J. Van Beneden, Histoire Naturelle des Ceétacés des Mers 
@ Europe, 1889. 

For fossil forms, in addition to the works of Van Beneden, Gervais, and Brandt, 
already cited, the reader may refer to various memoirs published by the former 
writer in the Bull. Ac. R. Belgique and Ann. Mus. R. Hist. Nat. Belg. 
See also R. Lydekker, ‘‘ The Cetacea of the Suffolk Crag,” Quart. Journ. Geol. 
Soc. vol. xlii. p. 7 (1887), and Catalogue of the Fossil Mammalia in the British 
Museum, pt. v. (1887). 


' Gray, Cat. Seals and Whales Brit. Mus. 2d ed. p. 393 (1866). 


CHAPTER IX 
THE ORDER UNGULATA 


UnpeEr this term may be included provisionally a large and rather 
heterogeneous group of mammals, the existing members of which 
form the Pecora and Bellue of Linnzus, the Ruminantia and 
Pachydermata of Cuvier. A few years ago it was found convenient 
to restrict the order to a well-marked and distinctly circumscribed 
group, comprising the two sections known as Perissodactyla and 
Artiodactyla, and to leave out such isolated forms as the Elephant 
and Hyrax ; but the discovery of a vast number of extinct species, 
which could not be brought under the definition of either perisso- 
dactyle or artiodactyle Ungulates, and yet are evidently allied to 
both, and to a certain extent bridge over the interval between 
these and the isolated groups just mentioned, makes it necessary 
either to introduce a number of new and ill-defined ordinal 
divisions, or to widen the scope of the original order so as to 
embrace them all. 

The existing forms are all animals eminently adapted for a 
terrestrial life, and in the main for a vegetable diet. Though a 
few are more or less omnivorous, and may under some circumstances 
kill living creatures smaller and weaker than themselves for food, 
none are distinctly and habitually predaceous. Their teeth are 
markedly heterodont and diphyodont,—the milk set being well 
developed and not completely changed until the animal attains its 
full stature. The molars have broad crowns with tuberculated or 
ridged surfaces. There are no clavicles.t Their toes are provided 
with blunt, broad nails, or in the majority of cases with hoofs, more 
or less enclosing the ungual phalanges. The scaphoid and lunar 
bones of the carpus are always distinct. The humerus has no 
entepicondylar foramen. The number of digits varies from five to 
one; and the radius and ulna may be united together. 

1 Since this was in type the discovery of transient rudimentary clavicles in 
the embryo of the Sheep has been announced by Wineza (Morpholog. Jahrh. xvi. 
p. 647). 

1 


[os 


274 ENGOL ATs 


The more generalised of the fossil forms do not conform in all 
respects to the above-mentioned characters ; clavicles being present 
in Typotherium, and perhaps in some of the Condylarthra, while in 
the latter group the humerus may have an enutepicondylar foramen, 
and thus approximate to the corresponding bone of the Carnivora. 
Wide as is the gap between existing Carnivores and Ungulates, there 
are indeed more or less strongly marked evidences of affinity 
between the earlier members of the two orders, as will be again 
noticed under the head of the suborder Condylarthra. A departure 
from the normal type of foot-struecture is exhibited by the extinct 
AMuacrotheriun, provisionally included in the Perissodactyla, where 
the digits terminated in long and curved clus. 

As a general rule, the cheek-teeth have distinct roots, and in 
those of the existing suborders a gradual increase in the height of 
the crowns of these teeth may be noticed in passing from the more 
generalised to the more specialised types. Those teeth in which 
the crowns are low, and their whole structure visible from the 

- evinding surface, ave termed brachydont (Fig. 122); while those with 
higher crowns, in which the bases of the infoldings of enamel are 
invisible from the grinding surface, ave known as hypsodont (Miz. 123). 
Again, when the tubercles on the crowns of the molars are more or 

less cone-like in form the tooth 
is said to be dunodont : but when 
they ave oxpanded in an antero- 
posterior direetion and curved into 

a crescent shape the tooth is 

deseribed as sclenodont. 

The whole order) may he 
divided into the Ungulata Vera, 
containing the suborders Perisso- 
dactyla and Artiodactyla, and a 
somowhat heterogeneous assem- 
Dlage of animals which may he 
ealled Subuneulata or Ungulata 
Polydactyla. Cope has pointed 
out a character in the structure 
of the carpus by which the latter 
are dillerentiated from the former. 
Thus in all the Subungulata the 
bones of the proximal and distal 

Fic, 98.—Right fore foot of Indian Kle- row aD the primitive OP TONG 
phant. x}. U, ulna; R, radius; ¢, cunci- typical relation to cach other (see 
form; 7, lunar; se, aeaphoid ; uw, unciferm ; Vig. 98) : the os mae of the 
bec cara es erred ines ERA second vow articulating mainly 

with the Iunar of the first, or 
with the cuneiform, but not with the seaphoid. But in the group to 


UNGULATA VERA 275 


which the vast majority of modern Ungulates belong the second or 
distal row has been shifted altogether towards the inner side of the 
limb (see Fig. 99), so that the magnum is brought considerably 
into relation with the scaphoid, and is entirely removed from the 
cuneiform, as in the great majority of existing mammals. 

It will be on the whole more convenient to commence our 
survey of the members of this suborder with the more specialised 
group of the Ungulata Vera, in which the Artiodactyla will be 
taken first. 


UNGULATA VERA. 


In the typical Ungulata the feet are never plantigrade, and the 
functional toes do not exceed four—the inner digit being suppressed, 
at all events in all forms which have existed since the Upper 
Eocene period.2. The os magnum of the carpus articulates freely 
with the scaphoid. The allantois is largely developed, and the 
placenta, so far as is known, is non-deciduate ; the chorionic villi 
being either evenly diffused or collected in groups or cotyledons (in 
Pecora). The testes descend into a scrotum. There is never an os 
penis. The uterus is bicornuate. The mamme are usually few 
and inguinal, or may be numerous and abdominal (as in Suina), but 
are never solely pectoral. The cerebral hemispheres in existing 
Ungulates are well convoluted. 

The group is now, and has been throughout almost the whole 
of the Tertiary period, composed of two perfectly distinct sections, 
differing from each other, not only in the obvious characters of the 
structure of the limbs, but in so many other parts of their organisa- 
tion that they must be considered as of the rank at least of 
suborders. The characters of these divisions, first indicated by 
Cuvier, were thoroughly established by Owen, by whom the names 
whereby they are now generally known were proposed. 


Suborder ARTIODACTYLA. 


This is a well-defined group, traceable from the Eocene period, 
though then apparently by no means so numerous as the Perisso- 
dactyles. Some of its types, as that represented in the existing 
Swine, have retained to the present time much of the primitive 
character of the group; but others have been gradually becoming 
more specialised and perfected in structure, and its latest modifica- 
tion, the Cavicorn Ruminants or Luride (Antelopes, Sheep, and 
Oxen), are now the dominating members of the great Ungulate 
order, widespread in geographical range, rich in generic and specific 
variation, and numerous in individuals—forming in all these 

1 Also known as Diplarthra. 
? The pollex is present in the manus of the extinct Cotylops. 


276 UNGULATA 


respects a great contrast to such decadent types as those represented 
by the Tapirs and Rhinoceroses. 

The principal anatomical characters by which the Artiodactyles 
are distinguished from the Perissodactyles are as follows. The 
premolar and molar teeth usually not alike, the former being 
single and the latter two-lobed. The last lower molar of both first 
and second dentition almost invariably three-lobed; and the first 
tooth of the upper cheek series always without a milk-predecessor. 
Nasal bones not expanded posteriorly. No alisphenoid canal. 


Fic. 99.—Bones of right fore foot of existing Artiodactyles. A, Pig (Sus scrofa), x4; B, 
Red Deer (Cervus elaphus), x}; C, Camel (Camelus bactrianus), x}. C, Ulna; R, radius; + 
cuneiform; /, lunar; s, scaphoid; u, unciform ; m, magnum; td, trapezoid; tin, trapezium, 
From Flower’s Osteology of Mammalia. 


Dorsal and lumbar vertebre together always nineteen, though the 
former may vary from twelve to fifteen. Femur without. third 
trochanter. Third and fourth digits of both feet almost equally 
developed, and their ungual phalanges flattened on their inner or 
contiguous surfaces, so that each is not symmetrical in itself, but 
when the two are placed together they form a figure symmetrically 
disposed to a line drawn between them. Or, in other words the 
axis or median line of the whole foot is a line drawn between the 
third and fourth digits, while in the Perissodactyles it is a line 
drawn down the centre of the third digit. Distal articular surface 


ARTIODACTYLA a7 


<3 


of the astragalus divided into two nearly equal facets, one for the 
navicular and the other for the cuboid bone. The caleaneum with 
an articular facet for the lower end of the fibula. Stomach almost 
always more or less complex. Colon convoluted. Ccum small. 
Placenta diffused or cotyledonary. Mamme few and inguinal, or 
numerous and abdominal. 

In treating of many sections of mammals, it is only from the 
existing species that our characters and classification can be derived, 
and to these chiefly our observations upon the group must be 
directed, many of the extinct forms being so little known that they 
can only be referred to incidentally. With the Ungulata, however, 
it is quite otherwise. The history of the Artiodactyla throughout 
the Tertiary period is now well known, and throws great light upon 
the position and relations of the existing groups. 

The principal modifications which have taken place in the type 
from its earliest known and most generalised manifestation have 
been the following :— 

1. As regards the teeth. Assumption by the grinding surfaces 
of the molar teeth either of a bunodont or of a selenodont form. 
Moditieation of the latter from a brachydont to a hypsodont type. 
Loss of upper incisors. Development of canines into projecting 
tusks. Loss of anterior premolars. 

2. As regards the limbs. Reduction of the ulna from a complete 
and distinet bone to a comparatively rudimentary state, in which it 
coalesces more or less firmly with the radius. Reduction of the 
fibula till nothing but its lower extremity remains. Reduction 
and final loss of external pair of digits (second and fifth), with coal- 
escence of the metapodial bones of the two middle digits. Union 
of the naviewlar and cuboid, and sometimes the ectocuneiform, 
bones of the tarsus. 

3. Change of form of the odontoid process of the axis vertebra 
from a cone to a hollow half-eylinder. 

4. Development of horns or antlers on the frontal bones, and 
gradual complication of form of antlers. 

3. Ry inference only, increasing complication of stomach with 
ruminating funetion superadded. Modification of placenta from 
simple diffused to cotyledonary form. 

The primitive Artiodactyles, with the typical number (44) of 
incisor, canine, and molar teeth, brachydont molars, conical odon- 
toid process, four distinct toes on each foot, with metapodium and 
all carpal bones distinct, no frontal appendages, and (in all proha- 
bility) simple stomach and diffused placenta, were separated at a 
very early period into Bunodonts and Selenodonts, although there 
is evidence of intermediate forms showing a complete transition 
from the one modification to the other. These and other fossil 
forms so completely connect the four groups—Suina, Tylopoda, 


278 UNGULATA 


Tragulina, and Pecora—into which the existing members of the 
suborder have become divided, that in a general classification 
embracing both living and extinct forms these divisions cannot be 
maintained. In the present work, however, it will be convenient 
to retain them, mention being made of some of the chief annectant 
forms in separate sections. 


SUINA. 


The existing members of this group are characterised by their 
bunodont molars, and the absence of a complete fusion of the third 
and fourth metapodials to form a “cannon-bone.” The full 
Eutherian dentition is very frequently present. 

Remains of very generalised swine-like animals have been 
abundantly found in Tertiary formations both in America and 
Europe. In the former continent they never (so far as present 
evidence indicates) underwent any great diversity of modification, 
but gradually dwindled away and almost died out, being only re- 
presented in the actual fauna by the two closely allied species of 
Peccary, among the smallest and most insignificant members of the 
group, which have existed almost unchanged since the Miocene age 
at least, if the evidence of teeth alone can be trusted. In the Old 
World, on the other hand, the swine have played a more important 
part in recent times, having become widely distributed, and throwing 
off some curiously specialised forms. At the present time, though 
not very numerous in species, they range through the greater part 
of the Old World, except within or near the Ar otic Circle, although, 
in common with all the other members of the great Ungulate order, 
they were completely absent from the whole of the Australian region, 
until introduced by man in very recent times. 

The existing swine-like animals may be divided naturally into 
three families:—I. Hippopotamide ; IL. Suide, or true Pigs; III. 
Dicotylide, or Peccaries.1 


Family HippopoTaMIp&. 


Muzzle very broad and rounded. Feet short and broad, havi ing 
four subequal toes, with short rounded hoofs, all reaching 
the ground in walking. Incisors not rooted, but continuously 
growing ; those of the upper jaw curved and directed downwards ; 
those of ‘the lower straight and procumbent. Canines very lar ge, 
curved, continuously growing ; those of the upper jaw directed 
downwards. Stomach complex. No cxeum. 

Hippopotamus.2—This genus may be taken to include all the 
known members of the family; it appears to have been always 

1 Tn the table on p. 89 the Peccaries are included in the Suide. 
? Linn. Syst. Net. 12th ed. vol. i. p. 101 (1766). 


AIPPCPOTAMID£E a9 


contined to the Old World. The dentition may be expressed by the 


== 


1 : 
formula i = ais) P5.m 


3 


wy w 


. The crowns of the molars (Fig. 100) 


when worn present rretuilchaiped surfaces of dentine: and those of 
tae premolars are sharp. The 
facial portion of the skull is much 
elongated, the orbits are tubular 
and very prominent, and che man- 
dible has a large rounded descend- 
ing dance at ics angle. The ears 
are small, the tail is short. and the 
legs are likewise so short that che 
belly i is raised a a little distance 
abore the ground. The brain is 
not richly eony calanod, and divers 
aS consider oly from thar of 
the Pics. approximating in some 
respects to that of the Camel and 
Laos bar on aes pha 


an axial length of 11 feet, and measurin 
the zreater curvature. Irs axcs is the pricras being 
cuated almost in the pelvis. and i dointo three distinet 
compartments, of which the hind is eviindrieal The liver of che 
adult is of extremely Simple form, elonzated sransverse.y. and narrow 
from above downwards. With the excepvion of a few vats of 


upwards of 15 feet along 


280 CNGULATA 


hair on the lips, on the sides of the head and neck, and at the 
extremity of the short compressed tail, the skin of the hippopotamus, 
some portions of which are two inches in thickness, is entirely desti- 
tute of covering. 

The common Hippopotamus (H. amphilius), widely distributed 
in the rivers and lakes of the African continent, is a huge bulky 
animal, characterised by having only two incisors on either side 
of each jaw; the central lower pair being very much larger than the 
outer ones. A male from the Upper Nile which lived for nearly 
thirty years in the gardens of the Zoological Society of London 
measured 12 feet along the back from the nose to the root of the tail. 

The Hippopotamus lives in herds of from twenty to forty 
individuals on the banks and in the beds of rivers, in the neighbour- 
hood of which it finds its food. This consists chiefly of grass and 
aquatic plants, of which it consumes enormous quantities, the 
stomach being capable of containing from 5 to 6 bushels. These 
animals feed principally by night, remaining in the water during the 
day, although in districts where they are undisturbed by man they 
are less exclusively aquatic. In such regions they put their heads 
boldly out of the water to blow, but when rendered suspicious by 
persecution, they become exceedingly cautious, only exposing their 
eyes and nostrils above the water, and even this they prefer 
doing amid the shelter of water plants. In spite of their enormous 
size and uncouth form, they are expert swimmers and divers, and 
can remain under the water from five to eight minutes. They 
are said to walk with considerable rapidity on the bottoms of 
rivers, beneath at least a foot of water. At nightfall they come 
on land to feed; and when, as often happens on the banks of 
the Nile, they reach cultivated ground, they do immense damage 
to growing crops, destroying by their ponderous tread even more 
than they devour. 

A much smaller species, known as the Pigmy Hippopotamus 
(Z. libertensis), inhabits some of the rivers of Western Africa, and 
is characterised by having only a single pair of lower incisors. 
Mainly on this account, it has been proposed to regard this species 
as representing a distinct genus, under the name of Cheropsis ; but 
since it agrees so essentially in other characters with the common 
form, and sometimes has two incisors on one side of the lower 
jaw, it appears preferable to include it in the type genus. The 
greater relative size of the brain-cavity as compared with the facial 
portion of the skull renders, indeed, the contour of the skull 
decidedly different from that of H. amphibius - but this is a feature 
generally found in young individuals of larger species, and also in 
the adults of allied smaller forms. 

Both the existing species are now exclusively confined to Africa, 
but in the Pleistocene and Pliocene periods the genus was widely 


SUIDE 281 


spread over the Old World. Thus in the Upper Pliocene of the 
Continent and the Pleistocene of England we meet with remains of 
a very large fossil Hippopotamus which cannot be specifically 
distinguished from H. amphibius. In the Pleistocene and Pliocene of 
India there are two species having three pairs of incisors in both 
jaws. Of these A. palwindicus has the second pair in the lower jaw 
very minute, and evidently just about to disappear ; from which we 
learn that it is this pair which is missing in H. amphibius. In the 
still more generalised H. siralensis the three incisors in’ the 
lower jaw are of equal size. Hexaprotodont species also occur 
in the Upper Tertiaries of Burma and Algeria. Small tetra- 
protodont species (A. pentlandi and H. minutus) have left their 
remains in enormous quantities in the caves and fissures of Sicily 
and Malta, 


Family Svip®. 


An elongated mohile snout, with an expanded, truncated, nearly 
naked, flat, oval terminal surface in which the nostrils are placed. 
Feet. narrow ; four completely developed toes on each. Hoofs of 
the two middle toes with their contiguous surfaces fattened. The 
outer (second and fifth) digits of existing forms not reaching to 
the ground in the ordinary walking position. Teeth variable in 
number, owing to the suppression in some forms of an upper incisor 
and one or more premolars. Incisors rooted. Upper canines 
curving more or less outwards or upwards. Stomach simple, _ 
except for a more or less developed pouch near the cardiae orifice. 
Acreum. Colon spirally coiled. Confined to the Old World. 

The mandible has no descending flange at the angle. The 
crowns of the molars do not wear into such distinct trefoils as in 
the Hippopotamus, and are oblong 
in shape. The last molar of both 
the upper and lower jaw (Fig. 102) 
has an additional hinder lobe or 
talon, varying in size in the different 
species. The upper premolars are 
simpler than the true molars. Fie. 102,.—Grinding surface of a worn 

Sus.t—Dentition: Ee c 1, p 4, m third right lower molar of the Wild Boar 
8; total 44. Upper incisors dimin- 6°72") Aferowen. 
ishing rapidly in size from the first to the third. Lower incisors 
long, narrow, closely approximated, and almost horizontal in position, 
their apices inclining towards the middle line ; the second shghtly 
larger than the first, the third much smaller. Canines strongly 
developed and with persistent roots and partial enamel-covering. 
those of the upper jaw not having the usual downward direction, 


1 Linn. Srsf, Not, 12th ed. vol. ip. 162 (1760). 


282 UNGULATA 


but curving strongly outwards, upwards, and finally inwards, while 
those of the lower jaw are directed upwards and outwards with 
a gentle backward curve, their hinder edges working and wearing 
against the front edges of the upper canines! They appear 
externally to the mouth as tusks, the form of the upper lip being 
modified to allow of their protrusion, but are much less developed 
in the females than in the males. The teeth of the molar series 
gradually increase in size and complexity from first to last, and 
are arranged in contiguous series, except that the first lower 
premolar is separated by an interval from the second. First and 
second upper premolars with compressed crowns and two roots. 
The third and fourth have an inner lobe developed on the crown, 
and an additional pair of roots. The first and second true molars 
have quadrate crowns, with four principal obtuse conical cusps, 
around which numerous accessory cusps are clustered. The length 
of the third molar is nearly equal (antero-posteriorly) to that of 
the first and second together, its crown. having, in addition to the 
four principal cusps, a large posterior talon or heel, composed of 
numerous clustered conical cusps, and supported by several additional 
roots. The lower molar teeth resemble generally those of the upper 
jaw, but are narrower. Milk dentition: 73,¢4,m3; total 28,— 
the first permanent premolar having no predecessor in this series. 
The third incisor, in both upper and lower jaws, is large, developed 
before the others, and has much the size, form, and direction of 
the canine. Vertebre: C7,D13-14, L 6,8 4, C 20-24. The hairy 
covering of the body varies much under different conditions of 
climate, but when best developed, as in the European Wild Boar, 
consists of long stiff bristles, mostly abundant on the back and 
sides, and of a close softer curling under-coat. 

The skull of the Pigs (Figs. 103-105) has the axis of the face 
bent down upon the basicranial axis, as is also the case with the 
Sheep. Its most striking feature is the elevation and backward 
slope of the occipital crest formed by the union of the supraoccipital 
and parietals. The broad and flat frontals have small postorbital 
processes, which do not join the zygomata, so that the orbits are 
open behind. The nasals are very long and narrow; and the pre- 
maxille send up long nasal processes, stopping short of the frontals. 
A peculiar prenasal bone is developed at the anterior extremity of 
the mesethmoid, which serves to strengthen the cartilaginous snout. 
The palate is long and narrow, and extends behind the last molar 


1 If from any accidental circumstances these teeth are not constantly worn 
down by friction, they grow into a complete circle, the point penetrating the 
bone of the jaw close to the root of the tooth. The natives of the Fiji Islands 
avail themselves of this circumstance to produce one of their most valued orna- 
ments—a circular boar’s tusk: the upper canines being extracted, the lower ones 
are allowed to grow to the desired form. 


SUIDE 283 


tooth. In most species the occipital crest is more nearly vertical 
than in the skull represented in Fig. 104. 

This genus occurs at present under three principal modifications 
or subgenera. 

A.—Sus proper comprises a number of animals 
found in a wild state throughout the greater part 
of Europe (except where extermin- 
ated by human agency), the north 
of Africa, southern continental 
Asia, and the 
great islands of 
the Malayan 
archipelago, 
Formosa, and 4 
Japan. The fol- 
lowing among 
others have 
been admitted 
by many zo- 
ologists as dis- 
tinct species : 


PN 


Y, @ ” 
—Sus ser of a, Fic. 103.—Left lateral view of the dentition of the Boar (Sus scrofa), 
the Wild Boar the roots of the teeth being exposed by removing the external lamina 


of Europe, Asia ie 
Minor, and North Africa, once common throughout the British 
Isles ; S. sennaurensis, North-East Africa; S. cristatus, India; 8. 


Fic. 104.—Left lateral view of the skull of Sus longirostris. } natural size. (From Nehring.) 


vittatus, Java, Borneo, Amboyna, Batchian ; %. pupuensis, New 
Guinea; S. timorensis, Timor and Rotti; S. andamanensis, Anda- 


284 “UNGULATA 


man Islands: S. feéranus, Formosa; 8. leucomystaz, Japan; 8. 
verrucosus, Java, Borneo, Ceram; 8. barbatus, Borneo; S. celebensis, 
Celebes, Philippines, and Moluccas; 8. longirostris, Borneo and 
Java. The last four species form an allied group in which the 
facial portion of the skull may be greatly 
elongated ; 8. barbatus and SS. celebensis 
being characterised by the small size and 
simple structure of the talon of the third 
molars. The skull of S&. longirostris is 
shown in Figs. 104 and 105. The small 
S. andamanensis also has very simple third 
molars. 8S. vittatus, S. leucomystaa, S. cris- 
tatus, S. tuéranus, and S. papuensis form 
another group, in which the third molar 
is generally of very complex structure, 
more or less closely allied to the Wild 
Boar; and Dr. Nehring is inclined to 
think that the whole five might be in- 
cluded under a single specific name. This 
list will give some idea of the geographical 
distribution of wild Pigs, but it must be 
borne in mind that through the whole of 
this region, and in fact now throughout 
the greater part of the habitable world, 
Pigs are kept by man in a domesticated 
state, and it is still an open question 
whether some of the wild Pigs of the 
islands named above may not be local 
races derived originally from, or crossed 
with, imported domestic specimens. In 
New Zealand a wild or rather “feral” 
race is already established, the origin of 
which is of course quite recent, since it is 
Fic. 105.—Frontal aspect of well ascertained that no animal of the 
the cranium of Sus longirostris. kind ever lived “pon th is] a ] 
} natural size. (From Nehring.) P € isian unt 
after its settlement by Europeans. 
Whether the various breeds of domestic Pigs have been derived 
from one or several sources is still unknown. As in so many 
similar cases, there is no historie evidence upon the subject, 
and the researches of naturalists, as Nathusius, Riitimerer, 
Rolleston, Nehring, and others, who have endeavoured to settle 
the question on anatomical evidence, have not led to any satis. 
factory conclusions. It is, however, tolerably certain that all 
the species or forms of wild Pigs enumerated above and all the 
domestic races are closely allied, and it is probable (though of 
this there has been no opportunity of proof) will breed freely 


SUIDE 285 


together. It is a curious circumstance that the young of all 
the wild kinds of Pigs (so far as yet is known) present a 
uniform coloration, being dark brown with longitudinal stripes of 
a paler colour, a character which completely disappears after the 
first few months. On the other hand, this peculiar marking is 
rarely seen in domestic Pigs in any part of the world, although it 
has been occasionally observed. It is stated by Darwin that the 
Pigs which have run wild in Jamaica and the semiferal Pigs of New 
Granada have resumed this aboriginal character, and produce longi- 
tudinally striped young; these must of course be the descendants 
of domestic animals introduced from Europe since the Spanish 


Fic. 106.—Wild Boar and Young. 


conquest, as before that time there were no true Pigs in the New 
World. Another character by which the European domestic Pig 
differs from any of the wild species is the concave outline of the 
frontal region of the skull, a form still retained by the feral Pigs 
in New Zealand. ; 

B.—The diminutive Pig of the Nipal, Terai, and Bhutan, Sus 
salvanius, has been separated from the rest by Hodgson under the 
generic name of Porcula, but all the alleged distinctive characters 
prove on more careful investigation to have little real value. Owing 
to its retired habits and power of concealment under bushes and 
long grass in the depths of the great Sal Forest, which is its 
principal home, very little has been known of this curious little 
animal, scarcely larger than a hare. The acquisition of living 


286 UNGOLATA 


specimens in the London Zoological Gardens has, however, afforded 
opportunities for careful anatomical observation.t 

C.—Two well-marked species of African Swine have been with 
more reason separated under the name of Potamecherus. The denti- 
tion differs from that of the true Svs, Inasmuch as the anterior 
premolars have a tendeney to disappear: sometimes in adult 
specimens the first upper premolar is retained, but it is usually 
absent, as well as the first and often the second lower premolars. 
The molar teeth are also less complex; the last especially having a 


Fi. 107.—The Red River-Heg (Sus poreus). From Selater, Guide to Animals 
in Zoological Society's Gardens, 1833, p. 183. 


much less developed talon. There are likewise characteristic cranial 
differences. The two species are very distinct in outward appearance 
and coloration. One is 8. africanus, the South African River-Hoe, 
or Bosch-Vark, of a gray colour, and the other S. percus, the West 
African Red River-Hog (Fig. 107), remarkable for its vivid colouring 
and long pencilled ears. It should be noted that the young of both 
these species, as well as of the pigmy S. sa/ranius, present the striped 
character of the true Sus, a strong indication of close attinities. 
whereas in all the following forms this is absent. 

The genus Svs, in the above exténded sense, is well represented 
in the Tertiaries of the Old World from the period of the Lower 
Pliocene upwards. In the Pliocene and Pleistocene of India 


1 See Garson, Proce. Zool. Sov. Lon’?, 1S82, p. £13. 


SIDE ost 


Fadromeni and Ss becwallansis are erceearauae : > 
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. —<Agrocharine 
(dorivchari.s, 

Orbits closed, a lachrymal fossa, no diastema, the last upper 
premolar with one outer column: outer wall of upper 
molars flattened.—Cotylopine Cotulops, Eporcedon, Me ve 
cocherus, Cyclopidius, eter 

II. Upper molars with five columns.—Protercentine Protoresdsn’. 


Anoploticriida. 
—This family in- 


BH. 
fe 3 some. eludes several 
gel rH Upper Eocene 
ee i . European genera. 
e =) a Vi with = selenodont 
ae BS \ upper molars, 
as ‘Y earrvinge five 
wat vine 
ASS columns arranged 
S : = 
aS Alu ays 
Fic. 113.—Restoration of Aner? therigns ooumnne ats ay Ale ens 
(Upper Eocene). Crvier. therium. One of 


1 This name (Leidy, 1551) is preoeeupied by Groves Agassiz. 1838 


294 UNGULATA 


the earliest known, .fuoplotherium, was fully described by Cuvier 
from remains found in the Paris gypsum-beds (Upper Eocene). 
Its forty-four teeth formed a series unbroken by « gap or diastema, 
and were of uniform height (as in Man alone of existing mam- 
mals). Its tail was long, with large chevron bones underneath, 
not usually found in Ungulates, and there were either three or 
two toes on each foot. It was in many respects a much- 
specialised form, apparently not on the line of descent of any of 
the existing groups. 

Dacrytherium is an allied genus whose dentition leads on to that 
of the smaller Viphodon. The latter genus is characterised by the 
compressed and elongated form of the crowns of the first three 
premolars, which thus approximate to those of the Chevrotains. 
There were only two functional digits to the feet. The so-called 
HHyopotamus pieteti, of the Swiss Eocene, is a species of Dacrytherium. 

Cenothertide.—The typical representatives of this family are 
small animals not larger than the Chevrotains, with the full comple- 
ment of teeth, generally no marked gap in the series, and the 
crowns of the upper molars carrying two columns on the anterior 
and three on the posterior half of the crown—precisely the reverse 
of the arrangement obtaining in the //nthracotheriidw. The known 
forms are from the Upper Eocene and Lower Miocene of Europe. 
In Cenotherium the molars are selenodont, while they are bunodont 
in Dichobunus. LHomacodun, of the Bridger Eocene of the United 
States, is closely allied to the the latter. The first lower premolar 
of Dichobunus assumes the form and function of a canine. Spunio- 
thertum (Metriotherium) is a much larger form, in which the molars 
are not unlike those of /uthrucotherium, if the arrangement of the 
cusps were reversed; it occurs in the Eocene Phosphorites of 
France. It is suggested that the Z'ylopulu may have originated 
from this group. 

Tapirulus is a small Eocene Artiodactyle with the columns of 
the upper molars, which are somewhat like those of Myopotanus, 
tending to form transverse ridges ; its family position is uncertain. 

Dichodontide,—The European genera included in this family all 
have quadritubercular selenodont molars, and show signs of approxi- 
mating more or less closely to existing types.  Michulon, from the 
Upper Eocene and Lower Miocene, has the full complement of teeth, 
which show no diastema, and have low crowns. The fourth upper 
premolar has four columns, like the true molars, and the corre- 
sponding lower tooth three complete lobes; these features being 
unknown in any other Sclenodonts. In Lophiomerys, of the same 
heds, the somewhat higher crowns of the molars approximate to 
those of the Cervidir, but the hinder lobes of the upper ones are 
imperfectly developed ; the genus may he allied to the Traqulide, 
In the small Gelocus, of the Lower Miocene, the molars are not 


CAMELID A 295 
unlike those of Dichodon bub the navicular and cubotl bones of 
the tarsus wero fused together, and the metatarsals had united to 
form oa “eannon-boue,” although the metacarpals still remained 
distinet, It is not improbable that upper incisors were wanting ; 
and it has boen suggested that we have in this genus the ancestral 
type of the Cragulide and Cereide, 


TYLOPODA, 
Manily CAMELUDY. 


This group is represented at the present day by the two species 
of Camels of the Old World and the Llamas of South America, 
collectively constituting the family Chanelide, Phe special characters 
which tho Llimas and Camels have in common, and the combina- 
tion of which distinguishes them from the vest of the Artiodaety les, 
areas follows. The promaxillas have the fall aumber of incisor 
tooth in the young state, and the outermost is persistent through 
life as an isolated Taniariform tooth, The canines are present in 
both jaws, and those of the mandible are differentiated from the 
long, procumbent, and spatulate incisors, being subereet and pointed. 
The crowns of the true molars belong to the crescentic or selen- 
odont type, and are very hypsodont; but one or more of the 
anterior prentohus is usually detached from the series, and is 
of simple pointed form. ‘The auditory bulke is filled with cancellous 
tissno, Tho hinder part of the body is much contracted, and the 
fom long and verticnlly plueed, se that the knee-joint is lower 
in position, and the thigh altogether more detached from. the 
abdomen than in most quadrapedal mammals. ‘The limbs are 
Tong, bat with only the third and fourth digits developed ; ioe 
traces of any of the othors being prosent. Tho trapezoid and mag- 
ume of the carpus, and) the cuboid and navientar of the tarsus ave 
distinet. The two metapodial bones of each limb are confluent: for 
the greater part of thet length, Chongh separnted for a considerable 
distanee at the lower end. ‘Pheir distal artienlar stirfiees, mstead 
of being pulley like, with deep ridges and grooves, as in other recent 
Artiodaetyles, ave simple, rounded, and smooth. ‘The proximal 
phaltnges ave expanded at their distal ends, and the wide, depressed 
middle phalanges ave embedded ina broad cutaneons pad, forming 
the sole of the foot, on whieh the animal rests in walking, iste 
ofon the hoofs. The ungnal phakiges are very small and nodular, 
not flattened: on thete inner or opposed surfaces, and noe completely 
oneased ta hoofs, but bearing mails on their upper sturfiee only, 
Tho cervical region is long and flexnous, and the vertebre of which 
it is composed are vetutrkable for the position of the canal for 


296 UNGULATA 


the transmission of the vertebral artery, which does not perforate 
the transverse process, but passes obliquely through the anterior 
part of the pedicle of the arch (a condition only found in two other 
genera of mammals, J/ucrauchenia and Ayrmecophaya). There are 
no horns or antlers. Though these animals ruminate, the stomach 
differs considerably in the details of its construction from that of 
the Pecora. The interior of the rumen or paunch has no villi on 
its surface, and there is no distinct psalterium or maniplies. Both 
the first and second compartments are remarkable for the presence 
of a number of pouches or cells in their walls, with muscular septa, 
and a sphincter-like arrangement of their orifices, by which they can 
be shut off from the rest of the cavity, and into which the fluid 
portion only of the contents of the stomach is allowed to enter.! 
The placenta is diffuse, as in the Suina and Tragulina, not. coty- 
ledonary, as in the Pecora. Finally, the Camelide differ not only 
from other Ungulates, but from all other mammals, in the fact 
that the red corpuscles of the blood, instead of being circular in 
outline, are oval, as in the inferior vertebrated classes. 
Camelus.2—Dentition of adult : 74, ¢4, p 3, m3; total 34. First 
upper premolar simple, placed immediately behind the premaxille, 
and separated by a long diastema from the penultimate tooth of 
that series. Lower incisors somewhat proclivous, the outermost the 
largest. Skull elongated, with an overhanging occiput, orbits com- 
pletely surrounded by bone, and the premaxille not articulating 
with the arched and somewhat elongated nasals. Vertebre: C 7, 
D 12, L 7, 84, C 13-15. Ears comparatively short and rounded. 
One or two dorsal adipose humps. Feet broad, with the toes very 
imperfectly separated. Tail well developed, tufted at the end. 
Hair nearly straight, and not woolly. Size very large and bulky. 
The genus is now represented by two species, viz. the single- 
humped Arabian Camel (Camelus dromedarius), and the double- 
humped Bactrian Camel (C. bactriunus, Fig. 114).3 The former 


* The stomach of the Camel inhabiting the Arabian desert is commonly 
looked upon as a striking example of specialised structure, adapted or modified 
in direct accordance with a highly specialised mode of life ; it is therefore very 
remarkable to find an organ exactly similar, except in some unessential details, 
in the Llamas of the Peruvian Andes and the Guanacos of the Pampas. No 
hypothesis except that of a common origin will satisfactorily account for this, 
and, granting that this view is correct, it becomes extremely interesting to 
find for how long a time two genera may be isolated and yet retain such close 
similarities in parts which in other groups appear readily subject to adaptive 
modifications. 

> 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. <A callosity on the inner side of the fore limb above 
the carpus. 

Fossil Species. —In the Pleistocene Horses of South America 
described as Hippidiwm, as well as in the closely allied ones from 
North America for which the name Pliohippus has been proposed, 
the upper molars are shorter and more curved than in the existing 
species, while their anterior pillar is not longer antero-posteriorly 
than in Hipparion ; the lateral claw-like hoofs persisting. Some of 
the European Pliocene species (like E. stenonis) agree with these 
species in the form of the grinding surface of the anterior pillar 
of the upper molars. In one of the species from the Lower 
Pliocene of India (£. sivalensis)—which was a contemporary of 
Hipparion—and in all the existing species, the grinding surface of 
the pillar in question is greatly elongated in the antero-posterior 
direction, as in Fig. 157, «. 

Fossil remains of Horses are found abundantly in deposits of 
the most recent geological age in almost every part in America, 
from Eschscholtz Bay in the north to Patagonia in the south. In 
that continent, however, they became quite extinct, and no Horses, 
either wild or domesticated, existed there at the time of the 
Spanish conquest, which is the more remarkable as, when intro- 
duced from Europe, the Horses that ran wild proved by their 
rapid multiplication in the plains of South America and Texas that 
the climate, food, and other circumstances were highly favourable 
for their existence. The former great abundance of Hguide in 
America, their complete extinction, and their perfect acclimatisation 


1 Linn. Syst. Nat. 12th ed. vol. i. p. 100 (1766). 


382 UNGULATA 


when reintroduced by man, form curious but as yet unsolved 
problems in geographical distribution. 

Existing Species—The existing species of the genus are the 
following :— 

The Horse, Equus caballus, is distinguished from the others by 
the long hairs of the tail being more abundant and growing quite 
from the base as well as the end and sides, and also by possessing 
a small bare callosity on the inner side of the hind leg, just below 
the “hock” or heel joint, in addition to the one on the inner side 
of the fore limb above the carpus, common to all the genus. The 
mane is also longer and more flowing, and the ears are shorter, 
the limbs longer, the hoofs broader, and the head smaller. 

Though the existing Horses are not usually marked in any 
definite manner, or only irregularly dappled, or spotted with light 
surrounded by a darker ring, many examples are met with showing 
a dark median dorsal streak like that found in all the other 
members of the genus, and even with dark stripes on the shoulders 
and legs indicating “the probability of the descent of all the 
existing races from a single dun-coloured, more or less striped, 
primitive stock, to which our horses still occasionally revert.” ! 

In Europe wild Horses were extremely abundant in the 
Neolithic or polished-stone period. Judging from the quantity of 
their remains found associated with those of the men of that time, 
the chase of these animals must have been among man’s chief 
occupations, and they must have furnished him with one of his 
most important food supplies. The characters of the bones 
preserved, and certain rude but graphic representations carved on 
bones or reindeers’ antlers, enable us to know that these Horses 
were rather small in size, and heavy in build, with large heads and 
rough shaggy manes and tails, much like, in fact, the present wild 
horses of the steppes of the south of Russia. They were 
domesticated by the inhabitants of Europe before the dawn of 
history, but it is doubtful whether the majority of the animals now 
existing on the Continent are derived directly from them, as it is 
more probable that they are descendants from Horses imported 
through Greece and Italy from Asia, derived from a still earlier 
domestication, followed by gradual improvement through long- 
continued attention bestowed on their breeding and training. 
Horses are now diffused by the agency of man throughout almost 
the whole of the inhabited parts of the globe, and the great modifica- 
tions they have undergone in consequence of domestication and 
selective breeding are well exemplified by comparing such extreme 
forms as the Shetland pony, dwarfed by uncongenial climate, the 
thoroughbred racer, and the London dray-horse. In Australia, 

! Darwin, Variation of Animals and Plants under Domestication, 1868, vol. 
i. chap. ii. 


EQUIDE 383 


as in America, horses imported by the European settlers have 
escaped into the unreclaimed lands, and multiplied to a prodigious 
extent, roaming in vast herds over the plains where no hoofed 
animal ever trod before. 

A wild Horse from Central Asia, named £. prezevalskii,) is 
described as having callosities on both limbs and broad hoofs like 
E. caballus ; but the long hairs of the tail do not begin until about 
half way down its length. It also differs from #. caballus in having 
a short erect mane and no forelock; neither is there any dorsal 
stripe. The ears are of moderate size; the whole body is of a 
whitish-gray, paler beneath, and reddish on the head and upper 
parts of the limbs. If rightly described this form would appear 
to be intermediate between the true Horses and the Asses. 

The second species is the domestic Ass (£. asinus), and the wild 
Asses of Africa (LZ. asinus, var. africanus and var. somalicus?). The 
domestic Ass, which is now nearly as widely diffused and useful 
to man as the Horse, was known in Egypt long before the latter, 
and is doubtless of African origin. The ears are long, the mane 
erect, the tail without long hairs at the base, and there are no 
callosities on the hind limbs. There is a dark dorsal stripe, and 
another across the shoulders ; while the limbs are frequently banded. 
Of the wild forms the Nubian race (var. africanus) has distinct 
dorsal and shoulder stripes, but the rings on the limbs are often very 
indistinct ; while in the Somali race the dorsal stripe is indistinct, 
and the shoulder stripe wanting, but the rings on the ‘limbs are 
very boldly marked. Teeth and bones from a Pleistocene cavern 
deposit in Madras have been referred to E. asinus. 

The Asiatic wild Asses, which roam in small herds in the open 
plains of Syria, of many parts of Persia, of the north-west of India, 
and the highlands of Tartary and Tibet, from the shores of the 
Caspian to the frontiers of China, differ from the last in being of a 
more rufous or isabelline colour, instead of pure gray, in wanting 
the dark streak across the shoulder, and having smaller ears. They 
have all a dark-coloured median dorsal stripe. Though it is con- 
sidered probable by many zoologists that they form but a single 
species ? (E. hemionus), they present such marked variations in size 
and form that they have commonly been divided into three—the 
Syrian Wild Ass (£. hemippus), the Onager (£. onager) from Persia, 
Baluchistan, the Punjab, Sind, and the desert of Kach, and the 
Kiang or Dzeggetai (£. hemionus) of the high table-lands of Tibet, 
where it is usually met with at an elevation of 15,000 feet and 
upwards above the sea-level. The last is considerably larger than 


1 See Nature, 21st August 1884, and Zool. Garten. vol. xxviii. p. 453. 

2 See Sclater, Proc. Zool. Soc. 1884, p. 542. 

3 See Blanford, Zoology and Geology of Eastern Persia (Journeys of the Persian 
Boundary Commission), p. 84. 


384 UNGULATA 


either of the others, and differs from them in external appearance, 
having more the aspect of the horse. They are all remarkably 
swift, having been known to outstrip the fleetest Horse in speed. 
Lastly, there are four striped species, all inhabitants of Africa. 
These constitute the genus Hippotigris of Hamilton-Smith, but they 
are not separable except by their coloration from the true Asses, 
and one of them, the Quagga (#. quagga), may he considered as 
intermediate. This animal was formerly met with in vast herds on 
the great plains of South Africa, between the Cape Colony and the 
Vaal River, but now, in common with most of the larger wild 
animals of that region, is becoming extremely scarce, owing to the 


Fig. 160.—The Quagga (Equus quagga). 


encroachments of European civilisation, if, indeed, it is not already 
extinct. In length of ears and character of tail it more resembles 
the Horse than it does the Ass, although it agrees with the latter in 
wanting the callosity on the inner side of the hind leg, just below 
the hock, characteristic of the Horse. The colour of the head, neck 
and upper parts of the body is reddish-brown, irregularly banded 
and marked with dark brown stripes, stronger on the head and 
neck and gradually becoming fainter until lost behind the shoulder 
There is a broad dark median dorsal stripe. The under surface - 
the body, the legs, and tail are nearly white, without stripes. The 
crest is very high, surmounted by a standing mane, banded alter- 
nately brown and white. Though never really domesticated 
Quaggas have occasionally been trained to harness. The accom. 
panying figure is reduced from a painting made from one of a pair 
which were driven in Hyde Park in the early part of the present 


EQUIDE 385 


century. The name is an imitation of the shrill barking neigh of 
the animal— ouag-ga, ouag-ga,” the last syllable very much pro- 
longed. It must be remembered, however, in reading books of 
African travel that the same word is very commonly applied by 
hunters to Burchell’s Zebra. 

Of the Zebras proper, the one which was first known to Europeans, 
and was formerly considered the most common, is the True Zebra 
(£. zebra), sometimes called the Mountain Zebra. It inhabits the 
mountainous regions of the Cape Colony; but now, owing to the 
advances of civilised man into its somewhat restricted range, it has 


Fic. 161.—True or Mountain Zebra (Equus zebra). 


become very scarce, and is even, like the Quagga, threatened with 
extermination at no distant date. The second species, Burchell’s 
Zebra (£. burchelli), still roams in large herds over the plains to the 
north of the Orange River, but in yearly diminishing numbers. 
Both species are subject to considerable individual variations in 
marking, but the following are the principal characters by which 
they can be distinguished. 

£. zebra (Fig. 161) is the smaller of the two (about 4 feet high 
at the shoulders), and has longer ears, a tail more scantily clothed 
with hair, and a shorter mane. The general ground colour is white, 
and the stripes are black ; the lower part of the face is bright brown. 
With the exception of the abdomen and the inside of the thighs, the 
whole of the surface is covered with stripes, the legs having narrow 
transverse bars reaching quite to the hoofs, and the base of the tail 

25 


386 UNGULATA 


being also barred. The outsides of the ears have a white tip and 
a broad black mark occupying the greater part of the surface, but 
are white at the base. Perhaps the most constant and obvious 
distinction between this species and the next is the arrangement 
of the stripes on the hinder part of the back, where there are a 
number of short transverse bands passing from the median longi- 
tudinal dorsal stripe towards, and sometimes joining with, the 
uppermost of the broad stripes which run obliquely across the 
haunch from the flanks towards the root of the tail. There is often 
a median longitudinal stripe under the chest. 


Fic. 162.—Burchell’s Zebra (Equus burchelli). 


E. burchelli (Fig, 162) is a rather larger and more robust animal 
with smaller ears, a longer mane, and fuller tail. The general 
ground colour of the body is pale yellowish-brown, the limbs nearly 
white, the stripes dark brown or black. In the typical form they 
do not extend on to the limbs or the tail; but there is a great 
variation in this respect, even in animals of the same herd, some 
being striped quite down to the hoofs (this form has been named 
E. chapmani). There is a strongly marked median longitudinal 
ventral black stripe, to which the lower ends of the transverse side 
stripes are usually united, but the dorsal stripe (also strong] 
marked) is completely isolated in its posterior half, and the ee 
most of the broad haunch stripes runs nearly parallel to it. A 
much larger proportion of the ears is white than in the other 
species. In the middle of the wide intervals between the broad 


ELQUID.E 387 


black stripes of the flanks and haunches fainter stripes are generally 
seen. 

£. grevyii—Under this name a Zebra has been described which 
was sent in 1882 to Paris from the Galla country, lying to the 
south of Abyssinia, the most northern locality in which Zebras have 
previously been met with. In many of its characters it resembles 
E. zebra, but the stripes are much finer and more numerous than in 
the typical examples of that species, and it has a strong, black, and 
isolated dorsal stripe. Even allowing for the great variations that 
are met with in the markings of animals of this group, the aberrant 
characters of this individual are quite sufficient to separate it specific- 
ally from the true Zebra of South Africa. Other similar specimens 
have been recently brought from the Somali country. 

The flesh of the Zebras is relished by the natives as food, and their 
hides are very valuable for leather. Although the many attempts 
that have been made to break in and train these animals for riding 
or driving have sometimes been rewarded with partial success, they 
have never been domesticated in the true sense of the word. 

There are thus at least seven modifications of the Horse type at 
present existing, sufficiently distinct to be reckoned as species by 
all zoologists, and easily recognised by their external characters. 
They are, however, all so closely allied that each will, at least in a 
state of domestication or captivity, breed with perfect freedom with 
any of the others. Cases of cross breeds are recorded between the 
Horse and the Quagga, the Horse and Burchell’s Zebra, the Horse 
and the Hemionus or Asiatic wild Ass, the common Ass and the 
Zebra, the common Ass and Burchell’s Zebra, the common Ass and 
the Hemionus, the Hemionus and the Zebra, and the Hemionus and 
Burchell’s Zebra. The two species which are perhaps the farthest 
removed in general structure, the Horse and the Ass, produce, as is 
well known, hybrids or Mules, which in some qualities useful to 
man excel both their progenitors, and in some countries, and 
for certain kinds of work, are in greater requisition than either. 
Although occasional instances have been recorded of female Mules 
breeding with the males of one or other of the pure species, it is 
doubtful if any case has occurred of their breeding inter se, although 
the opportunities of doing so must have been great, as Mules have 
been reared in immense numbers for at least several thousands of 
years. We may therefore consider it settled that the different 
species of the group are now in that degree of physiological differ- 
entiation which enables them to produce offspring with each other, 
but does not permit of the progeny continuing the race, at all events 
unless reinforced by the aid of one of the pure forms. 

The several members of the group show mental differences 
quite as striking as those exhibited by their external form, and 
more than perhaps might be expected from the similarity of their 


388 UNGULATA 


cerebral organisation. The patience of the Ass, the high spirit of 
the Horse, the obstinacy of the Mule, have long been proverbial. 
It is very remarkable that, out of so many species, two only should 
have shown any aptitude for domestication, and that these two 
should have been from time immemorial the universal and most 
useful companions and servants of man, while all the others remain 
in their native freedom to this day. It is, however, still a question 
whether this really arises from a different mental constitution 
causing a natural capacity for entering into relations with man, or 
whether it may not be owing to their having been brought gradually 
into this condition by long-continued and persevering efforts when 
the need of their services was keenly felt. It is quite possible 
that one reason why most of the attempts to add new species to 
the list of our domestic animals in modern times have ended in 
failure is that it does not answer to do so in cases in which existing 
species supply all the principal purposes to which the new ones 
might be put. It can hardly be expected that Zebras and Quaggas 
fresh from their native mountains and plains can be brought into 
competition as beasts of burden and draught with Horses and Asses, 
whose naturally useful qualities have been augmented by the train- 
ing of thousands of generations of progenitors. 

Not unfrequently instances occur of domestic Horses being 
produced with a small additional toe with complete hoof, usually on 
the inside of the principal toe, and, though far more ‘rarely, three 
or more toes may be present. These malformations are often cited 
as instances of reversion to the condition of some of the earlier 
forms of equine animals previously mentioned. Such explanations, 
however plausible they appear at first sight, are nevertheless very 
doubtful. All the feet of polydactyle horses which we have 
examined bear little resemblance to those of Hipparion or Anchi- 
therium, but look rather as if due to that tendency to reduplication 
of parts which occurs so frequently as a teratological condition, 
especially among domestic animals, and, whatever its origin, certainly 
cannot in many instances, as the cases of entire limbs super- 
added, or of six digits in man, be attributed to reversion. 

Anatomy.—The anatomical structure of the Horse has been de- 
scribed in great detail in several works devoted to the subject, which 
will be mentioned in the bibliography, though these have generally 
been written from the point of view of the veterinarian rather than 
of the comparative anatomist. The limits of the present work will 
only admit of the most salient points being indicated, particularly 
those in which the Horse differs from the other Ungulata. Unless 
otherwise specified, it must be understood that all that is stated 
here, although mostly derived from observation upon the Horse, 
applies equally well to the other existing members of the group. 

Skeleton.—The skull (Fig. 163) as a whole is greatly elongated, 


EQUIDE 389 


chiefly in consequence of the immense size of the face as compared 
with the hinder or true cranial portion. The basal line of the 
cranium from the lower border of the foramen magnum to the 
incisor border of the palate is very nearly straight. The orbit, of 
nearly circular form, though small in proportion to the size of the 
whole skull, is distinctly marked, being completely surrounded by a 
strong ring of bone with prominent edges. Behind it, and freely 
communicating with it beneath the osseous bridge (the postorbital 


Fic. 163.—Side view of skull of Horse, with the bone removed so as to expose the whole of 
the teeth. Px, Premaxilla; Jf, maxilla; Na, nasal; Ma, malar or jugal; L, lachrymal; Fr, 
frontal; Sq, squamosal; Pa, parietal; oc, occipital condyle; pp, paroccipital process; i1, i2, 
and i8, the three incisors ; c, the canine; pml, the situation of the rudimentary first premolar, 
which has been lost in the lower, but is present in the upper jaw; pm?, pm3, and pm4, the 
three fully developed premolars ; m1, m2, and m8, the three true molars. 


process of the frontal) forming the boundary between them, is the 
small temporal fossa occupying the whole of the side of the cranium 
proper, and in front is the great flattened expanse of the “ cheek,” 
formed chiefly by the maxilla, giving support to the long row of 
cheek-teeth, and having a prominent ridge running forward from 
below the orbit for the attachment of the masseter muscle. The 
lachrymal occupies a considerable space on the flat surface of the 
cheek in front of the orbit, and below it the jugal or malar does 
the same. The latter sends a horizontal or slightly ascending 
process backwards below the orbit to join the under surface of the 
zygomatic process of the squamosal, which is remarkably large, and, 
instead of ending as usual behind the orbit, runs forwards to join 
the greatly developed postorbital process of the frontal, and even 


390 UNGULATA 


forms part of the posterior and inferior boundary of the orbit, an 
arrangement not met with in other mammals. The closure of the 
orbit behind distinguishes the skull of the Horse from that of the 
Rhinoceros and Tapir, and also from all of the Perissodactyles of 
the Eocene period. In front of the cerebral cavity, the great 
tubular nasal cavities are provided with well-developed turbinal 
bones, and are roofed over by very large nasals, broad behind, and 
ending in front in a narrow decurved point. The opening of the 
anterior nares is prolonged backwards on each side of the face 
between the nasals and the elongated slender premaxille. The 
latter expand in front, and are curved downwards to form the semi- 
circular alveolar border supporting the large incisor teeth. The 
palate is narrow in the interval between the incisor and cheek- 
teeth, in which are situated the large anterior palatine foramina. 
Between the cheek-teeth it is broader, and it ends posteriorly in a 
rounded excavated border opposite the hinder edge of the penulti- 
mate molar. It is mainly formed by the maxille, as the palatines 
are very narrow. The pterygoids are delicate slender slips of bone 
attached to the hinder border of the palatines, and supported 
externally by, and generally ankylosed to, the rough pterygoid 
plates of the alisphenoid, with no pterygoid fossa between. They 
slope very obliquely forwards, and end in curved, compressed, 
hamular processes. There is a distinct alisphenoid canal for the 
passage of the internal maxillary or main branch of the external 
carotid artery. The base of the cranium is long and narrow; the 
alisphenoid is very obliquely perforated by the foramen rotundum, 
but the foramen ovale is confluent with the large foramen lacerum 
medium behind. The glenoid surface for the articulation of the 
mandible is greatly extended transversely, concave from side to 
side, convex from before backwards in front, and hollow behind, and 
is hounded posteriorly at its inner part by a prominent postglenoid 
process. The squamosal enters considerably into the formation of 
the temporal fossa, and, besides sending the zygomatic process for- 
wards, it sends down behind the meatus auditorius a post-tympanic 
process which aids to hold in place the otherwise loose tympano- 
periotic bone. Behind this the exoccipital gives off a very long 
paroccipital process. The periotic and tympanic are ankylosed 
together, but not with the squamosal. The former has a wide but 
shallow floccular fossa on its inner side, and sends backwards a 
considerable “ pars mastoidea,” which appears on the outer surface 
of the skull between the post-tympanic process of the squamosal and 
the exoccipital. The tympanic forms a tubular meatus auditorius 
externus directed outwards and slightly backwards. It is not 
dilated into a distinct bulla, but ends in front in a pointed styliform 
process ; and completely embraces the truncated cylindrical tym- 
panohyal, which is of great size, in correspondence with the large 


EQUIDA 391 


development of the whole anterior arch of the hyoid. This con- 
sists mainly of a long and compressed stylohyal, expanded at the 
upper end, where it sends off a triangular posterior process. The 
basihyal is remarkable for the long, median, pointed, compressed 
“glossohyal” process, which it sends forward from its anterior 
border into the base of the tongue. A similar but less developed 
process is found in the Rhinoceros. The mandible is largely 
developed, especially the region of the angle, which is expanded 
and flattened, giving great surface for the attachment of the 
masseter muscle. The condyle is greatly elevated above the 
alveolar border ; its articular surface is very wide transversely, and 
narrow and convex from before backwards. The coronoid process 
is slender, straight, and inclined backwards. The horizontal ramus, 
long, straight, and compressed, gradually narrows towards the 
symphysis, where it expands laterally to form with the ankylosed 
opposite ramus the wide, semicircular, shallow alveolar border for 
the incisor teeth. 

The vertebral column consists of seven cervical, eighteen dorsal, 
six lumbar, five sacral, and fifteen to eighteen caudal vertebrae. 
There may be nineteen rib-bearing vertebre, in which case five 
only will be reckoned as belonging to the lumbar series. The 
odontoid process of the atlas is wide, flat, and hollowed above, as 
in the Ruminants. The bodies of the cervical vertebre are elon- 
gated, strongly keeled, and markedly opisthoccelous, or concave 
behind and convex in front. Their neural lamine are very broad, 
the spines almost obsolete, except in the seventh, and the trans- 
verse processes not largely developed. In the trunk vertebre the 
opisthoccelous character of the centrum gradually diminishes. The 
spinous processes of the anterior thoracic region are high and com- 
pressed. To these is attached the powerful elastic ligament, 
ligamentum nuche, or “paxwax,” which passing forwards in the 
middle line of the neck above the neural arches of the cervical ver- 
tebrze, to which it is also connected, is attached to the occiput and 
supports the weight of the head. The transverse processes of the 
lumbar vertebre are long, flattened, and project horizontally out- 
wards or slightly forwards from the arch. The metapophyses are 
moderately developed, and there are no anapophyses. The caudal 
vertebre, except those quite at the base, are slender and cylindrical, 
without processes and without chevron-bones beneath. The ribs 
are eighteen or nineteen in number on each side, flattened, and 
united to the sternum by short, stout, tolerably well ossified sternal 
ribs. The sternum consists of six pieces; the anterior or pre- 
sternum being extremely compressed, and projecting forwards like 
the prow of a boat. The segments which follow gradually widen, 
and the hinder part of the sternum is broad and flat. 

As in all other Ungulates, there are no clavicles. The scapula 


302 UNGULATA 


is long and slender; the suprascapular border is rounded, and 
slowly and imperfectly ossified. The spine is very slightly devel- 
oped ; rather above the middle its edge is thickened and somewhat 
turned backwards, but it gradually subsides at the lower extremity 
without forming any acromial process. The coracoid process is a 
prominent rounded nodule. The humerus is stout and rather 
short, and has a double bicipital groove. The ulna is quite rudi- 
mentary, being only represented by little more than the olecranon. 
The shaft gradually tapers below, and is firmly ankylosed to the 
radius, The latter bone is of nearly equal width throughout. The 
three bones of the first row of the carpus (the scaphoid, lunar, and 
cuneiform) are subequal in size. The second row consists of a very 
broad and flat magnum, supporting the great third metacarpal, 
having to its radial side the trapezoid, and to its ulnar side the unci- 
form, which are both small, and articulate distally with the rudi- 
mentary second and fourth metacarpals. The pisiform is large and 
prominent, flattened, and curved ; articulating partly with the 
cuneiform and partly with the lower end of the radius. The large 
metacarpal is called in veterinary anatomy “cannon-bone”; the 
small lateral metacarpals, which gradually taper towards their 
lower extremities, and lie in close contact with the large one, are 
called “ splint-bones.” The single digit consists of a moderate-sized 
proximal (0s suffraginis, or large pastern), a very short middle (os 
corone, or small pastern), and a wide, semi-lunar, ungual phalanx 
(os pedis, or coffin-bone). There is a pair of large nodular sesamoids 
behind the metacarpo-phalangeal articulation, and a single large 
transversely extended sesamoid behind the joint between the 
second and third phalanx, called the “ navicular bone.” 1 

The carpal joint, corresponding to the wrist of man, is commonly 
called the “knee” of the Horse, the joint between the metacarpal 
and the first phalanx the “fetlock,” that between the first and 
second phalanges the “astern,” and that between the second and 
third phalanges the “ coffin-joint.” 

In the hind limb the femur is marked, as in other Perisso- 
dactyles, by the presence of a “third trochanter,” a flattened process, 
curving forwards, arising from the outer side of the bone, about 
one-third of the distance from the upper end. The fibula is reduced 
toa mere styliform rudiment of the upper end; its lower part being 
absent or completely fused with the tibia. The calcaneum has a 
long and compressed calcaneal process, The astragalus has a large 
flat articular surface in front for the navicular, and a very small one 
for the cuboid. The navicular and the external cuneiform bones 
are very broad and flat. The cuboid is small, and the internal and 
middle cuneiform bones are small and united together. The meta- 
podials and phalanges resemble very closely those of the fore limb, 


1 This must not be confounded with the navicular of the tarsus. 


EQUIDE 393 


but the principal metatarsal is more laterally compressed at its 
upper end than is the corresponding metacarpal. The joint 
between the femur and tibia, corresponding to the knee of man, is 
called the “stifle joint”; while that between the tibia and tarsus, 
corresponding to the ankle of man, is termed the “hock.” The 
bones and joints of the foot have the same names as in the fore 
limb. The Horse is eminently “digitigrade,” standing on the ex- 
tremity of the single digit of each foot, which is kept habitually in 
@ position approaching to vertical. 

The muscles} of the limbs are modified from those of the ordi- 
nary mammalian type in accordance with the reduced condition of 
the bones and 
the simple re- 
quirements of 
flexion and ex- 
tension of the 
joints, no such 
actions as pro- 
nation and 
supination, or 
opposition of 
digits, being 
possible or 
needed. The 
muscles, there- 
fore, which per- 7 
form these 
functions in 
other mammals 


are absent or 
rudimentary. 
Below the 
carpal and tar- 
sal joints the 
fore and hind 
limbs corre- 
spond almost 


Fic. 164.—Section of foot of Horse. 1, Metacarpal bone; 2, first 
phalanx (0s suffraginis); 3, second phalanx (os coronce); 4, third or 
ungual phalanx (0s pedis, or coffin-bone); 5, one of the upper sesamoid 
bones; 6, lower sesamoid or ‘‘navicular” bone; 7, tendon of anterior 
extensor of the phalanges ; 8, tendon of superficial flexor (fl. perforatus) ; 
9, tendon of deep flexor (jl. perforans); 10, suspensory ligament of 
fetlock; 11, inferior or short sesamoid ligament; 12, derma or skin 
of the foot, covered with hair, and continued into 18, the coronary 
cushion, 14, the podophyllous or laminar membrane, and 15, the kera- 
togenous membrane of the sole; 16, plantar cushion ; 17, hoof; 18, fatty 
cushion of fetlock. 


exactly in struc- 
ture as well as function. On the anterior or extensor surface of 
the limb a powerful tendon (7 in Fig. 164), that of the anterior 
extensor of the phalanges (corresponding to the extensor communis 
digitorum of the arm and eatensor longus digitorum of the foot of man) 
passes down over the metacarpal bone and phalanges, to be inserted 
mainly into the upper edge of the anterior surface of the last phalanx 


1 Want of space and of the necessary illustrations rendered it impossible to 
give an account of mammalian myology in the earlier chapters of this work. 


394 UNGULATA 


or pedal bone. There is also a much smaller second extensor on 
the outer side of this in each limb, the lateral extensor of the 
phalanges. In the fore leg the tendon of this muscle (which corre- 
sponds with the extensor minimi digiti of man) receives a slip from 
that of the principal extensor, and is inserted into the first phalanx. 
In the hind leg (where it is the homologue apparently of the 
peroneus brevis of man) the tendon becomes blended with that of the 
large extensor. 

A very strong ligamentous band behind the metapodium, 
arising from near the upper extremity of its posterior surface, 
divides into two at its lower end, and each division, being first 
connected with one of the paired upper sesamoid bones, passes by 
the side of the first phalanx to join the extensor tendon of the 
phalanges. This is called in veterinary anatomy the “suspensory 
ligament of the sesamoids,” or of the “ fetlock ” (10 in Fig. 164) ; but 
its attachments and relations, as well as the occasional presence of 
muscular fibres in its substance, show that it is the homologue of 
the short flexor muscle of other mammals, curiously modified both 
in structure and function to suit the requirements of the Horse’s 
foot. Behind or superficial to this are placed the two strong tendons 
of the long flexor muscles, the most superficial, or flexor perforatus 
(8), dividing to allow the other to pass through, and then inserted 
into the middle phalanx. The flexor perforans (9) is as usual in- 
serted into the terminal phalanx. In the fore leg these muscles 
correspond with those similarly named in man. In the hind leg, 
the perforated tendon is a continuation of that of the plantaris, 
passing pulley-wise over the tuberosity of the caleaneum. The 
perforating tendon is derived from the muscle corresponding with 
the long flexor of man, and the smaller tendon of the oblique flexor 
(tibialis posticus of man) is united with it. 

The hoof of the Horse corresponds to the nail or claw of other 
mammals, but is so constructed as to form a complete and very 
solid case to the expanded termination of the toe, giving a firm 
basis of support formed of a nonsensitive substance, which is con- 
tinually renewed by the addition of material from within as its 
surface wears away by friction against the ground. The terminal 
phalanx of the toe is greatly enlarged and modified in form to sup- 
port this hoof, and the size of the internal framework of the foot is 
further increased by a pair of lateral fibro-cartilaginous masses 
attached on each side to the hinder edges of the bone, and by a 
fibro-cellular and adipose plantar cushion in the median part. 
These structures are all enclosed in the keratogenous membrane or 
“subcorneous integument,” a continuation of the ordinary derma of 
the limb, but extremely vascular, and having its superficial extent 
greatly increased by being developed into papille or lamine. From 
this the horny material which constitutes the hoof is exuded. A 


EQUIDA 395 


thickened ring encircling the upper part, called coronary cushion 
(13), and the sole (15), are covered with numerous thickly set 
papille or villi, and take the greatest share in the formation of the 
hoof ; the intermediate part constituting the front and side of the 
foot (14), corresponding with the wall of the hoof, is covered with 
parallel, fine longitudinal lamine, fitting into corresponding depres- 
sions in the inner side of the horny hoof. 

The horny hoof is divided into a wall or crust consisting of the 
front and sides, the flattened or concave sole, and the “frog,” a 
triangular median prominence, notched posteriorly, with the apex 
turned forwards, situated in the hinder part of the sole. It is 
formed of pavement epithelial cells, mainly grouped in a concentric 
manner around the vascular papille of the keratogenous membrane, 
so that a section near the base of the hoof, cut transversely to the 
long axis of these papille, shows a number of small circular or oval 
orifices, with cells arranged concentrically round them. The nearer 
the surface of the hoof, or farther removed from the seat of growth, 
the more indistinct the structure becomes. 

Small round or oval plates of horny epidermis called “ chest- 
nuts,” growing like the hoof from enlarged papille of the skin, are 
found on the inner face of the fore limb, above the carpal joint, in 
all species of Hquide, and in the Horse (E. caballus) alone similar 
formations occur near the upper extremity of the inner face of the 
metatarsus. Their use is unknown. 

Behind the joint between the metapodium and the first phalanx 
is a prominence formed by the fatty cushion of the fetlock (18 in 
Fig. 164). On the middle of this is a small bare patch covered 
with thickened epidermis, the ergot or spur, generally concealed 
beneath the long hair which grows around it. This is the function- 
less vestige of the large callous pad found in this situation in the 
Tapir, and in fact in all mammals in which this part reaches the 
ground in walking. 

Dentition.—The dentition of the Horse, when all the teeth are 
in place, is, as stated before, expressed by the formula i 3, ¢ 4, p 4, 
m % = 42. The incisors of each jaw are placed in close contact, 
forming a semicircle. The crowns are broad, somewhat awl- 
shaped, and of nearly equal size. They have all the great peculi- 
arity, not found in the teeth of any other living mammal, of an 
involution of the external surface of the tooth (see Fig. 165) 
forming a deep fossa or pit, the bottom of which becomes partially 
filled up with cement. As the tooth wears, the surface, besides 
the external enamel layer as in an ordinary simple tooth, shows 
in addition a second inner ring of the same hard substance sur- 
rounding the pit, thus of course adding greatly to the efficiency 
of the tooth as an organ for biting tough, fibrous substances. This 
pit, generally filled in the living animal with particles of food, is 


396 UNGULATA 


conspicuous from its dark colour, and constitutes the “mark ” by 
which the age of the horse is judged, as in consequence of its 
extending only to a certain depth, it becomes obliterated as the 
crown wears away, when the tooth assumes the character of an 
ordinary incisor, consisting only of a core of dentine surrounded 
by the external enamel 
layer. It is not quite so 
deep in the lower as in 
the upper teeth. The 
canines are either quite 
rudimentary or entirely 
absent in the female. In 
the male they are com- 
pressed, pointed, and 
smaller than the incisors, 
from which they are 
separated by a slight in- 
terval. The teeth of the 
cheek series are all in 
contact with each other, 
but separated from the 
canines by a considerable 
toothless space. The 
anterior premolars are 


Fic. 165.—Longitudinal and transverse section of upper Quite rudimentary, often, 
incisor of Horse. p, Pulp cavity ; 4, dentine or ivory ; e, especially in the lower 
enamel ; c, outer layer of cement; c’, inner layer of cement, - 
lining a, the pit or cavity of the crown of the tooth. Jaw, not developed at all, 

and generally fall by the 
time the animal attains maturity, so that there are but six func- 
tional grinding teeth—three that have predecessors in the milk- 
dentition, and hence are considered as premolars, and three true 
molars, but otherwise, except the first and last of the series, 
not distinguishable in form or structure. These teeth in both 
upper and lower jaws are extremely long-crowned or hypsodont 
(Fig. 158), successive portions being pushed out as the sur- 
face wears away ;—a process which continues until the animal 
becomes advanced in age. The enamelled surface is infolded ina 
complex manner (a modification of that found in other Perissodac- 
tyles, see Figs. 155, 167), the folds extending quite to the base of 
the crown, and the interstices being filled and the surface covered 
with a considerable mass of cement, which binds together and 
strengthens the whole tooth. As the teeth wear, the folded enamel, 
being harder than the other constituents—the dentine and cement 
—forms projecting ridges on the surface arranged in a definite 
pattern, which give it great efficiency as a grinding instrument (see 
Fig. 157, b and c). The free surfaces of the upper teeth are 


EQUIDAS 307 


quadrate, except the first and last, which are nearly triangular. 
The lower teeth are much narrower than the upper. 

The milk dentition consists of 7 3, ¢ 9, m 3 = 24,—the canines 
and first or rudimentary premolars having apparently no pre- 
decessors. In form and structure they much resemble the 
permanent teeth, having the same characteristic enamel-foldings. 
Their eruption commences a few days after birth, and is complete 
before the end of the first year, the upper teeth usually appear- 
ing somewhat earlier than those of the lower jaw. The first 
teeth to appear are the first and second milk-molars (about 
five days), then the central incisor (from seven to ten days) ; this 
is followed by the second incisor (at one month), then by the third 
molar, and finally hy the third incisor. Of the permanent teeth the 
first true molar appears a little after the end of the first year, 
followed by the second molar hefore the end of the second year. At 
about two and a half years the first premolar replaces its predecessor. 
Between two and a half and three years the first incisor appears. 
At three years the second and third premolars and the third true 
molar have appeared ; at from three and a half to four years the 
second incisor; at four to four and a half years the canine; and, 
finally, at tive years the third incisor, completing the permanent 
dentition. Up to this period the age of the horse is clearly shown 
by the state of the dentition, and for some time longer indications 
can be obtained from the wear of the incisor teeth, though this 
depends to a certain extent upon the hardness of the food or other 
aceidental circumstances. As a general rule, the depression caused 
by the infolding of the surface of the incisor (the “mark ”) is 
obliterated in the first or central incisor at six years, in the second 
at seven years, and in the third at eight years. In the upper teeth, 
as the depressions are deeper, this obliteration does not take place 
until about two years later. After this period no certain indica- 
tions can be obtained of the age of the horse from the teeth. 

Digestive Organs.—The lips are flexible and prehensile. The 
membrane that lines them and the cheeks is quite smooth. The 
palate is long and narrow; its mucous surface has seventeen pairs 
of not very sharply defined oblique ridges, extending as far hack as 
the last molar tooth, beyond which the velum palati extends for 
about 3 inches, having a soft corrugated surface, and ending 
posteriorly in an arched border without uvula. This embraces the 
base of the epiglottis, and shuts off all communication between 
the cavity of the mouth and the nasal passages, respiration 
being, under ordinary circumstances, carried — on exclusively 
through the nostrils. © Between the mucous membrane and 
the bone of the hard palate is a dense vascular and nervous 
plexus. The membrane lining the fauces is soft and corrugated. 
An clongated raised glandular mass, 3 inches long and 1 inch from 


398 UNGULATA 


above downwards, extending backwards from the root of the tongue 
along the side of the fauces, with openings on the surface leading 
into crypts with glandular walls, represents the tonsil. The tongue, 
corresponding to the general form of the mouth, is long and narrow. 
It consists of a compressed intermolar portion with a flat upper 
surface, broad behind and becoming narrower in front; and of a 
depressed anterior part rather shorter than the former, which 
is narrow behind but widens towards the evenly rounded apex. 
The dorsal surface generally is very soft and smooth. There are 
two large circumvallate papille near the base, rather irregular in 
form, about a quarter of an inch in diameter and half an inch apart. 
The conical papille are very small and close set, though longer and 
more filamentous on the intermolar portion. There are no fungi- 
form papilla on the dorsum, but a few not very conspicuous ones 
scattered along the sides of the organ. 

Of the salivary glands the parotid is by far the largest; elongated 
in the vertical direction, and narrower in the middle than at either 
upper or lower extremity. Its upper extremity embraces the lower 
surface of the cartilaginous ear-conch ; its lower end reaches the 
level of the inferior margin of the mandible, along the posterior 
margin of which it is placed. Its duct leaves the inferior anterior 
angle, at first descends a little, and runs forward under cover of 
the rounded inferior border of the mandibular ramus, then curves 
up along the anterior margin of the masseter muscle, becoming 
superficial, pierces the buccinator, and enters the mouth by a simple 
aperture opposite the middle of the crown of the third premolar 
tooth. It is not quite so thick as a goose-quill when distended, and 
nearly a foot in length. 

The submaxillary gland is of very similar texture to the last, 
but much smaller ; it is placed deeper, and lies with its main axis 
horizontal. It is elongated and slender, and flattened from within 
outwards. Its posterior end rests against the anterior surface of 
the transverse process of the atlas, from which it extends forwards 
and downwards, slightly curved, to beneath the ramus of the jaw. 
The duct which runs along its upper and internal border passes 
forwards in the usual course, lying in the inner side of the sublingual 
gland, to open on the outer surface of a distinct papilla, situated 
on the floor of the mouth, half an inch from the middle line, and 
midway between the lower incisor teeth and the attachment of the 
frenum lingue. The sublingual is represented by a mass of vlands 
lying just beneath the mucous membrane of the floor of the mouth 
on the side of the tongue, causing a distinct ridge, extending from 
the frenum backwards, and the numerous ducts opening separately 
along the summit of the ridge. The buccal glands are arranged 
in two rows parallel with the molar teeth. The upper ones 
are the largest, and are continuous anteriorly with the labial 


EQUIDA 399 


glands, the ducts of which open on the mucous membrane of the 
upper lip. 

The stomach of the Horse is simple in its external form, with 
a largely developed right cul de ser, and is a good deal curved 
on itself, so that the cardiac and pyloric orifices are brought near 
together, The antrum pyloricum is small and not very distinctly 
marked off. The interior is divided by the character of the lining 
membrane into two very distinct portions, right and left. Over 
the litter the dense white smooth epithelial lining of the oesophagus 
is continued, terminating abruptly by a raised crenellated border. 
Over the right part (rather the larger portion) the mucous membrane 
has a grayish-red colour and a velvety appearance, and contains very 
numerous peptic glands, which are wanting in the cardiac portion. 
The cesophayenl orifice is very small, and is guarded by a strong 
crescentic or rather horse-shoe-like hand of muscul:w fibres, which is 
supposed to be the cause of the difficulty of vomiting in the Horse. 
The small intestine is of great length (80 to 90 feet), its mucous 
membrane being covered with numerous fine villi, The cecum is 
of conical form, about 2 feet long and nearly a foot in diameter ; 
its walls are sacculated, especialy near the base, having four longi- 
tudinal fibrous bands ; and its capacity is about twice that of the 
stomach, It lies with its hase near the lower part of the abdomen, 
wnd its apex directed towards the thorax. The colon is about one- 
third the length of the small intestine, and very capacious in the 
greater part of its course. As usual, it may be divided into an 
ascending, transverse, and descending portion; but the middle or 
transverse portion is folded into a great loop, which descends as low 
as the pubis ; so that the colon forms altogether four folds, generally 
parallel to the long axis of the body. The descending colon is much 
narrower than the rest, and not sacculated, and being considerably 
longer than the distance it has to traverse, is thrown into numerous 
folds. 

The liver (Fig. 166) is tolerably symmetrical in its general 
arrangement, being divided nearly equally into segments by a well- 
marked umbilical fissure. Each segment is again divided by lateral 
lissures, which do not extend quite to the posterior border of the 
orgin; of the central lobes thus cut off, the right is rather the larger, 
and has two fissures in its free border subdividing it into lobules. 
The extent of these varies, however, in different individuals, being 
not usually so marked as in the figure, which is from a feetal 
specimen. The two lateral lobes are subtriangular in form. The 
Spigelian lobe is represented by a flat surface between the portal 
fissure and the posterior border, not distinctly marked off from the 
left lateral by a fissure of the ductus venosus, as this vessel is buried 
deep in the hepatic substance, but the caudate lobe is distinct and 
tonguc-shaped, its free apex reaching nearly to the border of the 


400 UNGULATA 


right lateral lobe. In most works on the anatomy of the Horse this 
has been confounded with the Spigelian lobe of man. There is no 
gall-bladder (as in 
all other Perisso- 
dactyles), and the 
biliary duct enters 
the duodenum 
about 6 inches from 
the pylorus. The 
pancreas has two 
lobes or branches— 
a long one passing 
to the left and 
reaching thespleen, 
and a shorter right 
lobe. The principal 
duct enters the 
duodenum with the 
bile-duct, and there 
is often a second 


Fic. 165.—Uniler surface of the liver of the Horse. wu, Umbilical small duct which 
fissure ; lJ, left lateral lobe; lc, left central lobe; rc, right central opens separatel 
lobe; rl, right lateral lobe; s, Spigelian lobe; c, caudate lobe. pe ep y 

near to this. 


Circulatory and Resyirutory Organs.—The heart has the form of a 
rather elongated and pointed cone. There is one anterior vena cava, 
formed by the union of the two jugular and two axillary veins. 
The aorta gives off a large branch (the anterior aorta) very near its 
origin, from which arise—first, the left axillary, and afterwards the 
right axillary and the two carotid arteries. 

Under ordinary circumstances the Horse breathes entirely by 
the nasal passages, the communication between the larynx and the 
mouth being closed by the velum palati. The nostrils are placed — 
laterally, near the termination of the muzzle, and are large and 
very dilatable, being bordered by cartilages upon which several 
muscles act. Immediately within the opening of the nostril, the 
respiratory canal sends off on its upper and outer side a diverti- 
culum or blind pouch (called “false nostril”) of a conical form, and 
curved, 2 to 3 inches in depth, lying in the notch formed between 
the nasal and premaxillary bones. It is lined by mucous mem- 
brane continuous with that of the nasal passage, but its use is not 
apparent. It is longer in the Ass than in the Horse. A similar 
structure is found in the Rhinoceros, and in a much more developed 
condition in the Tapir. Here may be mentioned the guttural pouches, 
large air sacs, diverticula from the Eustachian tubes, and lying 
behind the upper part of the pharynx. These are likewise found 
in other Perissodactyles, but their use is also still not clearly 


EQUIDA 4ol 


understood. The larynx has the lateral sacculi well developed, 
though entirely concealed within the ale of the thyroid cartilage. 
The trachea divides into two bronchi, one for each lung. 

Nervous System.—The brain differs little, except in details of 
arrangement of convolutions, from that of other Ungulates. The 
cerebral hemispheres are rather elongated and subcylindrical, the 
olfactory lobes are large and project freely in front of the hemi- 
spheres, and the greater part of the cerebellum is uncovered. The 
eye is provided with a nictitating membrane or third eyelid, at the 
base of which the ducts of the Harderian gland open. 

Reproductive System.—The testes are situated in a distinct sessile 
or slightly pedunculated scrotum, into which they descend from the 
sixth to the tenth month after birth. The accessory generative 
glands are the two vesicule seminales, with the median third vesicle, 
or uterus masculinus, lying between them, the single bilobed pro- 
state, and a pair of globular Cowper’s glands. The penis is large, 
cylindrical, with a truncated, expanded, flattened termination. 
When in a state of repose it is retracted by a muscle arising from 
the sacrum, within the prepuce, a cutaneous fold attached below the 
symphysis pubis. 

The uterus is bicornuate. The vagina is often partially divided 
by a membraneous septum or hymen. The mamme (as in other 
members of the suborder), are two, inguinally placed. The surface of 
the chorion is covered evenly with minute villi, constituting a diffuse 
non-deciduate placenta. The period of gestation is eleven months. 


Bibliography.—M. 8. Arloing, ‘Organisation du pied chez le cheval,” Ann. 
Set. Nat, 1867, viii. pp. 55-81; H. Burmeister, Los caballos fosiles de la Pumpu 
Argentina, Buenos Ayres, 1875; Chauveau and Arloing, Zratté d’anatomie com- 
parée des animaux domestiques, Paris, 1871, and English edition by G. Fleming, 
1873; E. Cuyer and E. Alix, Le Cheval, 1886; A. Ecker, ‘‘ Das Europiische Wild- 
pferd und dessen Beziehungen zum domesticirten Pferd,’ Globus, Bd. xxxiv. 
Brunswick, 1878 ; Forsyth-Major, ‘‘ Beitrage zur Geschichte der fossilen Pferde 
besonders Italiens,” 4h. Schw. Pal. Ges. iv. pp. 1-16, pt. iv.; George, “ Etudes 
zool. sur les Hémiones et quelques autres espéces chevalines,” Ann. Sci. Nat. 
1869, xii. p. 5; E. F. Gurlt, dnatomische Abbildungen der Hausscugethicre, 1824, 
and Hand. der vergleich, Anat. der Hausstiugethicre, 2 vols. 1822; Huet, ‘ Croise- 
ment des diverses espéces du genre cheval,” Nowv. Archives du Muséum, 2d sér. 
tom. ii. p. 46, 1879; Leisering, Atlas der Anatomie des Pferdes, Leipsic, 1861 ; 
J. M‘Fadyean, The Anatomy of the Horse, 1884; O. C. Marsh, ‘‘ Notice of New 
Equine Mammals from the Tertiary Formation,” din. Journ. of Science and Arts, 
vol. vii. March 1874; Id. ‘‘ Fossil Horses in America,” Amer. Naturalist, vol. 
viii. May 1874 ; Id. ‘‘ Polydactyle Horses,” lim. Journ. of Science and Arts, vol. 
xvii. June 1879 ; Franz Miiller, Lehrbuch der Anatomie des Pferdes, Vienna, 1853 ; 
R. Owen, ‘‘Equine Remains in Cavern of Bruniquel,” Phil. Trans. vol. clix. 
(1870), p. 585; W. Percivall, The Anatomy of the Horse, 1832; G. Stubbs, 
Anatomy of the Horse, 1766. F. H. Huth’s Bibliographical Record of Hippology 
(1887) contains a list of nearly four thousand works on Horses and Equitation, 
published in the various languages of the civilised world. 

26 


402 UNGULATA 


Family RHINOCEROTIDA. 


Although the existing members of this family are readily dis- 
tinguished from the other living representatives of the suborder 
by the simple crescentoid form assumed by the ridges of the lower 
cheek-teeth, yet it is exceedingly difficult to give a definition by 
which they can be distinguished from the Lophiodontide, from some 
members of which they are, indeed, probably derived. The outer 
columns of the upper molars (Fig. 167) are, however, so excessively 
flattened as to produce 
a continuous thick and 
nearly straight outer 
wall, which is often pro- 
duced in advance of 
the anterior transverse 
ridge ; both transverse 
ridges being but little 
curved, and intimately 
connected with the 
outer wall. The upper 
premolars are in most 
cases nearly or quite as 
complex as the molars, 
and the ridges of the 
lower cheek-teeth are 
crescentoid. The last 
lower molar has no 
third lobe. The height 
of the crowns of the 
/ cheek-teeth is variable. 
Fic. 167.—A partially worn second right upper molar of The skull is large, with 


Rhinoceros untiquitatis. Letters as in Fig. 155 (p. 875), ex- the orbit confluent with 
cept k, which indicates a prolongation of the median valley. the temporal fossa 


(After Owen.) : 
wo There are either three 
or four digits in the manus, and three in the pes. One or more 


dermal horns are attached to the fronto-nasal region of the skull 
of existing forms, but these were wanting in some of the fossil 
species. 

Lhinoceros.'—Incisors variable, reduced in ‘number, often quite 
rudimentary, and early deciduous. Upper canines absent. Molar 
series, consisting of the full number of four premolars and three 
molars above and below, all in contact and closely resembling each 
other, except the first, which is much smaller than the rest and 
often deciduous ; and the last, in which the hinder lobe is partly 


1 Linn. Syst, Nat. 12th ed. vol. i. p. 104 (1766) 


RHAHINOCEROTIDE #68 


aborted, so that the contour of the crown is triangular. Head 
large, skull elongated, elevated posteriorly into a transverse occi- 
pital crest. No postorbital processes. Nasal bones large and stout, 
co-ossified, and standing out freely above the premaxille, from which 
they are separated by a deep and wide fissure; the latter small, 
generally not meeting in the middle line in front, often quite rudi- 
mentary. Tympanics small, not forming a bulla. Brain cavity very 
small for the size of the skull. Vertebre: C 7, D 19-20, L 3, 
S 4, C about 22. Limbs stout, and of moderate length. Three 
completely developed toes, with distinct broad rounded hoofs on each 
foot (Fig. 151, p. 36S), some fossil forms having a fourth in the 
manus. Eyes small. Ears of moderate size, oval, erect, prominent, 
placed near the occiput. Skin very thick, in many species thrown 
into massive folds) Hairy covering scanty. When one horn is 
present it is situated over the conjoined nasal bones ; when two, the 
hinder one is over the frontals. These horns, which are of a more 
or less conical form and usually recurved, often grow to a great 
length (three or even four feet), and are composed of a solid mass 
of hardened epidermic cells growing from a cluster of long dermal 
papilla. The cells formed on each papilla constitute a distinct 
horny fibre, like a thick hair, and the whole are cemented together 
by an intermediate mass of cells which grow up from the interspaces 
between the papille. It results from this that the horn has the 
appearance of a mass of agglutinated hairs, which, in the newly 
srowing part at the base. readily fray out on destruction of the 
softer intermediate substance ; but the fibres differ from true hairs in 
growing from a free papilla of the derm, and not within a follicular 
involution of the same. mE 

The large lower cutting - 
teeth of the typical Rhino- 
ceroses have been very gener- 
ally regarded as incisors, but 
comparison with fossil allied 
types, in which three lower in- ¢ 
cisors and canines are present, 
leaves little doubt but that 
they are really canines. The 
upper molarteethpresent some 
amount of specific variation ; Lae nem ese Sere ied 
thus while one type (Fiz. pee = ecuigies anise tee eee 
168, .4) has only a’ simple crotener; 2, r 
* erotchet ” projecting from valley; |, anterior intermediate column. Ozher 
the posterior transverse ridge "=" FS He PS 
into the median valley, in others (Fig. 168, £) this crotehet joins a 
“erista,” or “combing-plate.” projecting from the outer wall to cut 
off a distinet fossette from the median valley. Occasionally, however 


4O4 UNGULATA 


(as in Fig. 167), the crotchet and combing-plate do not completely 
join, although the fossette is distinctly indicated. The first upper 
premolar may occasionally be preceded by a milk-tooth. The Rhino- 
ceroses differ from the Horses and agree with the Tapirs in the 
direction of the cecum. 

The living species of Rhinoceros are all animals of large size, but of 
little intelligence, generally timid in disposition, though ferocious when 
attacked and brought to bay, using the nasal horns as weapons, by 
which they strike and toss their assailant. Their sight is dull, but 
their hearing and scent are remarkably acute. They feed on herbage, 
shrubs, and leaves of trees, and, like so many other large animals 
which inhabit hot countries, sleep the greater part of the day, being 
most active in the cool of the evening or even during the night. 
They are fond of bathing and wallowing in water or mud. None 
of the species have been domesticated. Animals of the group have 
existed in both the Old and New Worlds since the latter part of 
the Eocene period. In America they all became extinct before the 
end of the Pliocene period. In the Old World their distribution 
has become greatly restricted, and they are no longer found in 
Europe and North Asia, but only in Africa and portions of the 
Indian and Indo-Malayan regions. 

Existing Species.—The existing (as well as many of the extinct) 
species of Rhinoceroses naturally divide into three groups, which are 
regarded by some zoologists as of generic value. 

Lthinocerotic, or Typical Group.—The adults with a single large 
compressed incisor above on each side, and occasionally a small lateral 
one; below, a very small incisor and a very large, procumbent, 
pointed canine. Nasal bones pointed in front. A single nasal 
horn. Skin very thick, and raised into strong, definitely arranged 
ridges or folds. 

There are two well-marked species of one-horned Rhinoceroses. 
(1) The Indian Rhinoceros, R. wnicornis (Fig. 169) of Linneus,! the 
largest and best known, from being the most frequently exhihited 
alive in England, is at present only met with in a wild state in the 
terai region of Nipal and Bhutan, and in the upper valley of the 
Brahmaputra or province of Assam, though it formerly had a wider 
range. The first Rhinoceros seen alive in Europe since the time 
when these animals, in common with nearly all the large remark- 
able beasts of both Africa and Asia, were exhibited in the Roman 


? Many authors use Cuvier’s name, R. indicus, in preference to this, on the 
ground that there are more than one species with one horn, forgetting that the 
name substituted is equally inconvenient, as more than one species live in India. 
The fact of a specific name being applicable to several members of a genus is no 
objection to its restriction to the first to which it was applied ;_ otherwise 
changes in old and well-received names would constantly have to be made in 
consequence of new discoveries. 


RHINOCEROTIDE 405 


shows, was of this species. It was sent from India to Emmanuel, 
King of Portugal, in 1513; and from a sketch of it, taken in 
Lisbon, Albert Diirer composed his celebrated but rather fanciful 
engraving, which was reproduced in so many old books on natural 
history. Both in this and the following species the post-glenoid 
and post-tympanic processes of the squamosal bone of the skull 
unite below so as to completely surround the external auditory 
meatus. The molar teeth are hypsodont, and have a horizontal 
plane of wear; those of the upper jaw (Fig. 168, 0) being charac- 
terised by the presence of a combing-plate joining the crotchet, and 


Fie. 169.—Indian Rhinoceros (Rhinoceros wricornis). This figure, and also figures 170, 172, 
are reduced from drawings by J. Wolf, from animals living in the London Zoological Society’s 
Gardens. 
the absence of a distinct buttress at the antero-external angle. 
The stomach departs from the ordinary Perissodactyle type. The 
small intestine is beset over most of its surface with long and fine 
villi; and the Spigelian lobe of the liver is well developed. There 
is a gland behind the foot. Teeth from the Pleistocene of the 
Narbada valley in India apparently indicate the existence of the 
Indian Rhinoceros at that epoch. (2) The Javan Rhinoceros (R. 
sondaicus, Fig. 170) is a smaller form, readily distinguished by 
dental and internal characters, as well as by the different arrange- 
ment of the plications of the skin (as seen in the figures) ; the horn 
in the female appears to be very little developed, if not altogether 
absent. This species has-a more extensive geographical range, 
being found in the Bengal Sunderbans near Calcutta, Burma, the 
Malay Peninsula, Java, Sumatra, and probably Borneo. The molar 
teeth have shorter crowns than in the preceding species, and wear 
into ridges ; those of the upper jaw (Fig. 168, «) having no combing- 


406 UNGULATA 


plate, and a strongly-marked buttress at the antero-external angle 
(not distinctly shown in the figure). The visceral anatomy, accord- 
ing to Beddard,! does not differ materially from that of the next 
species. In respect to its dentition and anatomical characters 
this species is indeed more nearly allied to the Sumatran than to 
the Indian Rhinoceros; and therehy indicates that the division of 
the existing Rhinoceroses into separate genera is not advisable. 


Fic. 170.—Javan Rhinoceros (Rhinoceros sondaicus). 


Ceratorhine Group.—The adults with a moderate-sized compressed 
incisor above, and a laterally placed, pointed, procumbent canine 
below, which is sometimes lost in old animals. Nasal bones narrow 
and pointed anteriorly. A well-developed nasal, and a small frontal 
horn separated by an interval. The skin thrown into folds, but 
these not so strongly marked as in the former group. The 
smallest living member of the family, the Sumatran Rhinoceros, R. 
sumatrensis, Cuvier, now represents this group. Its geographical 
range is nearly the same as that of the Javan species, though not 
extending into Bengal; but it has been found in Assam, Chittagong, 
Burma, the Malay Peninsula, Sumatra, and Borneo. So far as 
can be determined during the life of the type specimen, it appears 
that the hairy form from Chittagong, described as R. Jusiotis, is only 
a variety of this species.2 The molar teeth of the Sumatran Rhino- 
ceros are almost indistinguishable from those of the Javan species, 


1 Trans. Zool. Soc. vol. xii.; see also Proc. Zool, Soc. 1889, p. 9. 
? See Beddard and Treves, Proc. Zool. Soc. 1889, Pp. 9 


RHINOCEROTIDAE 407 


and reference has already been made to the resemblance between 
the visceral anatomy of these species! The form of the stomach 
is very similar to that of the Horse. The liver (Fig. 171) has a 
comparatively large caudate lobe, but is chiefly remarkable for the 
peculiar shape of the Spigelian lobe, which mainly consists of a thin 
strip of tissue, 8 inches long, ? inch wide, and 4 inch deep. The 
small intestine, in place of the villi of 2. wnicornis, has throughout 
the greater part of its length a uniform series of thin and nearly or 
quite continuous transverse foldings, like the valvule conniventes 
of the human small intestine. There is no gland behind the foot. 


u SS) 


Fic. 171.—Posterior aspect of the liver of Rhinoceros sumatrensis. re, Right central lobe ; 
rl, right lateral lobe ; lec, left. central lobe ; 1, left lateral lobe; c, caudate lobe; sp, Spigelian 
lobe. (From Garrod, Proc. Zool. Soc. 1873, p. 102.) 


The post-glenoid and post-tympanic processes of the squamosal do 
not unite below the auditory meatus. The presence of a lateral 
nasal diverticulum, like that of the Horses and Tapirs, has been 
verified only in this species, although it doubtless occurs in the 
others. 

Atelodine Grouwp—In the adults the incisors and canines quite 
rudimentary or entirely wanting. Nasal bones thick, rounded and 
truncated in front. Well-developed anterior and posterior horns in 
close contact. Skin without any definite permanent folds. 

The two well-marked existing species are peculiar to the African 
continent. 


1 For the internal anatomy of R. swmatrensis see Garrod, Proc. Zool. Soc. 
1878, p. 92; and Beddard and Treves, loc. cit. 


408 UNGULATA 


The common Two-horned Rhinoceros, P. bicornis, is the smaller of 
the two, with a pointed prehensile upper lip, and a narrow compressed 
deep symphysis of the lower jaw. It ranges through the wooded 
and watered districts of Africa, from Abyssinia in the north to the 
Cape Colony, but its numbers are yearly diminishing, owing to the 
inroads of European civilisation, and especially of English sports- 
men. It feeds exclusively upon leaves and branches of bushes and 
small trees, and chiefly frequents the sides of wood-clad rugged 
hills. Specimens in which the posterior horn has attained a length 


Fic. 172.—Common African Rhinoceros (Rhinoceros bicornis). 


as great as, or greater than, the anterior have been separated under 
the name of &. keitloa, but the characters of these appendages are 
too variable to found specific distinctions upon. The Common 
African Rhinoceros is far more rarely seen in menageries in Europe 
than either of the three Oriental species, but one has lived in the 
gardens of the London Zoological Society since 1868. The molar 
teeth of this species are of the general type of those of R. sondaicus, 
having no combing-plate to join the crotchet in those of the upper 
jaw. The conch of the ear is much rounded at its extremity, and 
edged by a fringe of short hairs; while the nostrils are somewhat 
rounded. The eye is placed immediately below the posterior 
horn.’ Both in this and the following species the post-glenoid and 
post-tympanic processes of the squamosal do not unite below the 


1 These external points of distinction from R. simus are taken from a paper 
by Sclater in the Proc. Zool. Soc. 1886, p. 143. 


RHINOCEROTIDA 409 


auditory meatus. Nothing is known of the anatomy of the soft 
parts of cither of them. 

Burchell’s or the Square-mouthed Rhinoceros (L¢. s/ius), sometimes 
called the White Rhinoceros, though the colour (dark slate) is not 
materially different from that of the last species, is the largest of 
the whole group, and differs from all the others in having a square 
truncated upper lip and a wide, shallow, spatulate symphysis to 
the lower jaw. In conformity with the structure of the mouth, 
this species lives entirely by browsing on grass, and is therefore 
more partial to open countries or districts where there ave broad 
grassy valleys between the tracts of bush. It is only found in 
Africa south of the Zambesi, and of late years has become ex- 
tremely scarce, owing to the persecutions of sportsmen ; indeed, 
the time of its complete extinction cannot be far off. No specimen 
of this species has ever been brought alive to Europe. Myr. F. C. 
Selous! gives the following description of its habits from extensive 
porsonal observation :— 

“The square-mouthed rhinoceros is a huge ungainly -looking 
beast, with a disproportionately large head, a large male standing 
6 feet 6 inches at the shoulder. Like elephants and buttaloes they 
lie asleep during the heat of the day, and feed during the night 
and in the cool how's of early morning and evening. Their sight 
is very bad; but they ave quick of hearing, and their scent is very 
keen ; they ave, too, often accompanied by rhinoceros birds, which, 
by running about their heads, flapping their wings, and screeching 
at the same time, frequently give them notice of the approach of 
danger. When disturbed they go off at a swift trot, which soon 
leaves all pursuit from a man on foot far behind ; but if chased by 
a horseman they break into a gallop, which they can keep up for 
some distance. However, although they run very swiftly, when 
their size and heavy build is considered, they are no match for an 
average good horse. They are, as a rule, very easy to shoot on 
horseback, as, if one gallops a little in front of and on one side of 
them, they will hold their course, and come sailing past, offering 
au magnificent broadside shot, while under similar circumstances a 
prehensile-lipped rhinoceros will usnally swerve away in such a 
manner as only to present his hind-quarters for a shot. When 
either walking or running, the square-mouthed rhinoceros holds its 
head very low, its nose nearly touching the ground. When a small 
ealf accompanies its mother it always runs in front, and she appears 
to guide it by holding the point of her horn upon the little animal’s 
rump; and it is perfectly wonderful to note how in all sudden 
changes of pace, from a trot to a gallop or rice cers’, the same 
position is always exactly maintained. During the autumn and 
winter months (i.e. from March to August) the square-mouthed 

1 Prov, Zool. Soc, 1881, p. 726. 


410 CNGULATA 


rhinoceros is usually very fat; and its meat is then most excellent, 
being something like beef, but yet having a peculiar flavour of its 
own. The part in greatest favour among hunters is the hump, 
which, if cut off whole and roasted just as it is in the skin, ina 
hole dug in the ground, would, I think, be difficult to match either 
for juiciness or flavour.” : 

The molar dentition is of the type obtaining in R. wnicornis, so 
that in this respect 2. simus has the same relation to F. bicornis as 
is presented by R. unicornis to R. sondaicus. The ear-conch of the 
Square-mouthed Rhinoceros is very large, elongated, and pointed at 
its extremity, which bears only a slight tuft of hair; it is much ex- 
panded in the middle, and the lower portion has its edges united 
to form a short tube. The nostrils have a long slit-like aperture ; 
and the eye is situated behind the posterior horn. 

Extinct Species—Using the generic term Rhinocervs in its widest 
signification, a yery large number of fossil forms may be referred to 
it, the earliest of which date from the Upper Eocene (Oligocene) 
Phosphorites of Central France. Only a few of the more im- 
portant of these types can, however, be even mentioned in this 
place. 

In the Pliocene Siwaliks of India 2. sivalensis appears to have 
been the direct ancestor of R. sondaicus; while R. palwindicus was 
probably nearly related to R. wnicornis, although the upper molars 
had not developed a combing-plate. 

RR. schleirmacheri, of the Lower Pliocene of Europe, falls into 
the Ceratorhine group, although differing from R. sumatrensis by 
the union of the post-glenoid and post-tympanic processes of the 
squamosal beneath the auditory meatus. The Middle Miocene 
Zi. sansaniensis was a closely allied if not identical form. 

The Atelodine group was very widely spread in past epochs. 
Thus the huge &. platyrhinus of the Indian Pliocene, and the equally 
large L. wntiquitatis of the Pleistocene of Europe, were specialised 
forms with a dentition resembling that of R. simus, to which they 
were probably allied. An upper molar of R. antiquitatis—the so- 
called Tichorine, or Woolly Rhinoceros—is shown in the woodcut 
on p. 402. Of this species nearly whole carcases, with the thick 
woolly external covering, have been discovered associated with 
those of the Mammoth, preserved in the frozen soil of the north of 
Siberia. In common with some other extinct species it had a solid 
median wall of bone supporting the nasals, from which it is inferred 
that the horns were of a size and weight surpassing that of the 
modern species. In the Lower Pliocene of Attica R. pachygnathus 
appears to have been closely allied to R. dicornis. Several species, 
such as Lt. leptorhinus (Fig. 173), R. megarhinus, and R. cfruseis, 
occur in the European Pleistocene which do not present a marked 
relationship to any of the living forms. This group is also repre- 


RHINOCEROTIDE 411 


sented in the Pleistocene of Southern India by the small Lt. deccan- 
ensis and f. karnuliensis. 

In the Upper Miocene, or Lower Pliocene, of North America 
numerous Rhinoceroses with incisor teeth occur which have no 
nasal horn, although in those forms of which the limbs are known 
the fore feet resembled those of existing species in having only three 
digits. These species have been generically separated as Aphelops, 
but so closely do they resemble existing Rhinoceroses that at one 
time Professor Cope proposed to refer the hornless female of FR. 
sondaicus (described by Lesson as F. inermis) to the same genus. 
If these American types be included in hinoceros there seems no 
valid reason for separating the European Lower Pliocene and Mio- 
cene forms described as Aceratherium, at least some of which have 


Fia. 173.—Skull of Rhinoceros leptorhinus, from the Pleistocene of Essex. About } natural size. 


four digits in the manus. This group is represented in the Upper 
Eocene Phosphorites of France, and also by a very large species in 
the Pliocene of India. Lastly, R. minutus, of the Lower Miocene of 
France, and an allied North American species are distinguished by 
carrying a pair of very small horns placed transversely across the 
nasals, from which feature it has been proposed that they should 
be separated generically as Diceratherium. 

Extinct Generic Types.—The Tertiary deposits of different parts 
of the world have yielded remains of many extinct forms more or 
less closely related to the Rhinoceroses, and some of which should 
certainly be included in the same family; although others perhaps 
form the types of one or more distinct families. One of the most 
remarkable of these extinct types is the huge Elasmotheriuwm, from 
the Pleistocene of Siberia, in which the dentition was reduced to 
two premolars and three molars on either side of each jaw. The 


412 UNGULATA 


structure of the skeleton is essentially rhinocerotic, the skull having 
an ossified nasal septum, and a huge frontal prominence for the 
support of a very large horn. The teeth are extremely hypsodont, 
with the enamel plicated to a remarkable degree, and unlike those 
of Rhinoceros. The genus is evidently a very specialised one. 

The other genera we have to notice are more generalised types. 
Of these the North American Hyracodon, with the full typical 
number of teeth, and without nasal horn, appears to connect the 
Rhinoceroses with the Lophiodont Hyrachyus. The genera Amynodon 
and Metamynodon (Fig. 174), from the American Tertiaries, are 
forms allied to the Rhinoceroses, with the full number of incisors 
and canines, and the hinder lobe of the last upper molar not aborted. 
The lower canines are either upright, or less proclivous than in the 
Rhinoceroses ; in A/etamynodon the premolars are reduced to 3. 
Molar teeth from the Phosphorites of Central France, described 


i Ss ee WYN ‘ 
Fic. 174.—Right half of the palatal surface of the cranium of Metamynodon planifrons, from 
the Upper Miocene of North America. (After Scott and Osborn.) 


under the name of Cadurcotherium, are constructed on the general 
plan of those of the Rhinoceroses, although distinguished by their 
extreme narrowness; this type of tooth being very similar to that 
found in Homalodontotherium from Tertiary deposits in Patagonia. 
The latter has the full number of teeth, without any diastema in 
the series. Until we have some knowledge of the skeleton of these 
remarkable forms nothing definite can be said as to their serial 
position. 


Families LAMBDOTHERIID.E, CHALICOTHERIID, AND 
TITANOTHERIID&. 


These families contain a large number of more or less nearly 
related extinct types from Tertiary beds of both the Old and New 
Worlds, some of which present most remarkable deviations from 
the ordinary Ungulate structure. All are characterised by their 
brachydont molars, which depart widely from the normal lophodont 
type. The upper molars consist of four columns, of which the two 
external ones are expanded to form an outer wall; the posterior 
pair being connected in some cases by an oblique transverse ridge, 


LAMBDOTHERIHDA, ETC. . 413 


while there may be traces of an anterior ridge. The premolars 
are simpler. 

Lambdotheriide.—This family is confined to the Upper Eocene 
and Miocene of North America, where it is represented by Lambdo- 
therium, Paleosyops, and Limnosyops ; it presents the normal type 
of foot structure, and all the genera except the first have the full 
complement of teeth. There were four digits in the manus. The 
last lower molar has a third lobe. Limnosyops differs from Paleosyops 
in having two inner columns to the last upper molar. 

Chalicotheriide.—The genus Chalicotherium, which is found in the 
Tertiaries of Europe, Asia, and North America, differs so remark- 
ably in the structure of the feet from all other Ungulates that it has 
been proposed to regard it as the representative of a distinct order, 
Ancylopoda. The molars are, however, almost indistinguishable 
from those of the preceding and following families ; while the cervi- 
cal vertebree and portions of the limbs are of a Perissodactyle type. 
On the other hand, the femur has lost its third trochanter ; while 
the phalanges are strangely modified, the terminal ones forming 
long curved claws, while the others > (Fig, 175) have strong ging- 
lymoid distal articulations. 
These phalanges were, indeed, 
long regarded as referable to 
Edentates, being described in 
Europe as Macrotherium, and 
in the United States as Moro- 
therium and Moropus.  An- 
cylotherium, of the Grecian 
Pikermi beds, is founded upon y 
phalanges which indicate an 1G. 175.—Anterior and distal aspects of a 
allied genus. The Indian phalangeal bone of Chalicotherium sivalense. (From 
species of Chalicotherium is dis- ‘® Pwontolegia Indica.) 
tinguished by the loss of the incisors and the upper canine; while 
all the species want the first premolar. 

Titanothertide.—This exclusively North American family in- 
cludes gigantic forms closely allied to the Lambdotheriide, but with 
the last upper premolar as complex as the molars, and frequently 
with large bony protuberances in the nasal region. The best 
known genus, Titanotheriwum (Menodus, Brontotheriwm, Symborodon, 
Allops, etc.), may either have the full complement of teeth, or the 
incisors may be reduced to #2. The canines and incisors are small, 
and there is no diastema when the full dental series is developed. 
The skull is very like that of the Rhinoceroses; but has a trans- 
verse pair of large bony prominences on the nasal region, varying 
considerably in shape and size in the different species, which in the 
living animal were probably covered with horny sheaths. The third 

1 This name is the earliest, but is preoccupied. 


414 UNGULATA 


trochanter of the femur was.aborted. These huge animals— 
inferior in size only to the Elephant—appear to have been abundant 
in the United States during the Miocene period. 


Family MACRAUCHENIIDA. 


This extinct South American family is best known by the genus 
Afacrauchenia, as represented by Jf. patachonica and AL. boliviensis, 
which are apparently from Pleistocene formations. 'They are very 
singular and specialised forms, quite out of the line of descent of 
any of the existing Perissodactyles, and the steps by which they 
are connected with the rest of the group have not yet been 
discovered. Of the larger species, Jf patachonica, the skeleton is 
completely known. It had the full number of forty-four teeth, 
forming an almost uninterrupted series. The cervical vertebrae 
resemble those of the Camels in the position of the vertebrarterial 
canal, but the ends of the centra are flat, and not opisthoccelous as 
in the allied forms. In some of the limb characters it resembles 
the Hquide, but in the articulation of the fibula with the calcaneum 
it agrees with the Artiodactyles. The structure of the feet is, 
however, distinctly Perissodactylate, there being three toes on each. 
The teeth approximate to a Rhinocerotine structure ; and the incisors 
have an infolding of the enamel of their crowns, as in those of the 
Horses. The nares open on the top of the skull, and it is probable 
that the muzzle was produced into a short proboscis. Several 
other South American forms have been referred to this family, 
some of which have received distinct generic names, but further 
evidence is required before many of them can be accepted. Pos- 
sibly Homalodontotherium should be placed here. 


Family PROTEROTHERIID. 


Proterotherium.—Here may be noticed certain very remarkable 
Perissodactyles from the South American Tertiaries, for which the 
name Proterotherium has been proposed. The cheek-teeth are so 
like those of Anchitherium that they have been described under 
that name. The upper jaw has one pair of canine-like incisors and 
no canines, while the lower jaw carries two pairs of incisors. In 
the skull the orbits were completely closed, as in the Horses. The 
feet were tridactyle, like those of Hipparion, but the tarsus was 
constructed on an Artiodactyle type. 


SUBUNGULATA. 


By far the greater number of the Subungulata are extinct, and 
of many of those whose former existence has been revealed, chiefly 
by the labours of the American paleontologists, our knowledge is 
at present necessarily imperfect, though daily extending. It will 


HYVRACID.E 415 


only be possible here to give details of some of the more interest- 
ing ov best-known forms. 

The characters by which the skeleton of the feet of the Sub- 
ungulata are distinguished from those of the Ungulata Vera have 
been already mentioned on p. 275. In addition to these it may 
be observed that the feet frequently have five functional digits, 
and may be plantigrade; while the upper surface of the astragalus 
is generally flattened, instead of presenting the strongly-marked 
pulley-like ridges and groove so characteristic of the Ungulata 
Vera, 


Suborder HYRACOIDEA,. 
Family HYRACID.©. 


This division is constituted to receive a single family of mam- 
mals, the affinities of which have long constituted a puzzle to 


zoologists. They were first placed among the Rodents, to which 
animals their small size and general appearance and habits vive 
them much superficial resemblanee. Cuvier’s investigations into 
their anatomical structure, and especially their dental characters, 
led him to place them among the Ungulates, near the genus 
Rhinoceros, a position long accepted by many zoologists. Further 
knowledge of their organisation and mode of development. caused 
Milne-Edwards, Huxley, and others to disassociate them from this 
connection, and, failing to find any agreement with any other known 
forms, to place them in an order entirely apart. Paleontology has 
thrown no light upon the aftinities of this anomalous and isolated 
group, as no extinet animals possessing their distinetive characters 
have as yet been discovered. 


416 UNGULATA 


The dentition, according to the usual interpretation, consists 
only of incisors and molars, the formula in all known species being 
2 4,¢9,p4, m3. The upper incisors have persistent pulps, and 
are curved longitudinally, forming a semicircle as in Rodents. 
They are, however, not flattened from before backwards as in that 
order, but prismatic, with an antero-external, an antero-internal, 
and a posterior surface, the first two only being covered with 
enamel; their apices are consequently not chisel-shaped, but sharp 
pointed. They are preceded by functional, rooted milk-teeth. 
The outer lower incisors, which should perhaps be regarded rather 
as canines, have long tapering roots, but not of persistent 
growth. They are straight, procumbent, with awl-shaped, trilobed 
crowns. Behind the incisors is a considerable diastema. The 
molars and premolars are all contiguous, and formed almost exactly 


Fic. 177.—Skull and dentition of Dendrohyraz dorsalis. x 3. 


on the pattern of some of the Perissodactyle Ungulates. The hyoid 
arch is unlike that of any known mammal. The dorsal and lumbar 
vertebrae are very numerous, 28 to 30, of which 21 or 22 bear 
ribs. The tail is extremely short. There are no clavicles. In 
the fore foot the three middle toes are subequally developed, 
the fifth is present, but smaller, and the hallux is rudimentary, 
although, in one species at least, all its normal bones are present. 
The ungual phalanges of the four outer digits are small, somewhat 
conical, and flattened in form. The carpus has a distinct os 
centrale. There is a slight ridge on the femur in the place of a 
third trochanter. The fibula is complete, thickest at its upper 
end, where it generally ankyloses with the tibia. The articulation 
between the tibia and astragalus is more complex than in other 
mammals, the end of the malleolus entering into it. The hind 
foot is very like that of Rhinoceros, having three well-developed 
toes. There is no trace of a hallux, and the fifth metatarsal is 
represented only by a small nodule. The ungual phalanx of the 


AVRACIDEA 417 


inner (or second) digit is deeply cleft, and has a peculiar long 
curved claw, the others have short broad nails. The stomach is 
formed upon much the same principle as that of the Horse or 
Rhinoceros, but is more elongated transversely and divided by a 
constriction into two cavities—a large left cul de sac, lined by 
a very dense white epithelium, and a right pyloric cavity, with a 
very thick, soft, vascular lining. The intestinal canal (Fig. 178) 
is long, and has an 
arrangement per- 
fectly unique among 
mammals, indeed 
among vertebrated 
animals, for, in addi- 
tion to the ordinary 
short, but capacious 
andsacculated cecum 
(em) at the com- 
mencement of the 
colon, there is, lower 
down, an additional 
pair of large, conical, 
pointed, supplemen- 
tal ceca (c). The 
liver is much sub- 
divided, and there is 
no gall-bladder. The 
brain resembles that 
of the typical Un- 
_ gulates far more than 
the Rodents. The 
testes are perman- 
ently abdominal. Fic. 178.—Diagrammatic view of the alimentary canal of 
The ureters open into Hyrax capensis, the intestines being somewhat abbreviated. 


the fundus of the d, Duodenum ; i, ileum; cm, ceecum ; c, supplemental colic czca ; 
r, rectum. 


bladder, as in some 
Rodents. The female has six teats, of which four are inguinal 
and two axillary ; and the placenta is zonary, as in the Elephant 
and Carnivora. 
There are two distinct forms of Hyrax, differing both in 
structure and habits, which may be accorded generic rank. 
Hyraz—Molar teeth having the same pattern as those of 


1 Hermann, Tab. Affinit. Anim. p. 115 (1783). It has recently been pro- 
posed to substitute the earlier name Procavia in lieu of Hyrax, The anatomy of 
Hyrax was first described by Pallas (Spicilegia Zoologica). Besides minor 
memoirs, two detailed accounts of its structure have appeared—one by Brandt, 
in Mém. Acad. Nat. Scien. St. Pétersbowrg, 7*™* ser. vol. xiv. No. 2, 1869; and 

27 


418 UNGULATA 


Rhinoceros. Interval between upper incisors less than the width of 
the teeth. Lower incisors slightly notched at the cutting edge. 
Vertebre: C 7, D 22, L 8,8 6,0 6. Of this form the earliest 
known species, H. capensis (Fig. 176) is the type. There are several 
other species, as H. habessinicus and syriacus, from Eastern Africa 
and Syria. They inhabit mountainous and rocky regions, and live 
on the ground. 

Dendrohyraa.1—Molar teeth having the same pattern as Paleo- 
therium (except that the third lower molar has but two lobes). 
Interval between upper incisors exceeding the width of the teeth. 
Lower incisors with very distinctly trilobed crowns. Vertebre : 
C7, D 21,L7,85,C10. The members of this section frequent 
the trunks and large branches of trees, sleeping in holes. There 
are several species, not distinctly defined, from western and south 
Africa, as D. arboreus and D. dorsalis. The members of both groups 
appear to have a power like that possessed by the Lizards called 
Geckos of clinging to vertical surfaces of rocks and trees by the 
soles of their feet. 

It should be added that some writers separate three of the 
African species usually included in Hyrax (viz. H. bocagei, H. bakeri, 
and H. blainvillet) under the designation of Heterohyraz.? 


Suborder PROBOSCIDEA. 


This name has been appropriated to a well-marked group of 
animals, presenting some very anomalous characters, allied in many 
respects to the typical Ungulata, but belonging neither to the Artio- 
dactyle nor Perissodactyle type of that order. It has been thought 
that they possess some, though certainly not very close, affinities 
with the Rodentia, and also with the Sirenia. It is certain, 
however, that the two species of Elephant, which are the sole living 
representatives of the group, stand quite alone among existing 
mammals, differing widely from all others in many points of their 
structure. In some respects, as the skull, proboscis, and dentition, 
they are highly specialised ; but in others, as in the presence of two. 
anterior vene cave and in the structure of the limbs, they retain 
a low or generalised condition. A considerable series of extinct 
forms, extending back through the Pliocene and Miocene epochs, 
show the same type under different modifications, and in still more 


another by George, in Annales des Sciences Naturelles, 6%me sér. tom. i. 1874, in 
which references to all the previous literature will be found. The mechanism 
by which the sole of the foot is enabled to adhere to smooth surfaces is fully 
described by G. E. Dobson, Proc. Zool. Soc. 1876, p. 526. 

’ Gray, Ann, Mag, Nat, Hist. ser. 4, vol. i, p. 48 (1868). 

° See a paper by J. V. Barboza du Bocage, in the Jorn. Set. Phys. Nat. Lisboa 
(2), vol. i. p. 186 (1889), where a list of all the known species will be found. 


PROBOSCIDEA 419 


generalised outlines; and certain forms from the Eocene of North 
America, if their affinities are rightly interpreted, appear to link 
the true Proboscidea to some unknown primitive type of Ungulata. 

The following are the principal characters common to existing, 
and, by inference, to the extinct, Proboscidea. The nose extended 
into a long, muscular, very flexible and prehensile proboscis, at the 
end of which the nostrils are situated, and from which the name 
given to the group is derived. The teeth consisting of ever-growing 
incisors of very great size, but never exceeding one pair in each 
jaw, and often present in one jaw only; no canines; large and 
transversely ridged molars. No clavicles. Limbs strong, the 
upper segment, especially in the hind limb, the longer. Radius 
and ulna distinct, the latter articulating extensively with the carpus. 
Fibula and tibia distinct. Astragalus very flat on both surfaces. 
Manus and pes short, broad, and massive, each with five toes, 
though the outer pair may be more or less rudimentary, all encased 
in a common integument, though with distinct, broad, short hoofs. 
Third digit the largest. Two anterior venz cave entering the 
right auricle. Stomach simple. A capacious cecum. Testes per- 
manently abdominal. Uterus bicornuate. Placenta nondeciduate 
and zonary. Mamme two, pectoral. 

With regard to the teeth, the incisors,! which project largely 
out of the mouth, and are commonly called “tusks,” are of an 
elongated conical form, and generally curved. They are composed 
mainly of solid dentine, the fine elastic quality and large mass of 
which renders it invaluable as “ivory” for commerce and the arts. 
A peculiarity of the dentine of most Proboscidea is that it shows, in 
transverse fractures or sections, strie proceeding in the arc of a 
circle from the centre to the circumference in opposite directions, 
and forming by their decussations curvilinear lozenges, as in the 
“ engine-turning ” of the case of a watch. The enamel-covering in 
existing species is confined to the extreme apex, and very soon 
wears off, but in some extinct species it forms persistent longitudinal 
bands of limited breadth. The tusks have small milk-predecessors, 
shed at an early age. 

The molar teeth present a remarkable series of modifications, 
from the comparatively simple form in Dinotherium, with two or 
three strongly pronounced transverse ridges and a normal mode of 
succession, to the extremely complex structure and anomalous mode 


1 These teeth are by some writers classed as canines, as their roots are im- 
planted in the maxille ; but, as in Rodents, they are originally developed in the 
gum covering the premaxille, in which bones their primitive alveoli are sunk. 
As growth proceeds, however, firm support for such massive and weighty bodies 
can only be obtained by their roots gradually sinking through the premaxille 
into the great and specially modified alveolar processes of the maxille, but this 
does not vitiate their homology with the incisors of other mammals. 


420 UNGULATA 


of replacement found in the true Elephants. The intermediate 
conditions occur in the various species of J/ustoden. In this genus 
the enamel-covered transverse ridges of each tooth are generally 
more numerous than in Dinotherium, and often complicated by 
notches dividing their edge or by accessory columns attached to 
them, but in the unworn tooth they stand out freely on the surface 
of the crown, with deep valleys between (Fig. 179, I). In the 
Elephants the ridges are still further increased in number, and con- 
sequently narrower from before backwards, and are greatly extended 


renga 


NNN ng 


RU UER 
OTA ae 
beeen, 


vi sym 
ees } 


Ai 


Fia. 179.—Longitudinal sections of the crown of a molar tooth of various Proboscideans 
showing stages in the gradual modification from the simple to the complex form. I, Mastodon 
americanus ; II, Elephas insignis; III, Elephas africanus; IV, Elephas primigenius. The dentine 
is indicated by transverse lines, the cement by a dotted surface, and the enamel is black. 


in vertical height, so that, in order to give solidity to what would 
otherwise be a laminated or pectinated tooth, it becomes necessary 
to envelop and unite the whole in a large mass of cement, which 
completely fills up the valleys, and gives a general smooth appear- 
ance to the organ when unworn ; but as the wear consequent upon 
the masticating process proceeds, the alternate layers of tissue 
of different hardness—cement, dentine, and enamel—which are 
disclosed upon the surface form a fine and very efficient trituratine 
instrument. The modification of the tooth of a Mastodon into that 
of an Elephant is therefore precisely the same in principle as that 
of the molar of a Palzotherium into that of a Horse, or of the 


PROBOSCIDEA 421 


corresponding tooth of one of the primitive Artiodactyles into that 
of an Ox. The intermediate stages, moreover, even in the present 
state of our knowledge, are so numerous that it is not possible to 
draw a definite line between the two types of tooth structure (see 
Fig. 179, I, II, Il, IV). 

As regards the mode of succession, that of modern Elephants is, 
as before mentioned, very peculiar. During the complete lifetime 
of the animal there are but six molar teeth on cither side of each 
jaw, with occasionally a rudimentary one in front, completing the 
typical number of seven. The last three represent the true molars 
of ordinary mammals; those in front appear to be milk-molars, 
which are never replaced by permanent successors, but the whole series 
gradually moves forwards in the jaw, and the teeth become worn 
away and their remnants cast out in front, while development of 

_ others proceeds behind. The individual teeth are so large, and the 
processes of growth and destruction by wear take place so slowly, 
that not more than one, or portions of two, teeth are ever in place 
and in use on either side of each jaw at one time, and the whole series 
of changes coincides with the usual duration of the animal’s life. 
On the other hand, the Dinotherium, the opposite extreme of the 
Proboscidean series, has the whole of the molar teeth in place and 
use at one time, and the milk-molars are vertically displaced by 
premolars in the ordinary fashion. Among Mastodons transitional 
forms occur in the mode of succession as well as in structure, many 
species showing a vertical displacement of one or more of the milk- 
molars, and the same has been observed in one extinct species of 
Elephant (£4. plunifrons) as regards the posterior of these teeth. 

All known Proboscideans are animals of comparatively large 
dimensions, and some are the most colossal of land mammals. The 
head is of great proportionate size ; and, as the brain case increases 
but little in bulk during growth, while the exterior wall of the 
skull is required to be of great superficial extent to support the 
trunk and the huge and ponderous tusks, and to afford space for 
the attachment of muscles of sufficient size and strength to wield 
the skull thus heavily weighted, an extraordinary development of 
air-cells takes place in the cancellous tissue of nearly all the bones 
of the cranium (Fie. 180). These cells are not only formed in the 
walls of the cranium proper, but are also largely developed in the 
nasal bones and upper part of the premaxilla and maxilliv, the bones 
forming the palate and the basicranial axis, and even extend into 
the interior of the ossified mesethmoid and vomer. Where two 
originally distinct bones come into contact, the cells pass freely 
from one to the other, and almost all the sutures become obliterated 
in old animals. The intercellular lamelle in the great mass which 
surrounds the brain cavity superiorly and laterally mostly radiate 
from the ‘inner to the outer table, but in the other bones their 


422 UNGULATA 


direction is more irregular. Like the similar but less developed 
air-cells in the skulls of many other mammals, they all communicate 
with the nasal passages, and they are entirely secondary to the 
original growth of the bones, their development having scarcely 
commenced in the new-born animal, and they gradually enlarge as 
the growth of the creature proceeds towards maturity. The nasal 
bones are very short, and the anterior narial aperture is situated 
high in the face. The zygomatic arch is slender and straight, the 
jugal bone being small, and forming only the middle part of the 
arch, the anterior part of which (unlike that of typical Ungulates) is 


Fic. 180.—A vertical section of the skull of the African Elephant (Elephas africanus) taken 
to the left of the middle line, and including the vomer (Vo) and the mesethmoid (ME). 
an, Anterior, and pn, posterior narial aperture. yz natural size. (From Flower’s Osteology of 


the Mammalia.) 

formed only by the maxilla. The maxillo-turbinals are but rudi- 
mentary, the elongated proboscis supplying their place functionally 
in warming and clearing from dust the inspired air. 

The neck is very short. The limbs are long and stout, and 
remarkable for the great length of the upper segment (especially 
the femur) as compared with the distal segment, the manus, and 
pes. Itis owing to this and the vertical position of the femur that 
the knee-joint in the hind leg is placed much lower, and is more 
conspicuous externally than in most quadrupedal mammals; and 
this having been erroneously compared with the hock-joint or ankle 
of typical Ungulates, the popular fallacy that the joints of the 
Elephant’s leg bend in a contrary direction to that of other mam- 
mals has arisen. There is no round ligament in the hip-joint, or 
third trochanter to the femur. The radius and ulna are distinct, 
though fixed in a crossed or prone position. The fibula also is 


PROBOSCIDEA 423 


quite distinct from the tibia. The feet are short and broad, the 
carpal and tarsal bones being very square, with flattened surfaces 
for articulation ; the astragalus especially differs from that of typical 
Ungulates in its flatness, in the absence of a distinct pulley-like 
articular surface at either extremity, and in having no articular 
facet for the cuboid. The fibula articulates with the caleaneum, as 
in Artiodactyles. Of the five toes present on each extremity (see 
Fig. 98), the middle one is somewhat the largest, and the lateral 
ones smallest, and generally wanting (especially in the hind foot) 
the complete number of phalanges. The ungual phalanges are all 
small, irregular in form, and late in ossification. The whole are 
encased in a common integument, with a flat, subcircular, truncated 
sole, the only external indication of the toes being the broad oval 
nails or hoofs arranged in a semicircle around the front edge of the 
sole. The hind foot is smaller and narrower than the front. The 
liver is small and simple, and there is no gall-bladder. In form 
the brain resembles that of the Rodents and other lower orders of 
mammals, the cerebellum being entirely behind and uncovered by the 
cerebrum, but the hemispheres of the latter are richly convoluted. 

The Proboscidea are exclusively vegetable feeders, living chiefly 
on leaves and young branches of forest trees and various kinds of 
herbage, which they gather and convey to their mouth by the very 
mobile proboscis, an organ which combines in a marvellous manner 
strength with dexterity of application, and is a necessary compensa- 
tion for the shortness and inflexibility of the neck, as by it many 
of the functions of the lips of other animals are performed. By its 
means the Elephant is enabled to drink without bending the head 
or limbs; the end of the trunk being dipped into the stream or 
pool, a forcible inspiration fills the two capacious air-passages in 
its interior with water, which, on the tip of the trunk being turned 
upwards and inserted into the mouth, is ejected by a blowing action, 
and swallowed ; or if the animal wishes to refresh and cool its skin, 
it can throw the water in a copious stream over any part of its 
surface. Elephants can also throw dust and sand over their bodies 
by the same means and for the same purpose, and wild animals 
have been frequently observed fanning themselves with leafy boughs 
held in the trunk. The species are at present limited in their 
geographical distribution to the Ethiopian and Oriental regions, but 
they formerly had a far more extensive range. 


Family ELEPHANTID.£. 


Cheek-teeth succeeding one another in an are of a cirele, and 
portions of only two, or at most three, of the hinder teeth in use 
at any one time. Premolars frequently lost, and in any case of no 
functional importance. 


424 UNGULATA 


Elephas.\—Dentition: i2, ¢ 8, dm 3, m $=26. The incisors 
variable, but usually of very large size, especially in the male sex, 
directed somewhat outwards, and curved upwards, without enamel 
except on the apex before it is worn. The molars composed of 
numerous flattened enamel-covered plates or ridges of dentine, 
projecting from a common many-rooted base, surrounded and united 
together by cement, and extending straight across the crown, with- 
out (in most forms) any median division into inner and outer 
columns. The number of plates increases from the anterior to the 
posterior molar in regular succession, varying in the different species, 
but the third and fourth (or the last milk-molar and the first true 
molar), and these only, have the same number of ridges, which 
always exceeds five. Premolars nearly always wanting. Skull 
of adult very high and globular. Mandible ending in front in a 
short, deflected, and spout-like symphysis. Vertebre: C 7, D 19- 
21, L 3-4, 8 4, C 26-33. 

The existing species of the genus differ so much that they have 
been referred by some writers to distinct genera ; fossil forms show, 
however, such a transition from the one to the other that it is 
scarcely possible to regard them even as the representatives of 
distinct groups. 

In the well-known Indian or Asiatic Elephant (£. indicus) the 
average number of plates of the six successive molar teeth is 
expressed by the “ridge-formula,” 4, 8, 12, 12, 16, 24. The 
plates are compressed from before backwards, the anterior and 
posterior surfaces (as seen in the worn grinding face of the tooth, 
Fig. 181) being 
nearly parallel. 
Ears of moder- 
ate size. Upper 
margin of the 
end of the pro- 
boscis devel- 
oped into a 
distinct finger- 


Fic. 181.—Grinding surface of a half-worn lower molar of the Indian like process, 
Elephant (Elephas indicus). d, Dentine; e, enamel; c, cement. (From much lon ger 


ne than the lower 
margin. Five nails on the fore feet, and four (occasionally five) on 
the hind feet. 

This species inhabits in a wild state the forest lands of India, 
Burma, the Malay Peninsula, Cochin China, Ceylon, and Sumatra. 
The elephants from the last-named islands, presenting some variations 
from those of the mainland, have been separated under the name of 
F. sumatranus, but the distinction has not been satisfactorily estab- 

1 Linn. Syst. Vat. 12th ed. vol. i. p. 48 (1766). 


LIFEPHANTIP 2 A3% 


Nshed. The appearance ci the Asiatic Elephant is familiar <o all. 
Though rarely breeding in captivity. it has been domesticated from 
the most remete antiquity, and is still extensively used in the East 
as a beast of burden. In the wild state it is gregarious, associating 
in herds of ten. twenty, or more individuals, and though it may, 
under certain circumstances. become dangerous. it is generally 
inoffensive and even timid. fond ci shade and solitude and the 
neizhbourhood ci water. The height of the male at the shoulder 
when full grown is usually from $ to 10 feet, but occasionally as 
much as 1]. The female is somewhat smaller. 

In the African Elephant (£. 27rieanus) the molars (Fig. 152: are 
ef ecarse constrnetion, with fewer and larger plates and thicker 
enamel. Ridge-formula: 3. 5. 7, 7. >. 10. _ The plates not 
flattened, but thicker in the middle than at the edges. so that their 
worn grinding suriaces are lozenge-shaped. Ears very large. The 


pas The lef Ge af oo feuress che Son 
(From Owen.) 


h. and the lower sil: the onter bonier. 


upper and lower marzgirs ¢i the end of the trunk forming iv: 
nearly equal prehensile ics. But three bests on the hind foot. 
This species now inhabits the wooded districts of the whole oi 
Afiea south <i the except where i: has been driven away 
by human serlements. Fossil remains of Pleiscocene age. undis- 
tinguishable specifically, have been Zound in Alzeria, 
Sealy. Ir was trained for war and show by the ancient Cartha- 
gimians and Romans, and recent experience of the species in ay 
in Haglan! shows that it is as mnvelizent as its 
net more so. while surpassing it in evurage. av 
Nevertheless. in modern times, no people in Africa have been 
sufticiently civiised or enterp: Tsing To eare to train it tor domestic 
yourmeses. It is buntea chiefly for the sake of the ivory ¢f its 
immense tusss. of which it vVields the principal source of supply to 
the European market, and the cexre to obtain which is rapidly 
leading to the extermination ci the species. Im size the male 
/ rpasses that of Asis. but the female js 
sumierence of the fore Ioct is halt the 


oar 


426 UNGULATA 


height at the shoulder, a circumstance which enables the hunters to 
judge from the footprints the exact size of the animals of which 
they are in pursuit. The African Elephant also differs from its 
Indian congener in having tusks in both sexes, whereas in the latter 
the male only is so armed. Moreover, the eye is relatively larger, 
the forehead more convex, and the colour somewhat darker. 
Whereas the Indian Elephant frequents the depths of forests and 
seldom leaves their shade during the daytime, the following 
account by Sir Samuel Baker indicates different habits in the 
African species. This traveller observes: “In Africa, the country 


Fic. 183.—African Elephant (Elephas ayricanus). From a young specimen in the 
London Zoological Gardens. 


being generally more open than in Ceylon, the Elephant remains 
throughout the day either beneath a solitary tree or exposed to 
the sun in the vast prairies, where the thick grass attains a height 
of from nine to twelve feet. The general food of the African 
Elephant consists of the foliage of trees, especially mimosas. Many 
of the mimosas are flat-headed, about thirty feet high, and the 
richer portion of the foliage confined to the crown. Thus the 
Elephant, not being able to reach to so great a height, must over- 
turn the tree to obtain the coveted food. The destruction caused 
by a herd of Elephants in a mimosa forest is extraordinary, and I 
have seen trees uprooted of so large a size that I am convinced no 
single elephant could have overturned them, I have measured 
trees four feet six inches in circumference and about thirty feet 
high uprooted by elephants. The natives have assured me that 


ELEPHANTIDE 427 


the elephants mutually assist each other, and that several engage 
together in the work of overturning a large tree.” 

Extinct Species of Elephant.—Abundant remains of Elephants are 
found embedded in alluvial gravels, or secreted in the recesses of 
caves, into which they have been washed by streams and floods, or 
dragged as food by Hyznas and other carnivorous inhabitants of 
these subterranean dens. Such remains belonging to the Pleistocene 
and Pliocene periods have been found in many parts of Europe, 
including the British Isles, in North Africa, throughout the North 
American continent from Alaska to Mexico, and extensively dis- 
tributed in Asia, where the deposits of the sub-Himalayan Siwalik 


Fic. 184.—Restored skeleton of the Mammoth (Elephas primigenius). From Tilesius in 
Mém. Acad. Imp. Sc. St. Pétersbourg, vol. v. (1815). s, Scapula; h, humerus; 7, radius; w, ulna; 
c, carpus; rs, ischium ; f, femur; t, tibia; ji, fibula; ta, tarsus. 


Hills, and equivalent deposits in the Punjab, Perim Island,! and 
Burma, belonging to the earliest Pliocene, are rich in the remains 
of Elephants of varied form. These species are chiefly known and 
characterised at present by the skulls and teeth ; some of the latter 
resemble the existing Indian and some the African type, but the 
majority are between the two, and make the distinction between 
the two existing species as of generic importance quite impractic- 
able. Others again approach so closely in the breadth and coarse- 
ness of the ridges and paucity of cement to Mastodon as to have 
been placed by some zoologists in that genus. These form the 
subgenus called Stegodon by Falconer, and may be regarded as a 
distinct group of the genus. 


1 In the Gulf of Cambay,—not the island of the same name in the Red Sea. 


428 UNGULATA 


Among the best known extinct Elephants is £. primigenius, the 
Mammoth,! very closely resembling the existing Indian species, and 
one of the most recently extinct and extensively distributed of all 
the fossil forms. Probably no animal which has not survived to 
the historic period has left such abundant and well-preserved evi- 
dence of its former existence. The discovery of immense numbers, 
not only, as in the case of most extinct creatures, in the form of 
fragmentary bones and teeth, but often as more or less nearly 
entire carcases, or “mummies,” as they may be called, with the 
flesh, skin, and hair im situ, in the frozen soil of the tundras of 
Northern Siberia, has for along time given great interest to the 
species, and been the cause of many legendary stories among the 
natives of the lands in which they occur. Among these one of the 
most prevailing is that the Mammoth was, or still is, an animal which 
passes its life habitually in burrows below the surface of the ground, 
and immediately dies if by any chance it comes into the upper air. 

Of the whole group the Mammoth is in many respects, as in the 
size and form of the tusks, and especially the characters of the 
molar teeth, the farthest removed from the primitive Mastodon-like 
type, while its nearest surviving relative, £. indicus, has retained 
the slightly more generalised characters of the Mammoth’s con- 
temporaries of more southern climes, Z. columbi of America, and 
L. armeniacus of the Old World, if, indeed, it can be specifically 
distinguished from them. 

The tusks or upper incisor teeth were doubtless present in both 
sexes, but probably of smaller size in the female. In the adult 
males they often attained the length of from 9 to 10 feet measured 
along the outer curve. Upon leaving the head they were directed 
at first downwards and outwards, then upwards and finally inwards 
at the tips, and generally with a tendency to a spiral form not seen 
in other species of Elephant. Different specimens, however, present 
great variations in curve, from nearly straight to an almost com- 
plete circle. 

It is chiefly by the characters of the molar teeth that the 
various extinct modifications of the Elephant type are distinguished. 
Those of the Mammoth (Fig. 185) differ from the corresponding 
organs of allied species in the great breadth of the crown as 
compared with the length, the narrowness and close approximation of 
the ridges, the thinness of the enamel and its straightness, parallel- 
ism, and absence of “ crimping,” as scen on the worn surface, or in a 
horizontal section of the tooth. Dr. Falconer gave the prevailing 


? The word Mammoth was introduced into the languages of Western Europe 
about two centuries ago from the Russian, and is thought by Pallas and Norden- 
skiéld to be of Tartar origin, but others, as Witzen, Strahlenburg, and Howorth, 
have endeavoured to prove that it is a corruption of the Arabic word Behemoth, 
or great beast. 


ELEPHANTIDE 429 


“ridge-formula” as 4, *, 12, 12, 16, 24. Dr. Leith Adams, work- 
ing from more abundant materials, has shown, however, that the 
number of ridges of each 
tooth, especially those at 
the posterior end of the 
series, is subject to very 
great individual variation, ~ | 
ranging in each tooth of 

the series within the fol- 
lowing limits: 3 to 4, 6 
to % 9 to 12, 9 to 15, 
14 to 16, 18 to 27, ex- on ee i 

cluding the small plates 71+ Grating sures of soe meas of the 
called talons at each end ,, enamel. (From Owen.) 

of the tooth. Besides these 

variations in the number of ridges or plates of which each tooth is 
composed, the thickness of the enamel varies so much as to have 
given rise to a distinction between a “thick-plated ” and a “thin- 
plated” variety—the latter being most prevalent among the speci- 
mens from the Arctic regions, and most distinctively characteristic 
of the species. From the specimens with thick enamel plates 
the transition to the other species or varieties mentioned above, 
including £. indicus, is almost imperceptible. 

The hones of the skeleton generally more resemble those of the 
Indian Elephant than of any other known species, but the skull 
differs in the narrower summit, narrower temporal fozs#, and more 
prolonged incisive sheaths required to support the roots of the 
enormous tusks. Among the external characters by which the 
Mammoth was distinguished from either of the existing species of 
Elephant was the dense clothing, not only of long coarse outer hair, 
but also of close woolly under hair, of a reddish-brown colour, 
evidently in adaptation to the colder climate which it inhabited. 
This character, for a knowledge of which we are indebted to the 
well-preserved remains found in Northern Siberia, is also represented 
in the rude but graphic drawings of prehistoric age found in caverns 
in the south of France.t In size different individuals varied con- 
siderably, but the average height does not appear to have exceeded 
that of either of the existing species of Elephant. 

The geographical range of the Mammoth was very extensive. 
There is scarcely a county in England in which some of its remains 
have not been found either in alluvial deposits of gravel or in 
caverns, and numbers of its teeth are from time to time dredged 


1 The best known of these is the etching upon a portion of tusk found in the 
cave of La Madelaine in the Dordogne, figured in Lartet and Christy’s Reliquie 
Aquitanice, and in many other works bearing on the subject of the antiquity of 
man. 


430 UNGULATA 


up from the bottom of the sea by the fishermen who ply their 
trade in the German Ocean, having been washed out of the water- 
worn cliffs of the eastern counties of England. In Scotland and 
Ireland its remains are less abundant, but they have been found in 
vast numbers at various localities throughout the greater part of 
Central Europe (as far south as Santander in Spain and Rome), 
Northern Asia, and the northern part of the American continent, 
though the exact distribution of the Mammoth in the New World 
is still a question of debate. It has not hitherto been met with in 
any part of Scandinavia or Finland. 

In point of time, the Mammoth belongs exclusively to the 
Pleistocene epoch, and it was undoubtedly contemporaneous with 
man in France, and probably elsewhere. There is evidence to show 
that it existed in Britain before, during, and after the glacial period. 

As before indicated, it is in the northern part of Siberia that 
its remains have been found in the greatest abundance, and in 
quite exceptional conditions of preservation. For a very long 
period there has been from that region a regular export of 
Mammoth ivory in a state fit for commercial purposes, both east- 
ward to China and westward to Europe. In the middle of the tenth 
century an active trade was carried on at Khiva in fossil ivory, 
which was fashioned into combs, vases, and other objects, as related 
by Abu ’l Kasim, an Arab writer of that period. Middendorff 
reckoned that the number of tusks which have yearly come into 
the market during the last two centuries has been at least a hundred 
pairs, and Nordenskiéld, from personal observation, considers this 
calculation as probably rather too low than too high. They are 
found at all suitable places along the whole line of the shore 
between the mouth of the Obi and Behring Straits, and the farther 
north the more numerous do they become, the islands of New 
Siberia being now one of the most favourite collecting localities. 
The soil of Bear Island and of Liachoff Islands is said to consist only 
of sand and ice with such quantities of Mammoth bones as almost 
to compose its chief substance. The remains are not only found 
around the mouths of the great rivers, as would be the case if the 
carcases had been washed down from more southern localities in 
the interior of the continent, but are imbedded in the frozen soil 
in such circumstances as to indicate that the animals had lived not 
far from the localities in which they are now found, and they are 
exposed either by the melting of the ice in unusually warm 
summers or by the washing away of the sea cliffs or river banks 
by storms or floods. In this way the bodies of more or less nearly 
perfect animals, often standing in the erect position, with the soft 
parts and hairy covering entire, have been brought to light. 

References to the principal recorded discoveries of this kind, 
and to the numerous speculations to which they have given rise, 


ELEPHANTIDA 431 


both among ignorant peasants and learned academicians, will be 
found in Nordenskiéld’s /’vyage of the Vega (English translation, 
vol. i. 1881, p. 398 sy.) and a series of papers in the Geologicud 
Magazine for 1880 and 1881, by H. H. Howorth, as well as ina 
separate work on the Mammoth by the same writer. For the 
geographical distribution and anatomical characters, see Falconer’s 
Patwontological Memoirs, vol, ii. 1868; Boyd Dawkins, “Zlephas 
primigenius, its Range in Space and Time,” Quurt. Journ. Geol. Soc. 
xxxv. p. 138 (1879); and Leith Adams, “Monograph of British 
Fossil Elephants,” part ii, Paleontegraphical Society (1879). 

E. antiquus, of the Enropean Pleistocene, has a lower ridge- 
formula than in the Mammoth, the molars being narrower, and 
approximating to those of the African Elephant in structure. 
Small allied forms occur in the rock-fissures and caverns of Malta, 
and have been described as £. mnaidriensis and E. melitensis ; some 
of the individuals of the latter not exceeding 3 feet in height. The 
European #. meridionalis is a southern form of somewhat carlier 
age, very common in the Upper Pliocene of Italy and France, and 
also in the so-called Forest-bed of the Norfolk coast. It attained 
very largo dimensions, its height being estimated at upwards of 
15 feet. The ridge-formula is lower than in £. antiguas, the 
molars are broad, with the worn enamel-dises generally expanded 
in the middle, and the enamel itself is crenulated. 

Elephant remains are very abundant in the Pleistocene and 
Pliocene deposits of India, those from the latter beds being the 
oldest. representatives of the genus. Of these the Pleistocene 
fH. namadicus appears closely allied to £. antigens, from which it is 
distinguished by a bold ridge across the forehead. Among the Plio- 
cone forms Z. hysudricus may be an ancestral type allied to the Indian 
Elephant ; while £. planifrons is closely velated to £. meridionalis, 
although retaining the ancestral feature of developing premolars. 

The Stegodont group is peculiar to the castern parts of the 
Old World, and, as already observed, connects the true Elephants 
intimately with the Mastodons. The molars (Fig. 179, I) are 
characterised by tho lowness of the ridges, while the intervening 
valleys may have but little cement, and there may be a more 
ov less distinct longitudinal groove in the crown dividing each 
ridge into an inner and an outer moiety. In species like LZ. insignis 
the ridge-formula is neatly the same as in A. meridionalis, but in 
BK. cliftt, some of the molars carry only six ridges, and premolars 
were present, so that we thus have such a complete transition to 
tho next genus that it is very difficult to know where to draw the 
line between the two. 

AMastodon.A—Deutition : 4 ew ¢ 8, dn 3, m3. Upper incisors 


Y Cuvier, fn. du Muséan, vol. villi, p. 270 (1806). 


432 UNGULATA 


very large, as in Elephas, sometimes with longitudinal bands of 
enamel, more or less spirally disposed. Lower incisors variable ; 
when present comparatively small and straight, sometimes per- 
sistent, sometimes early deciduous, and in some species never 
present. Grinding surface of molars with transverse ridges, the 
summits of which are divided more or less into conical or mam- 
millary cusps, and often with secondary or additional cusps between 
and clustering against the principal ridges; enamel thick; cement 
very scanty, never filling up the interspaces between the ridges. 
The third, fourth, and fifth cheek-teeth (ie. the last milk-molar, 
and the first and second molars) having the same number of ridges,! 
which never exceeds five. 

In the upper jaw the incisors, though of large size, were 
apparently never so much curved as in some species of Elephant, 
and they often have longitudinal bands of enamel, more or less 
spirally disposed upon their surface, which are not met with in 
Elephants. Lower incisors were present throughout life in some 
species, which have the symphysis of the lower jaw greatly elon- 
gated to support them (as in If. angustidens, M. pentelici, and M. 
longirostris). Inthe common North American species (Af. americanus) © 
the mandibular symphysis is short, but it may have a small incisor 
on one side. In other species no inferior tusks have been found, 
at all events in adult life (see figure of MZ. arvernensis). 

The molar teeth increase in size from before backwards, but as 
many as three of these teeth may be in place in each jaw at one 
time. There is in many species a true vertical succession, affecting 
either the third, or the third and second, or (in M. productus) the 
first, second, and third of the six molariform teeth. These three 
are therefore reckoned as milk-molars, and their successors as pre- 
molars, while the last three, which are never changed, correspond 
to the true molars of those animals in which the typical dentition 
is fully developed. The study of the mode of succession of the 
teeth in the different species of Mastodons is particularly interest- 
ing, as it exhibits so many stages of the process by which the very 
anomalous dentition of the modern Elephants may have been 
derived by gradual modification from the typical heterodont and 
diphyodont dentition of the ordinary mammal. It also shows that 
the anterior molars of Elephants do not correspond to the pre- 
molars of other Ungulates, but to the milk-molars, the early loss of 
which in consequence of the peculiar process of horizontal forward- 


1 This, and the larger number of ridges in the latter, are the only absolute 
distinctions which Falconer could find between Mastodon and Elephas (Paleont. 
Memoirs, ii. p. 9), and it is clear that they are somewhat arbitrary. The line 
between the two genera is drawn at this point more as a matter of convenience 
for descriptive purposes than as indicating any great natural break in the 
sequence of modifications of the same type. 


ELEPHANTIDA 433 


moving succession does not require, or allow time for, their replace- 
ment by premolars. It must be noted, however, that, in the 
Mastodon in some respects the least specialised in tooth-structure, 
the A. americanus of North America, no vertical succession of the 


A 


Sere 


Fic. 186.—Restoration of the skeleton of Mastodon arvernensis, from the Pliocene of Europe. 
(After Sismonda.) 


molars has yet been observed, although vast numbers of specimens 
have been examined. 

The Mastodons have fewer ridges on their molar teeth than 
the Elephants; the ridges are also less elevated, wider apart, have 
a thicker enamel-cover- 
ing, and scarcely any 
cement filling up the 
space between them. 
Sometimes (as in Jf. 
americanus) the ridges 
are simple transverse 
wedge-shaped — eleva- 
tions, with straight or 
concave edges. In 


other species the sum- Fic. 187.—Oblique side and crown view of the last upper 
mits of the ridges are molar of Mastodon arvernensis. (From Owen.) 


more or less subdivided into conical cusps, and may have accessory 
cusps clustering around them (as in Jf. arvernensis, see Fig. 187). 
When the apices of these are worn by mastication, their surfaces 
present circles of dentine, surrounded by a border of enamel, and 
as the attrition proceeds different patterns are produced by the 
union of the bases of the cusps, a trilobed or trefoil form being 
characteristic of some species (Fig. 188). 
28 


434 eee cem: E Em 


As already mentioned, certain of the molariform teeth of the 
middle of the series in 
Mastodons have the 
same number of princi- 
pal ridges, those in 
front of them having 
fewer and those behind 
a greater number. 
These teeth were dis- 
tinguished as “ inter- 
mediate” molars by 
Dr. Faleoner, and are 


Fic. 188.—Grinding surface of the partially worn last three in number, hame- 
left lower milk-molar of Mustofon angustidens, from the Jy the last milk-molar 


Upper Miocene of India. The lower side of the figure is ~ Sane eee ene 
the outer border of the tooth. and the first and se cond 


true molars (or the 
third, fourth, and fifth of the whole series). The number of ridges 
on these intermediate molars is nearly always three or four, and the 
tooth in front has usually one fewer and that behind one more, so 
that the ridge-formula of most Mastodons can be reduced either to 
1, 2, 3, 3, 3, 4, or 2, 3, 4, 4, 4, 5. The former characterises the 
section called Tr ilophodon (of which an intermediate molar is shown 
in Fig. 188), and the latter that called Zetralophodon by Dr. Faleoner. 
These divisions are very useful, as under one or the other all the 
present known species of Mastodon can be ranged, but observations 
upon a larger number of individuals have shown that the number 
of ridges upon the teeth is not quite so constant as implied hy the 
formule given above. Their exact enumeration is even difficult in 
many cases, as “talons” or small accessory ridges at the hinder end 
of the teeth occur in various stages of development, until they take 
on the character of true ridges. Transitional conditions have also 
been shown, at least in some of the teeth, between the trilophodont 
and the tetralophodont forms, and again between the latter and 
what has been called a “pentalophodont” type, which leads on 
towards the condition of dental structure characteristic of the true 
Elephants. 

The range of the genus Jfustedon in time was from the middle 
of the Miocene period to the end of the Pliocene in the Old World, 
when it became extinct ; but in America several species—especially 
the one best known, owing to the abundance of its remains, which 
has been variously called i. americanus, M. ohioticus, and MW. giganteus 
—survived to a late Pleistocene period. 

The range in space will be best indieated by the following list 
of some of the better known species. (1) Trilophodont series 
AL. angustidens,! borsoni, pentelici, turicensis, from Europe ; 1. falconert 

1 Also found beyond the extreme north-western froutier of India. 


DINOTHERUD& 435 


and pandionis, from India ; M. americanus, obscurus, and productus, 
North America; and M. cordillerwm and humboldti, South America. 
(2) Tetralophodont series—M. arvernensis, M. longirostris, from 
Europe ; M. latidens, sivalensis, and perimensis, from India ; AZ. mari- 
jficus, from North America. Adastodon arvernensis and M. longirostris, 
together with a trilophodont species, occur in the crag-deposits of 
Norfolk and Suffolk. 


Family DINOTHERIID. 


An extinct family distinguished from the Elephantide by the whole 
series of permanent cheek-teeth being in use at the same time. 

Dinotherium4—Dentition of adult: i %, ¢ 8, p 3, m 3=22; all 
present at the 
same time, there 
being no hori- 
zontal succes- 
sion, but the 
premolars re- 
placing milk- 
teeth in the or- 
dinary manner. 
The presence or 
absence of upper 
incisors has not 
yet been clearly 
ascertained. 
Lower incisors : 
large, conical, descending, and slightly 
curved backwards, implanted in a gr ‘eatly 
thickened and deflected beak or pro- 
Iongation of the symphysis. In section 
they do not show the decussating striz 
characteristic of Mastodons and Ele- 
phants. Crowns of molars carrying strong 
transverse, crenulated ridges, with deep 
valleys between, much resembling the 
lower ones of the Tapirs. Ridge-formula Fic. 189.—Skull of Dinotherium 
of the permanent molar series: 2, 2, 3, feet seg a aa 
2,2. The three ridges of the first true Le ee aw eon 

, $ Pp) p, 3, 4, pre : 

molar are constant in both upper and 12 4 molars. 
lower jaws, although it is quite an anomalous character among 
Proboscideans for this molar to have more ridges than those which 
come behind it. The last milk-molar has also three ridges, the 


1 Kaup, Isis, vol. xxii. p. 401 (1829). 


436 UNGULATA 


penultimate but two. The cranium is much depressed, with com- 
paratively little development of air-cells. The remainder of the 
skeleton is imperfectly known, but apparently agrees in its general 
characters with that of the other Proboscideans. 

Remains of Dinotherium giganteum, an animal of elephantine 
proportions, strikingly characterised by the pair of huge tusks 
descending nearly vertically from the front of the lower jaw, were 
first discovered at Eppelsheim, near Darmstadt, and described by 
Kaup. They have since been met with in various Lower Pliocene 
and higher Miocene formations in the south of Germany, France, 
Greece, and Asia Minor. Closely allied forms also occur in the 
Lower Pliocene and Upper Miocene of India, but none are known 
from America. 


Suborder AMBLYPODA. 


Uintatherium.i—Among the most remarkable of the compara- 
tively recent discoveries in the higher Eocene formations of the 
western states of North America has been one of a group of 
animals of huge size, approaching that of the largest existing 
Elephants, presenting a combination of characters quite unlike 
those known among other recent or extinct creatures, and of which 
there were evidently many species living contemporaneously, but 
all of which became extinct before the close of the Eocene period. 
To form some idea of their appearance, we must imagine animals 
very elephantine in general proportions and in the structure of their 
limbs. The feet had five short toes. The tail, as in the Elephants, 
was long and slender, but the neck, though still short, was not so 
much abbreviated as in the Proboscideans, and there is no evidence 
that these animals possessed a trunk. The head differed greatly 
from that of the Elephants, being long and narrow, more like that 
of a Rhinoceros, and, as in that animal, was elevated behind into a 
great occipital crest, and it had developed upon its upper surface 
three pairs of conspicuous, laterally diverging protuberances—one 
pair in the parietal region, one on the maxillaries in front of the 
orbits, and one (much smaller) near the fore part of the elongated 
nasal bones. Whether these were merely covered by bosses of 
callous skin, as the rounded form and ruggedness of their extremities 
would indicate, or whether they formed the bases of attachment for 
horns of still greater extent, like those of the Rhinoceros or of the 
Cavicorn Ruminants, can only be a matter of conjecture. There 
were no upper incisors, but usually three on each side below, of 
comparatively small size, as was also the lower canine. A huge, 
compressed, curved, sharp-pointed canine tusk, very similar in form 


‘ Leidy, Proc. te, Nat. Sei, Philad, 1872, p. 169. 


AJSBLYPODA 437 


and position to that of the Musk-Deer, descended from each side 
of the upper jaw. These were present in both sexes, but very 
much smaller in the female, as was also the flange-like process of 
the lower jaw by which they were guarded. Behind these, and 
at some distance from them, were on each side above and below 
six cheek-teeth, of comparatively small size, placed in continuous 
series, each with a pair of oblique ridges conjoined internally and 
diverging externally in a V-like manner, and provided with a 
stout basal cingulum. The normal dental formula was therefore 
iQ ef p Rm Z= 34; and the dentition had thus already attained 
a remarkable degree of specialisation, although the brain was 
smaller and more rudimentary in characters than in almost any other 


Fig. 190.—Skeleton of Uintatkerium minabile. w natural size. (From Marsh, 
am, Journ, Sei, vol. xii. pl. 2.) 


known mammal. In its comparative length and the absence of a 
third trochanter the femur of these animals resembles that of the 
Proboscidea. The first discovered evidences of the existence of 
animals of this group were deseribed by Leidy in 1872. under the 
name of Uinfatherium (from the Uinta mountains, near which they 
were found). Subsequently the names Dinoecras, Tinoceras, Loro- 
lophodon, ete., have been applied to various members of the group. 
but the characters by which they are distinguished do not seem of 
suttcient importance to allow of their separation from the type 
genus Cinfatherium. 

Coryphodon* —Another interesting form referred to this suborder 
is Coryphodon, which appears to connect the Cintatheriide with the 
most primitive Perissodactrla. It was first described by Owen in 

1 For detailed descriptions and figures of this group, see Marsh. *t Monograph 


of the Dinocerata.” Rep. US. Geol. Stare. vol. x. (1SS4°. 
> Owen, Bris, Foss. Mamm, and Birds, p. 299 (1846. 


438 UNGULATA 


1846 from a fragment of a jaw from the London Clay. — Other 
remains were afterwards discovered in France, and lately in great 
abundance, indicating many species from the size of a Tapir to that 
of a Rhinoceros, in the Lower and Middle Eocenes of New Mexico 
and Wyoming in the United States. Coryphodon had forty-four 
teeth; the canines of both jaws were large and sharp pointed, 
and the molars had strongly pronounced oblique ridges. The 
general proportions were those of a Bear, but the tail was of 
moderate length, and the feet short and wide. The femur had 
a third trochanter ; and the cranium was devoid of protuberances. 


Fic. 191.—Palatal aspect of the cranium of Coryphodon hamatus, from the Wasatch Eocene of 
New Mexico. # natural size. (After Cope.) 


The genus should be regarded as the type of a distinct family 
Coryphodontide. 


Suborder CONDYLARTHRA. 


The term Condylarthra has been proposed by Professor Cope 
for a number of generalised and mostly comparatively small Ungu- 
lates, which were probably allied both to the Perissodactyla and 
Artiodactyla, but present characters separating them from those 
divisions as commonly defined. In the structure of the carpus 
and tarsus these forms (which are chiefly known to us from 
the Eocene of the United States) come nearer to the Hyracoidea 
than to any other existing type. As a rule they have the full 
dental formula; the molars are brachydont, generally bunodont, 
and in many instances also tritubercular ; while the premolars are 
always simpler than the molars. 

The humerus is quite peculiar among Ungulates in having an 


TOXODONTIA 439 


entepicondylar foramen; the femur has a third trochanter; and 
the form and relations of the astragalus are similar to those obtain- 
ing in the Carnivora. The feet are usually furnished with five 
functional digits, of which the ungual phalanges are pointed. In 
many respects the skeleton of these remarkably generalised Ungu- 
lates approximates so decidedly to a Carnivorous type as to have 
led paleontologists to conclude that the Ungulata and Carnivora 
are branches of an original common stock. 

In this work space only permits of allusion to a few of the 
more important types of this group. Leriptychus, which occurs in 
the lowest Eocene of New Mexico, is a bunodont type readily dis- 
tinguished by the vertical flutings of the premolars, and the small 
size of the incisors and canines. It has been suggested that this 
genus is closely related to the stock of the bunodont Artiodactyla. 
Of greater interest is the genus Phenucodus, which is regarded as the 
lowest factor in the series from which the modern Horse has been 
evolved, where it holds the position immediately below Hyraco- 
therinm or Systemodon (see p. 374). One of the species was about 
the size of a Bull-dog, while another might be compared to a small 
Leopard. The structure of the cheek-teeth is such as might readily 
be modified into that obtaining in Hyracotherivm , all the feet had 
five fully developed digits, and the tail was long. J/eniscotherinin 
and Hyracodontotherium are more specialised forms of somewhat 
later age, with a lophodont dentition; the latter genus heing 
European. 


Suborder TOXODONTIA. 


In addition to'the Macraucheniide and certain other forms 
noticed under the head of the Perissodactyla, the Tertiarics of 
South America have yielded some very remarkable forms of mam- 
malian life, the nature and affinities of which have greatly puzzled 
all zoologists who have attempted to unravel them. 

Nesodon and Torodon.—Among these Nesodon, from Patagonia, 
has the full typical Eutherian number of teeth; the crowns of the 
incisors being short, and the molars having a complex rhinocerotic 
type of structure somewhat intermediate between Homalodonto- 
therium (p. 412) and the following genus Toxulon, The typical 
species of Nvsodon was about as large as a Sheep, but nothing 
more is known of it than the teeth and portions of the skull. 

Tocodon is an animal about the size of a Hippopotamus ; it was 
first discovered by Darwin, and many specimens have since been 
found in Pleistocene deposits near Buenos Ayres, and described by 
Owen, Gervais, and Burmeister. The teeth consist of large incisors, 
very small lower canines, and strongly curved molars, all with 
persistent roots, the formula being apparently «3, ¢ 2, p 4, m #= 38. 


440 UNGULATA 


The cranial characters exhibit a combination of those found in both 
Perissodactyles and Artiodactyles, but the form of the hinder part 
of the palate and the absence of an alisphenoid canal belong to the 
latter ; and the tympanic, firmly fixed in between the squamosal 
and the exoccipital, ankylosed to both, and forming the floor of a 
long upward-directed meatus auditorius, is so exactly like that of 
the Suina that it is difficult to believe it does not indicate some 
real affinity to that group. These characters seem to outweigh in 
importance those by which some zoologists have linked Toxodon to . 
the Perissodactyla, and the absence of the third trochanter and the 
articulation of the fibula with the calcaneum tell in the same direc- 
tion. According to the recent observations of Ameghino the hind 
feet were certainly tridactylous, and the front feet probably so. 
The earlier allied genera Protorodon and Adinotherium are definitely 
known to have tridactylous front and hind feet, which conform to 
the Perissodactylate type, the bones of the proximal and distal 
rows of the carpus interlocking. Acrotherium, which has similar 
feet, differs from all other Ungulates, and indeed from all Eutherians 
except some individuals of the existing carnivorous genus Otocyon, 
in having eight cheek-teeth, five of which have been reckoned as 
premolars. 


Fic. 192.—Cranium and Lower Jaw of Typotherium eristatum. } natural size. From Gervais. 


Lypotherium. — Typotherium (Fig. 192), also called Mesotherium, 
from the same locality as Tovodon, was an animal rather larger than 


TILELODON? I 44t 


a Capybara, and of much the same general appearance. 1ts skeleton 
is completely known, and shows a singular combination of charactors, 
resembling Jeraden ora generalised Uneulate ou the one hand, and 
the Rodents, especially the Lerceitoay en the other la the presence 
of clavicles it ditfers from all known Uneulates, and in having two 
pairs of lower incisors from all Rodents. The teeth ave ioe Spy. 
meow. 

Peom the Tortiaries of various parts of South Amerion a number 
ot forms move ov less closely allied to Tevedan and Cypotheriiani have 
booew recently dosertbed. but as many of them are very impertectly 
known, and there is much doubt as to their generic position, it will 
bo unnecessary to refer to them further. 

It will thus be seen that, although our knowledge of many of 
these forms is still very limited, we may trace among them a eurious 
chain of attinines, which would seem to unite the Ungulates on the 
one hand with the Rodents on the other: bur further materials 
ave voqiured botore we ean establish with certainty so important a 
relationship, ene which, if true, would alter materially some of the 
prevailing views npon the elassitiention of mammals. 


Group TULODEN TIA. 


Here may be noticed a remarkable group of animals, eatled by 
Marsh, Tillodontia, the remains of which are found abundantly in 


Pra it- Shall ot Pilotheriwam Adiew.  }atualsss. Prem Moos): 


the Lower and Middle Rocene beds of Nerth Amertea. They seem 
to combine the characters of the Ungulata Rodentia, and Carnivora. 
In the genus Cideticrium of Marsh (probably identical with the pre- 
viously deseribed oii diccadus of Leidy) the skull (Pig. 198) resembled 
that of the Bears but the molar teeth were of the Cngnlate type, 
while the large incisers were very similar to these of the Rodents. 
The dental formula is i Soe hp doe S$. The tess pur of incisors 


a 


442 UNGULATA 


was very small; the upper molars were tritubercular, while the 
lower ones had crescentoid ridges as in Paleotherium. The skeleton 
resembled that of the Carnivores, but the scaphoid and lunar bones 
were distinct, and there was a third trochanter on the femur. The 
feet were plantigrade, and each had five digits, all with long pointed 
claws. In the allied genus Stylinodon all the teeth were rootless. 
Some forms were as large as a Tapir. 

These, with other more or less closely allied animals, such as 
Calamodon and Psittacotherium, constituting a group called Teeni- 
odonta, are included by Cope in his large order Bunotheria, to which 
also the existing Insectivora are referred. The dentition of some 
of these forms makes a remarkable approximation towards a Rodent 
type, while it has been suggested that there are also signs of remote 
Edentate affinities. The constantly increasing knowledge of these 
annectant forms adds to the difficulty so often referred to in this 
work of establishing anything like a definite classification of the 
heterodont mammals. An incisor tooth from the Swiss Eocene 
has recently been referred to Calamodon. 


Bibliography of Tagulata.—In addition to the works and memoirs mentioned 
under the different sections of the order, the following may be referred to :— 
W. Kowalevsky, ‘‘ Monographie des genus Anthracotherium,” Palwontographica 
1873; Id. ‘‘Sur l’Anchitherium aurelianense et sur V’histoire paléontologique 
des Chevaux,” Meém. de U'Acad. Imp. des Sciences de St. Pétersbourg, 1873; Id. 
“On the Osteology of the Hyopotamide,” Philosophical Transactions, 1873; 
L. Riitimeyer, ‘‘ Versuch einer natiirlichen Geschichte des Rindes,” etc., Neue 
Denks. der allgem. Schweiz. Gesellsch. fiir Naturwissenschaften, 1867: 1d. “Die 
Rinder der Tertiiir-Epoche,” bhand. der Schweiz. Paléont. Gesellsch. 1877 and 
1878; Id. ‘‘ Beitriige zu einer Natiirliche Geschichte der Hirsche,” ibid. 1880-1881; 
C. J. Forsyth-Major, “ Beitriige zur Geschichte der Fossilen Pferde,” bid. 1880: 
M. Schlosser, ‘‘Beitriige zur Kenntniss der Stammesgeschichte der Hufthiere 
und Versuch einer Systematik der Paar-und Unpaarhufer,” Morph. Jahrb. 1886 : 
E. D. Cope, ‘The Perissodactyla,” mer. Natural. 1887; M. Pavlow, “ Etudes 
sur histoire paléontologique des Ongulés,” Bull. Soc. Imp. Naturalistes Moscow, 
1887-1890. W. B. Scott and H. F. Osborn, “‘The Mammalia of the Uinta For- 
mation,” Trans. Amer. Phil. Soc. vol. xvi. (1889). 


CHAPTER X 


THE ORDER RODENTIA 


THE Rodentia, or Rodents, form a well-defined order, readily dis- 
tinguished by their large scalpriform incisors and the absence of any 
trace of canines. The existing forms are mostly of comparatively 
small size, and are generally of terrestrial habits, although a 
few are arboreal or natatorial. The dentition is diphyodont ; the 
mandible never has more than a single pair of incisors; the pre- 
molars are always below the full number, being very generally 4, or 
altogether wanting. The feet are plantigrade or semi-plantigrade, 
generally with five digits, and usually unguiculate, although occa- 
sionally of a subungulate type. Clavicles are present as a rule, 
although they may be imperfeet or rudimentary. 

The upper incisors resemble the lower in growing uninter- 
ruptedly from persistent pulps, and, except in the suborder 
Duplicidentata, agree with them in number; the premolars and 
molars may be rooted or rootless, with tuberculated or laminated 
crowns, and are arranged in an unbroken series. The orbits com- 
municate freely with the temporal fosse; the condyle of the 
mandible is clongated in the antero-posterior direction, and, through 
the absence of a postglenoid process to the squamosal, admits of a 
backward and forward motion of the jaw. The intestine (except 
in the J/yoride) has a large excum; the testes are inguinal or 
abdominal; the uterus is two-horned, the cornua either opening 
separately into the vagina or uniting to form a corpus uteri; the 
placenta is discoidal and deciduate ; and the smooth cerebral hemi- 
spheres do not extend backwards so as to cover any part of the 
cerebellum, 

The Rodents inelude by far the greatest number of species, and 
have the widest distribution of any of the orders of terrestrial 
mammals, being in fact cosmopolitan, although more abundant in 
some parts than in others. The total number of known existing 
species exceeds 900. South America may be regarded as their head- 


qt RODENTIA 


quarters at the present day; while in Australia and Madagascar 
they are represented only by a few genera. All the Rodents are 
exclusively herbivorous, and the whole of them gather their food 
by gnawing. They present considerable diversity of habits. Thus 
the Squirrels are arboreal, and some of them provided with a para- 
chute for taking flying leaps from tree to tree; the Hares are 
cursorial; the Jerboas agile jumpers; the Mole-Rats fossorial ; 
while the Beavers and Water-Voles are aquatic. In spite, how- 
ever, of this diversity of habits the Rodents present a remarkable 
similarity in general structure ; so much so, indeed, that the char- 
acters employed for distinguishing the various families and genera 
are comparatively trivial, and of slight structuralimportance. The 
skull of the Rodents is characterised by the invariable presence of 
the zygomatic arch, of which the middle portion is formed by the 
jugal (Fig. 7, p. 37); and, as already mentioned, the orbit communi- 
cates freely with the temporal fossa. There is invariably a long 
diastema separating the incisors from the cheek-teeth; and, with 
the exception of the Duplicidentata, the glenoid cavity of the squa- 
mosal is elongated antero-posteriorly. Postorbital processes of the 
frontals exist only in the Squirrels, Marmots, and Hares; in all 
other genera they are rudimentary or altogether absent; the 
zygoma never sends upwards a corresponding process; the lachry- 
mal foramen is always 
within the orbital margin; 
in many species the infra- 
orbital foramen is very 
large (in some as large as 
the orbit), and transmits 
part of the great masseter 
muscle (Fig. 194, m), by 
means of which the jaws 
are worked. The zygo- 
matie arch varies in its 

Fro. 194.—Skull of Hystrix cristata (juv.) t, Temporal degree of development, 
muscle ; m, masseter; m‘, portion of masseter transmitted and the position of the 


through the infraorbital foramen, the superior maxillary . baa 
nerve passing outwards between it and the maxillary bone. jugal therein is used as a 


distinguishing character 
for grouping the families ; the nasals are, with few exceptions, large, 
and extend far forwards; the parietals are moderate, and there is 
generally a distinct interparietal. The palate is narrow from before 
backwards—this being especially pronounced in the Hares, where it 
is reduced to a mere bridge between the premolars ; while in other 
cases, as in the Mole-Rats (Bathyergine), it is extremely narrow 
transversely, its width being less than that of one of the molar teeth. 
Auditory bulle are always present, and generally large ; in some 
genera, as in the Gerbilles and Jerboas, there are also supplemental 


RODENTIA 445 


mastoid bull forming great hemispherical bony swellings at the back 
of the skull (see Fig. 7, Per); and in these genera, and in the true 
Hares, the meatus auditorius is tubular and directed upwards and 
backwards. The mandible is characterised by the abruptly nar- 
rowed and rounded symphysial part supporting the large incisors, 
as well as by the small size of the coronoid process and the great 
development of the angular portion. 

The dental formula varies from i 3, ¢ $, p 3, m 2 (total 28) 
in the Duplicidentata to i 4, ¢ 2, p 2, m 2 (total 12) in Hydromys, 
eromys, and one species of Heterocephalus; but in’ the great 
majority of forms it is very constant, i4,c 9, p ay m & being 
very typical. Only in the Duplicidentata is there a second pair of 
upper incisors, which are of very small size, and situated immedi- 
ately behind the large normal pair. This group is also peculiar in 
that the enamel of the incisors is not confined to their anterior 
surfaces, but extends partially on to their sides. It is by reason 
of the thick layer of enamel on their anterior surface and its 
absence from the posterior surface that the incisors maintain their 
sharp chisel-like edge, which is so essentially characteristic of the 
order. Both the upper and the lower incisors are regularly curved 
—the curvature being somewhat greater in the upper ones—and 
since they grew continuously from persistent pulps, it is quite 
evident that should any accident, such as the loss of one of them, 
or displacement by 
fracture of the jaw, 
prevent the regula- 
tion of the length 
by attrition against 
one another, the 
unopposed tooth 
will gradually 
curve upon itself 


until a complete Fic. 195.—Vertical and longitudinal section, through skull of 
circle or more has the Beaver (Castor fiber) showing the cerebral cavity, the greatly 

developed turbinal lamellz, the mode of implantation of the large 
been formed, the incisor, and the curved rootless molars. 


tooth, perhaps, 
passing during its growth through some part of the animal’s head. 
The molars, as already mentioned, may be rooted or rootless, tuber- 
culated or laminated; this diversity of structure occurring even 
in the same family. When there are more than three cheek- 
teeth those in front of the last three have succeeded milk-teeth, 
and must therefore be considered premolars. In some species, as 
in the Agoutis (Dasyproctide), the milk-teeth are long retained, 
while in the allied Cavies (Caviide) they are shed before birth. 
There are generally nineteen dorso-lumbar vertebre (thirteen 
dorsal and six lumbar), their form varying in the different genera. 


446 RODENTIA 


In the cursorial and leaping species the lumbar transverse processes 
are generally very long, and in the Hares there are large com- 
pressed hypapophyses. The caudal vertebre exhibit great variety 
in structure, being in a rudimentary condition in the Guinea-Pig, 
while in the Jumping Hares and prehensile-tailed Porcupines they 
are of very large dimensions. The scapula is usually narrow, with 
a long acromion ; the clavicles may be altogether absent or imper- 
fect, as in the Porcupines, Cavies, and Hares, but in most species 
they are well developed. In all existing forms the humerus has 
no entepicondylar foramen, and the radius and ulna are distinct. 
In most species the manus has five digits, with phalanges normally 
developed ; the pollex being rarely rudimentary or absent. The 
pelvis has well-developed ischia and pubes, meeting in a long, and 
usually bony, symphysis. The femur varies considerably in form, 
but generally has a well-defined third trochanter; in the Sciurine 
and Hystricine Rodents the tibia and fibula are distinct, but in the 
Rats and other Murines, and in the Hares, these bones are united, 
often high up ; the pes is much more variable than the manus, the 
digits varying in number from five, as in the Squirrels and Rats, to 
four, as in the Hares, or even three, as in the Capybara, Viscacha, 
and Agouti; in the Dipodide the metatarsals are greatly elongated, 
and in some of the species, as in the Jerboas, they are ankylosed 
together. 

The mouth is divided into two cavities communicating by a 
constricted orifice, an anterior one containing the large incisors, and 
a posterior one in which the molars are placed; the hairy integu- 
ment of the face being continued inwards behind the incisors. This 
peculiar arrangement evidently prevents substances not intended 
for food getting into the mouth, as when the animal is engaged in 
gnawing through an obstacle. In the Hares and Pacas the inside 
of the cheeks is hairy, and in some species, as in the Pouched Rats 
and Hamsters, there are large internal cheek-pouches lined with 
the hairy integument, which open near the angles of the mouth 
and extend backwards behind the ears. In the New World 
Pouched Rats (Geomyidw) the pouches open externally on the 
cheeks. The tongue presents little variability in length, being 
always short and compressed, with an obtuse apex never protruded 
beyond the incisors. In most species there are three circumvallate 
papille at the base; and the apical portion is generally covered 
with small filiform papille, some of which in the Porcupines 
(Hystriz) become greatly enlarged, forming toothed spines. The 
stomach varies in form from the simple oval sac of the Squirrel ‘to 
the complex ruminant-like organ of the Lemming. In the Water- 
Vole (Arvicola umphibius) and the Agouti (Dusyprocta agutt) it is 
strongly constricted between the cesophagus and pylorus. In the 
common Dormouse the esophagus immediately before entering the 


RODENTIA 447. 


stomach is much dilated, forming a large egg-shaped sac with 
thickened glandular walls; and in some other species, as in 
Lophiomys imhausi and in the Beaver, glandular masses are 
attached to and open into the 
cardiac or pyloric pouches. The 
alimentary canal (Fig. 196) of 
all Rodents, with the exception 
of the Dormice (J/yoridw), has 
a czecum, which is often of great 
length and sacculated, as in the 
Hares, Water-Voles, and Poreu- 
pines. In some instances, as in 
the Hamster and Water-Vole, 
the long colon is spirally twisted 
upon itself near its commence- 
ment. The liver is typically 
divided in all, but the lobes are 
variously subdivided in the 
different species (in Capromys \ 
they are divided into minute 
lobules); and the gall-bladder, 
though present in most, is absent 
in a few. In most species the Fie. 19.—Alimentary canal of Rat (Mus decu- 
penis (which is generally pro- puvwtty grater ort of th gm testo 
vided with a bone) can be more j, jteam: em, excum; ¢, colon. , 
or less completely retracted 

within the fold of integument surrounding the anus, where it lies 
curved backwards upon itself under cover of the integument. It 
may, however, be carried forward some distance in front of 
the anal orifice, from which in the breeding season, as in the 
Voles and Marmots, the prominent testicular mass separates it. 
The testes in the rutting season form projections in the groins, 
but (except in the Duplicidentata) do not completely leave the 
cavity of the abdomen. Prostatic glands and, except in the 
Duplicidentata, vesiculze seminales are present in all. The uterus 
may be double, each division opening by a separate aperture into 
a common vagina, as in Leporide, Seiurida, and Hydrocharus, or 
completely two-horned, as in most species. The mamme vary in 
number and position from the single abdominal pair of the Guinea- 
Pig to the ten thoracico-abdominal pairs found in some of the 
Rats. In the Octodontidw the mamm are placed high up on the 
sides of the body. 

The peculiar odour evolved by many Rodents is due to the 
secretions of special glands, which may open either into the 
prepuce, as in Jus, Arvicola, Cricetus, ete., or into the reetum, as in 
Arctomys and tulacodus, or into the passage common to both, as in 


448 RODENTIA 


the Beaver, or again, into pouches opening near the anus, as in the 
Hare, Agouti, and Jerboa. 

The integument is generally thin, and the panniculus carnosus 
(the sheet of muscle underlying the skin) rarely much developed. 
The fur varies exceedingly in character. Thus it may be very 
fine and soft, as in the Chinchillas and Hares, in others more 
or less replaced by spines on the upper surface, as in the Spiny- 
Rats and Porcupines; in several genera, as in Xerus, Acunthomys, 
Platacanthomys, Echinothriz, Loncheres, and Echinomys, the spines are 
flattened. In the muscular structures the chief peculiarities are 
noticeable in the comparatively small size of the temporal muscles, 
and in the great double masseters (Fig. 194), which are the prin- 
cipal agents in gnawing ; the digastrics also are remarkable for their 
well-defined central tendon, and in many species their anterior bellies 
are united between the mandibular rami ; the cleidomastoid generally 
arises from the basioccipital, and the pectoralis major is connected 
with the latissimus dorsi; in the Poreupines and Hares the tendons 
of the flexor digitorum longus and flexor hallucis longus are con- 
nected in the foot, while in the Rats and Squirrels they are separate, 
and the flexor digitorum longus is generally inserted into the 
metatarsal of the hallux.t 

Rodents are tolerably well represented in a fossil condition from 
the period of the Upper Eocene, while if Decticadapis, of the Lower 
Eocene of Rheims, is rightly referred to it the order dates from the 
oldest Tertiary. All the fossil forms at present known are, however, 
essentially true Rodents, and afford no clue as to the relations of 
the order with other mammals. The remote affinities of the 
Rodents to the Proboscidea, as well as their more marked resem- 
blances to Typotherium, have been already mentioned. Whether 
there is a real genetic affinity (as Professor Cope suggests) with the 
Tillodontia cannot be decided with the evidence at present available. 


Suborder SIMPLICIDENTATA. 


Only one pair of upper incisors, having their enamel confined to 
their front surfaces. Incisive foramina moderate and distinct : 
fibula not articulating with the caleaneum. Testes abdominal, and 
descending periodically only into a temporary sessile scrotum. 


Section SCIUROMORPHA. 


Zyzomatie arch slender, chiefly formed by the jugal, which is 
not supported by a long maxillary process extending backwards 
beneath it; postorbital processes of frontal present or absent ; 


1 See G. E. Dobson, Journ. -trat. Phys. vol. xvii. 


ANOMALURIDE 449 


infraorbital opening small (except in Anomalurus); mandible with 
the angular part arising from the inferior surface of the bony 
socket of the lower incisor ; clavicles well developed; fibula distinct. 


Family ANOMALURID. 


Arboreal forms, having their limbs connected by a cutaneous 
expansion supported by a cartilaginous process arising from the 
olecranon ; tail long and hairy, with large imbricated scales on its 


Fic. 197.—Anomalurus fulgens. From Alston, Proc. Zool. Soc. 1875. 


inferior surface near the root ; sixteen pairs of ribs; no postorbital 
processes on the frontals; p 4; molars not tuberculate, with 
transverse enamel-folds. Confined to the Ethiopian region. 
Anomalurus,! with several species from West and Central Africa, 
alone represents the family. The peculiar caudal scales, which 
evidently assist the animal in climbing, and the position of the 
cartilaginous support of the parachute, are well shown in Fig. 197. 
All the species but two are from Western Africa ; 4. orientalis occurs 
near Zanzibar, and A. pusillus is from the equatorial regions of that 


1 Waterhouse, Proc. Zool. Soc, 1842, p. 124. 
29 


450 RODENTIA 


continent. According to Mr. O. Thomas,! the latter “little animal 
is most nearly allied to the West-African 4. beccrofti, but differs 
from that species in its duller and less yellow upper side, in the 
entire absence of rufous on its neck and belly, and, as from all the 
other described species, in its diminutive size.” 


Family Scrcrw-£. 


Arboreal or terrestrial forms, with cylindrical hairy tails, with- 
out scales, and with 
twelve or thirteen 
pairs of ribs. Skull 
(Figs. 198, 199) with 
distinct postorbital 
processes; infra- 
orbital opening 
small; palate broad; 
P%; first upper pre- 
molar very small or 
deciduous; molars 
rooted, tubercular. 
Subfamily Seiur- 
ine.—TIncisors com- 
Fic. 198,—Lateral view of skull of American Marmot pressed form slen- 
(Arctomys monax). der ; tail long and. 
hairy. Cosmopolitan (excluding Australian region). 

This subfamily includes the true Squirrels, of which seven 
existing genera are usually recognised. 

Seturus.2—Tail long and bushy ; ears generally well developed, 
pointed, often tufted ; 
feet adapted for climb- 
ing, the anterior hav- 
ing four digits and 
a rudimentary pollex, 
and the posterior with 
five digits, all of which § 
have long, curved, and 
sharp claws. Mamme, 
from four to six. Skull 
(Fig. 199) lightly built, 
with long postorbital 
processes. Penultimate 


Fig. 199.—Palatal Aspect of cranium of Squirrel (Sefurus 
upper premolar, when bicolor). Natural size, 


present, minute. 


» Proce. Zool. Sov, 1882, p. 8. * Linn. Syst, Vat. 12th ed. vol. i. p. $08 (1766). 


SCIURIDE 451 


True Squirrels are found in most of the temperate and tropical 
regions of the world, exclusive of Madagascar and the Australian 
region. They are, however, most abundant in the Malayan part of 
the Oriental region, and attain their largest size and most brilliant 
coloration in the tropics. Their size is very variable, so that 
whereas S. soricinus, of Borneo, is no larger than a Mouse, S. bicolor, 
of the Malayan region, is nearly as large as a Cat. The common 


Fic. 200.—Burmese Squirrel (Sciurus pygerythrus). After Anderson. 


English Squirrel (S. vzlyaris) is found over the whole of the Palearctic 
region, reaching in one direction from Ireland to Japan, and in the 
other from the north of Italy to Lapland ; its remains occur in the 
Norfolk “Forest-bed.” In the Malayan region “nearly all the 
numerous species are brilliantly marked, and many are ornamented 
with variously coloured longitudinal stripes along their bodies. One 
of the commonest and best known of the striped species is the little 
Indian Palm-Squirrel (S. palimeruim), which in large numbers runs 
about every Indian village. Another Oriental species (S. cuniceps) 
presents almost the only known instance among mammals of the 


452. RODENTIA 


temporary assumption during the breeding season of a distinctly 
ornamental coat, corresponding to the breeding-plumage of birds. 
For the greater part of the year the animal is of a uniform gray 
colour ; but about December its back becomes a brilliant orange- 
yellow, which lasts until about March, when it is again replaced by 
gray. The Squirrel shown in Fig. 200 is a native of Burma and 
Tenasserim, and is closely allied to S. caniceps, but goes through no 
seasonal change of colour. 

“The number of species in the genus is about 75, of which 3 
belong to the Palearctic, 15 to the Ethiopian, about 40 to the 
Oriental, and 16 to the combined Nearctic and Neotropical regions ” 
(Thomas). 

Fossil species referred to Sctwrus are found in the European 
Tertiaries down to the Phosphorites of Central France, while others 
occur in the White River Miocene of the United States. 

ELhithrosciwrus.i—A very striking Squirrel, confined to Borneo, 
and as yet only known from three or four examples, bas been 
separated generically under this name. The general shape of its 
skull is very different from that of other Squirrels; but its most 
peculiar characteristic is the presence of from seven to ten minute 
parallel vertical grooves running down the front face of its incisors ; 
no other Squirrel having really grooved incisors, and no other 
member of the whole order incisive grooves resembling these. 
Its premolars number 3, and its molars are simpler and less ridged 
than in the other genera. This Squirrel (2. macrotis) is far larger 
than the English, with an enormously long bushy tail, long tufted 
ears, and black and white bands down its sides. 

Xerus.2—Fur coarse and spiny. Claws long and comparatively 
straight. Ear-conchs minute or absent. Skull with the postorbital 
processes short and directed backwards, the bony palate prolonged 
considerably behind the tooth-row, and the external ridge on the 
front face of the anterior zygomatic root more developed, and 
continued much farther upwards than in Sciurus. Premolars 2; 
molars as in Sciurus. Mammztwo. This genus contains four well- 
marked species, known as Spiny Squirrels, all natives of Africa. 
They are terrestrial in their habits, living in burrows which they 
dig for themselves. X. getulus, a striped species of North Africa, 
has much the size and appearance of the Indian Palm-Squirrel ; 
all the others are a little larger than the English Squirrel. 

Tamias.*—All the members of this genus are characterised by 
the possession of internal cheek-pouches, and by their style of colora- 
tion ; being ornamented on the back with alternate light and dark 
bands. Their skulls are slenderer and lighter than those of the 

1 Gray, Ann. Mag. Nat, Hist. ser. 3, vol. xx. p- 272 (1867). 
? Hemprich and Ehrenberg, Symbol. Phys. Mam. vol. i. (1882). 
3 Illiger, Prodromus Syst. Mam. p. 83 (1811). 


SCLURIDAE 453 


true Squirrels, from which they differ in several unimportant 
details. There is only one functional premolar—the small anterior 
one usually found in Sciwrus being either absent altogether or quite 
small and functionless. There are some four well-defined species, 
all found in North America, one (7. asiaticus) extending also through 
Siberia into Eastern Europe:!. They are generally known as Ground- 
Squirrels, but in America, where they are among the commonest 
and best known of the indigenous Rodents, as “ Chipmunks.” The 
members of this genus seem to lead into the genus Spermophilus, 
so that the division of the Sciuride into two subfamilies, although 
convenient for classification, is rather artificial. 

Remains of Tams, probably belonging to existing species, occur 
in the Pleistocene deposits of Europe and Nebraska. 

Pteromys* and Sciuropterus.2—The Flying Squirrels, although in- 
capable of true flight, can yet float through the air for considerable 
distances by the aid of an extension of skin connecting their fore 
and hind limbs, and forming a sort of parachute. This parachute 
is merely a lateral extension of the ordinary skin of the body, 
which passes outwards between the limbs and terminates at the 
wrists and ankles. In addition to the lateral membrane there is a 
narrow and inconspicuous one passing from the cheek along the 
front of the shoulder to the front of the wrist, and another—at 
least in the larger species—stretching across behind the body from 
ankle to ankle and involving the base of the tail. The Flying 
Squirrels are divided into three genera. Of those with a normal 
dentition Pteromys contains the larger and Sciwropterus the smaller 
species. The two differ in certain details of dentition, as well as 
in the greater development in the former of the expanded mem- 
branes, especially of the “interfemoral” or posterior membrane, 
which in the latter is almost wholly absent. In Pteromys the tail 
is cylindrical and comparatively thin, while in Sciuropterus it is 
broad, flat, and laterally expanded, and evidently compensates for 
the absence of the interfemoral membrane by acting as a supple- 
mentary parachute. In appearance Flying Squirrels resemble the 
other forms, although they are even more beautifully coloured. 
Their habits, food, etc., are also very similar to those of the 
true Squirrels, except that they are more decidedly nocturnal, 
and are therefore less often seen by the traveller; their peculiar 
shrill cry is, however, well known to all who have camped out in 
the regions which they inhabit. Their mode of flight is precisely 
similar to that of the Flying Phalangers of Australia. Of each of 
the two genera there are about thirteen or fourteen species, all 


1 Some American zoologists have recently proposed to raise a large number of 
the forms usually regarded as local races to the rank of species. 

2 Cuvier, Légons d’ Anatomie Comp. (1800). : 

3 Cuvier, Ann. du Muséum, vol. x. p. 126 (1825). 


a5 RODENTIA 


natives of the Oriental region, except that one of Sciwropterus is found 
in North America, and another in Siberia and Eastern Europe. 

Eupetaurus.1—Externally as in Pteromys, except that the claws 
are less sharp. Skull with a more produced muzzle than in the 
latter, more distinct supraorbital notches, longer anterior palatal 
foramina, and a shorter bony palate. Cheek-teeth differing from 
those of all other Sciurid in their hypsodont character. One large 
species (E. cinereus), from Gilgit and adjacent districts on the 
extreme north-west of Kashmir territory. This fine Flying Squirrel 
is chiefly known by one entire specimen and some imperfect skins. 

Extinet Genero.—The genera Pseudosciurus and Sviuroides, from 
the Upper Eocene of Europe, have the molar teeth more elongated 
than in Seiurus. Gymunaptychus with p +, from the North American 
Miocene, approximates in the structure of its molars to Tamias. 
Meniscomys (p 2), from the latter deposits, together with Sciurodon 
of the French Phosphorites, are regarded as Squirrels showing signs 
of affinity with the Huplodontide. 

Subfamily Aretomyinze.—Incisors not compressed: typically 
the form stout, and the tail comparatively short. This subfamily 
comprises burrowing forms which may be collectively known as 
Marmots ; as already mentioned, they are so intimately connected 
with the preceding subfamily that the division into two groups is 
purely a matter of convenience. They are confined to the Pale- 
arctic and Nearetic regions. 

_{rctomys.°—External form stout and heavy, ears short, tail 
short and hairy, chéek-pouches rudimentary or absent. Fore feet 
with four well-developed digits, and a rudimentary pollex provided 
with a flat nail. Skull (Fig. 198) large and heavy, with the post- 
orbital process stout, and at right angles to the axis. Incisors 
broad and powerful. First upper premolar nearly as large as the 
second, Molar series nearly parallel, scarcely converging behind 
at all. 

The various species of true Marmot, which exceed a dozen in 
number, are all much alike in general appearance, ranging in size 
from about 15 to 25 inches in length, with tails from 3 to 12 inches 
long. 

The Alpine Marmot (Fig. 201) is peculiar to Europe, being 
found in the Alps, Pyrenees, and Carpathians ; its remains occur in 
European Pleistocene deposits. 4. bobac occurs in Eastern Europe 
and Siberia. Several species (¢.g. 4. monar, Fig. 198) are found 
in the Nearctic region, and many in Kashmir and Central Asia. 
The long-tailed Red Marmot (.4. caudatus) is a fine Himalayan 
species, which may be seen on the mountain passes to the north of 
the valley of Kashmir, as soon as the snow begins to disappear, 


2 0. Thomas, Journ, As. Soc. Bengal, vol. vii. p. 256 (1888). 
* Schreber, Seéugethierv, vol. iv. p. 721 (1792). 


SCIURIDE 455 


sitting at the entrance to its burrow, which is generally beneath a 
rhubarb plant. 

The following account of the habits of the Alpine Marmot is 
given by Professor Blasius: “ Marmots live high up in the snowy 
regions of the mountains, generally preferring exposed cliffs, whence 
they may have a clear view of any approaching danger, for which, 
while quietly basking in the sun or actively running about in search 
of food, a constant watch is kept. When one of them raises the cry 
of warning, the loud piercing whistle so well known to travellers 


Fic. 201,—Alpine Marmot (dArctomys marmotta). After Brehm. 


in the Alps, they all instantly take to flight and hide themselves in 
holes and crannies among the rocks, often not reappearing at the 
entrance of their hiding-places until several hours have elapsed, and 
then frequently standing motionless on the look-out for a still longer 
period. Their food consists of the roots and leaves of various 
Alpine plants, which, like squirrels, they lift to their mouths with 
their fore paws. For their winter quarters they make a large 
round burrow, with but one entrance, and ending in a sleeping-place 
thickly lined with hay. Here often from ten to fifteen Marmots 
pass the winter, all lying closely packed together fast asleep until 
the spring.” 

Cynomys.1—Size and form intermediate between <Arctomys and 
Spermophilus. Ears and tail short. Cheek-pouches shallow. Fore 

1 Rafinesque, Amer. Monthly Mag. vol. ii. p. 45 (1817). 


456 RODENTIA 


feet with five claws, that on the pollex as large as that on the fifth 
toe. Skull (Fig. 202) heavily built, with the postorbital processes 
directed outwards. Dentition (as shown in Fig. 202) remarkably 
heavy, the molar teeth 
differing from those 
of Arctomys and Sper- 
mophilus by having 
three instead of two 
transverse grooves on 
their crowns. First 
premolar nearly as 
large as the second. 
Molar series strongly 
convergent behind. 

Two species of 
Prairie Marmots, or, 
Fra. 202.—Palatal aspect of the craniuin of the Prairie Marmot as they areoften called, 

(Cynomys ludovicianus). “ Prairie - Dogs,” are 
found in North America. They live together in large communities, 
inhabiting burrows excavated at short distances apart, and feeding 
on the buffalo-grass which covers the plains. The small burrowing 
owl (Athene cuntcularia) and the rattlesnake are often found inhab- 
iting their burrows; the former probably availing itself of the 
convenience of a ready-made habitation, the latter coming there to 
feed on the young Marmots. 

Spermophilus..Size much smaller than in either of the preced- 
* ing genera; form more slender and squirrel-like. Tail very variable, 
from 1 to 8 or 9 inches in length. Cheek-pouches always present. 
Fore feet with four well-developed toes and a rudimentary pollex, 
of which the claw may be either present or absent. Skull more 
lightly built than in the other preceding genera, with the postorbital 
processes slender and directed backwards. Molar series nearly 
parallel, as in Aretomys, but all these teeth much smaller and lighter ; 
first premolar simply rounded, never more than about one-third of 
the size of the second. 

The Pouched Marmots, or Sousliks, have nearly the same dis- 
tribution as Tamias, and are represented by a considerable number 
of species. They present a far greater range of variation than 
is found among the true Marmots, some of them, such as the 
European species, being scarcely as large as a common squirrel, 
almost entirely without external ears, and with the tail reduced to 
a mere stump, barely an inch long, while others are more than 
three times this size, with large and often tufted ears, and long 
bushy squirrel-like tails. Professor Blasius gives the following 
details of the habits of the common European Souslik (8. citillus) : 

1 F, Cuvier, Mem. du Muséum, vol. vi. p. 298 (1822). 


CASTORIDAE 457 


“Tt lives in dry treeless plains, especially on a sandy or clayey soil, 
and is never found either in forests or on swampy ground. It 
forms burrows, often 6 or 8 feet deep, in which food is stored up 
and the winter sleep takes place. Each burrow has but one 
entrance, which is closed up when winter approaches,—a second 
hole, however, being previously formed from the sleeping-place to 
just below the surface of the ground. The second hole is opened 
the next year, and used as the ordinary entrance, so that the 
number of closed-up holes round a burrow gives an indication of 
the length of time that it has been occupied. Sousliks ordinarily 
feed on roots, seeds, berries, etc., but occasionally also on animal 
food, preying readily on eggs, small birds, and mice, the remains of 
these latter being often found in their burrows. They bring forth 
in the spring from four to eight young ones, which, if taken early, 
may be easily tamed. They are often eaten by the peasants, the 
inhabitants of the Russian steppes considering their flesh an 
especial delicacy.” 

Remains of Spermophilus are not uncommon in European Tertiary 
deposits, some belonging to living and others to extinct species. 

Extinct Genera.—Plesispermophilus, from the Upper Eocene Phos- 
phorites of Central France, appears to be closely allied to the 
Sousliks. Plesiarctomys (Sciuravus or Paramys), which is common 
to the Middle Tertiaries of Europe and North America, appears to 
be a generalised form, showing some resemblance both to Arctomys 
and Seiwrus, but with tritubercular upper molars and no postorbital 
processes to the skull; in the latter respect agreeing with the next 
family. In the size of the preorbital vacuity the skull resembles the 
Hystricomorpha. 


Family HAPLODONTID&. 


Distinguished from the Sciwride hy the absence of postorbital 
processes to the frontals, the depressed skull, and the rootless cheek- 
teeth. Premolars ?; the penultimate upper one small. 

Haplodon1—H. rufus and H. inajor, of North America, west’ of 
the Rocky Mountains, are the only representatives of the family; 
their habits are similar to those of Cynomys. 


Family CASTORIDE. 


Skull massive, without postorbital processes, the angle of the 
mandible rounded, and the cheek-teeth rootless, with re-entering 
enamel-folds. Premolars +. Habits natatorial. 

Castor.2—The upper molars are subequal, each with one internal 


1 Richardson, Zool. Journ. vol. iv. p. 334 (1829). Amended. 
2 Linn. Syst. Nat. 12th ed. vol. i. p. 78 (1766). 


458 RODENTIA 


and two external enamel-folds ; the stomach has a large glandular 
mass situated to the right of the esophageal orifice; the anal and 
urethro-genital orifices open within a common cloaca; the tail is 
broad, horizontally flattened, and naked; and the hind feet are 
webbed. One or two species, Palearctic and Nearctic. 

Zoologists are not yet of accord as to whether the European 
and American Beavers should be regarded as distinct species or as 
local races; the general concensus of opinion being in favour of 
the latter view. 

The European Beaver (C. fiber) was at one time an inhabitant 
of the British Isles, having been found, according to Pennant, in 
certain Welsh rivers so late as the twelfth century, while subfossil 
remains of it occur in the peat-beds of many parts of the country. 
In Scandinavia Beavers are still found in the neighbourhood of 
Arendal. Isolated pairs are occasionally met with on the banks of 
the Rhone, Weser, and Elbe; and a considerable number are kept 
in a park belonging to the Emperor of Austria, on the banks of 
the Danube. They also occur sparingly in Russia and Poland, 
in the streams of the Ural Mountains, and in those which flow 
into the Caspian. They live in burrows on the banks of rivers, 
like the Water-Rat, and show little of the architectural instinct 
so conspicuous in the American form, but this may be owing to 
unfavourable external conditions rather than to want of the 
faculty ; for there is a well-authenticated instance of a colony of 
Beavers,-on a small stream near Magdeburg, whose habitations 
and dam were exactly similar to those found in America. 

The American Beaver (C. canadensis) extends over that part of 
the American continent included between the Arctic circle and 
the tropic of Cancer; owing, however, to the gradual spread of 
population over part of this area, and still more to the enormous 
quantity of skins that, towards the end of last and the beginning 
of the present century, were exported to Europe, numbering about 
200,000 annually, this species is in imminent danger of extirpation. 
It is distinguished from the European Beaver by the shorter and 
somewhat wider nasals. 

Remains of extinct species of Castor occur in the Pliocene of 
Europe, and in the North American Miocene; the one from the 
last-mentioned deposits being of small size, and separated by some 
writers as Lucastor. 

Extinct Genera.—A very large Beaver known as T, rogontherium 
(Diobroticus), and distinguished by the nature of the enamel-folds of 
the molars, occurs in the Upper Pliocene and Pleistocene of Europe. 
Chalicomys (Steneofiber) is a considerably smaller form from the 
Miocene of Europe and the United States, distinguished from all 
existing Rodents by the presence of an entepicondylar foramen in 
the humerus. Palwocastor, of the North American Miocene, is allied. 


MVOXIDE 459 


Section MYoOMORPHA. 


Skull (Fig. 203), with slender zygomatic arch, in which the 
jugal seldom extends 
far forwards, being 
usually supported by 
the long zygomatic 
process of the maxilla; 
no postorbital process ; 
infraorbital vacuity 
variable; angle of 
mandible, except in 
the Bathyergine, rising 
from the inferior sur- 
face of the incisive 
alveolus. Clavicles 
well developed, except 
in Lophiomys. Tibia 
and fibula united. 


Fic. 203.—Side view of skull of Fiber zibethicus, natural size. 


Fanily Myoxip&. 


Small arboreal forms, with long hairy tails, large eyes and ears, 
and short fore limbs. No cecum in the intestine. Skull with 
narrow frontals, a high and narrow infraorbital vacuity of moderate 
size, and a long and slender coronoid process to the mandible. 
Premolars +; molars rooted, with transverse enamel-folds. 

The Dormice form a natural family allied to the Squirrels in 
form and habits, and confined to the Palwarctic and Ethiopian 
regions. The absence of the cecum distinguishes them from all 
other members of the order. They are usually divided into the 
following five genera, but some of these are of very doubtful value, 
and it might be preferable to retain Muscardinus and include all 
the others in Myoxus.+ 

Myoxus.2—Represented by the European JZ. glis, and charac- 
terised by the bushy distichous tail, simple stomach, and the large 
size and complex enamel-folds of the molars, which have flat crowns. 

Eliomys.2—Tail tufted and distichous; stomach simple; and 
the molars small, with concave crowns and indistinct enamel-folds. 
Some seven species, Ethiopian and Palearctic. 

Graphiurus.A—Tail short, cylindrical, and tufted at the end; 


1 For a monograph of the Afyoxide, see C. L. Reuvens, Die Myoxide, ete., 
4to, Leyden, 1890. : 2 Schreber, Stiugethiere, vol. iv. p. 824 (1792). 

3 Wagner, Abh. baier. Ahad. vol. iii. p. 179 (1843). 

4 F. Cuvier, Zammiferes, 60me livr. (1845). 


460 RODENTIA 


molars very small, with the enamel-folds almost absent. Some 
three Ethiopian species. 

Claviglis..Represented by one West African species, said to be 
distinguished from all other forms by the shorter tail, which is 
more distinctly pencilled. The right to generic distinction is, how- 
ever, very problematical. 

Muscardinus.2—Includes the Common Dormouse (Jf. avellanarius) 
of Europe, distinguished by the cylindrical bushy tail, and thickened 
glandular walls of the cardiac extremity of the cesophagus; the 
molars have flat crowns, with complex enamel-folds. 

Fossil Dormice.—Using the generic term Myoxus in a more 
extended sense than the above, it has existed in Europe from the 
date of the Upper Eocene. A species nearly as large as a Guinea- 
Pig, with very complex molars, is common in the Pleistocene of 
Malta. 


Family LoPHIOMYID2. 


The genus Lophiomys,? represented only by L. imhausi (Fig. 


Fic. 204.—Lophiomys imhausi. From Milne-Edwards. 


204) of North-East Africa, differs from the typical Muwridew in 
having the temporal fossee roofed over by a thin plate of bone, | 
rudimentary clavicles, and an opposable hallux. On these grounds 
it has been made the type of a family, but since all the features 
are Murine—the dentition being that of a typical Cricetine—it 


 Jentink, Notes Leyd. Mus. vol. x. p. 41 (1888), 
* Kaup, Entwickl. Europ. Thierwelt, p- 189 (1829). 
* A, Milne-Edwards, Z’Jnstitut, vol. xxxv. p. 46 (1867). 


MURIDA 461 


appears doubtful whether that distinction is justifiable. The hair 
forms a crest along on the back, and is of a peculiar structure. 
The habits of this Rodent are arboreal. 


Family MuRIDA. 


Skull (Fig. 203) with contracted frontals ; a short and slender 
jugal, generally reduced to a splint between the zygomatic pro- 
cesses of the maxilla and squamosal ; the lower root of the former 
process more or less flattened into a perpendicular plate ; typically, 
the infraorbital vacuity tall, and wide above and narrow below. 
Lower incisors compressed ; no premolars ;! molars rooted, or root- 
less, tuberculate, or with angular enamel-folds. Pollex rudimental ; 
tail generally nearly naked and scaly. Habits various, but mostly 
terrestrial. 

This large and cosmopolitan family, which includes more than 
a third of the existing Rodents, is represented by about forty 
genera. 

Subfamily Hydromyinz.—Molars 2 in number, rooted, and 
divided into transverse lobes. Represented by two Australasian 
genera. 

Hydromys.2—External form modified for an aquatic life. Tip 
of muzzle extensively haired, so that the nostrils can be closed. 
Skull with the infraorbital vacuity crescentic, scarcely narrowed 
below, and its external wall without the perpendicular zygomatic 
plate characteristic of most of the family ; incisive foramina very 
small. 

Two species, with habits like those of the Water Voles, are 
known from Australia, Tasmania, and New Guinea. In the 
typical H. chrysogaster the colour of the back is black, with an 
admixture of golden-coloured hairs; the belly being of a dark 
golden hue.® 

Xeromys.t—External form Murine. Tip of muzzle as in Mus, 
not as in Hydromys. Toes unwebbed. Tail scaly, very finely 
haired. Skull as in Mus, with the exception of the rounding of the 
supraorbital edges. Teeth as in Hydromys. 

Represented by X. myoides, of Queensland; a species about 
twice the size of the Common Mouse. This genus serves to con- 
nect Hydromys with the other Murines, although it is difficult to 
say to which group it comes nearest. 

Subfamily Plataeanthomyinze.—Molars rooted, with transverse 


1 Sminthus is referred to the Dipodida. 

2 Geoffroy, Ann. du Muséum, vol. vi. p. 81 (1805). 

3 For the anatomy of this animal see B. C. A. Windle, Proc. Zoo/. Soc. 1887, 
p. 53. 4 0. Thomas, Proc. Zool. Soc. 1889, p. 247. 


462 RODENTIA 


lamine. Flattened spines mingled with the hair; tail thickly 
haired. Represented by one genus. 

Platacanthomys..The one representative of this genus is P. 
lasiurus, found in the clefts of rocks and hollow trees in Southern 
India at elevations of about 3000 feet. This elegant little animal 
closely resembles a Dormouse ; the tail and body having a length 
of 6 inches. 

Subfamily Gerbillinze.—Incisors narrow; molars with transverse 
lamine (Fig. 205). Auditory bulle very large in most cases. 
Hind limbs elongated. Tail usually long and hairy. Ranges over 
the Palearctic, Oriental, and Ethiopian regions. 

Gerbillus.2-—Upper incisors grooved; first molar with three 
lamine, second with two, and third with one. 
There are some sixty species, with a range 
coextensive with that of the family. The 
Gerbils, with their large and bright eyes and 
long tufted tails, are very graceful creatures, 
inhabiting sandy plains, where they form ex- 
tensive burrows. Remains of existing species 


are found in cavern-deposits in Madras (Fig. 
Fic. 205.—The left ramus 205). 


of the mandible of Gerbillus ‘ " 3 i 
indicus, with an enlarged Pachyuromys.2—The African genus Pachy- 


view of the molars, froma wromys is distinguished by the very large size 

rei i pe of the auditory bulla, as well as by the short 

Tndica.) and fleshy tail, which is club-shaped. The 
incisors are narrow and faintly grooved. 

Mystromys,* Otomys,® and Dusymys.\—These genera, also from 
South Africa, differ from Gerbillus in the form of the molars, and 
are represented by a few species. 

Malacomys.'—The one known species of this genus is from the 
Gaboon, and is in some respect intermediate between the true 
Gerbils and the Rats. Thus the dentition and feet are those of the 
former, but the long scaly tail resembles that of the latter. 

Subfamily Phleomyins.’—This subfamily is represented only 
by Phicomys® cumingt, of the Philippine Islands, in which the incisors 
are very broad, the molars are divided into transverse lamin, and 
the claws are large. The muzzle is blunt; the ears are hairy 


} Blyth, Proe. ds. Soe. Bengal, vol. xxviii. p. 289 (1859), 
* Desmarest, Nouv. Dict. @ Hist. Nat. vol. xxiv. p. 22 (1804). 
* Lataste, Le Nut. vol. i. p. 314 (1880). 

4 Wagner, Wiegmann’s Archiv, 1841, p. 132. 

5 F. Cuvier, Dents des Manmiferes, p. 168 (1825). 
® Peters, Monatsber. Ak. Berlin, 1875, p. 12. 

" A, Milne-Edwards, Bull. Soc. Philom. ser. 6, vol. xi. p. 9 (1877). 

8 Nesocia was included by Alston in this subfamily. 

® Waterhouse, Proc. Zovl. Soc. 1839, p. 108. 


MURIDAE 463 


externally ; the tail is moderate, and thickly haired; and the 
auditory bulle are very small. The first upper molar has three, 
and the others two lamine. 

Subfamily Dendromyinze.—Incisors convex in front; molars 3, 
rooted and tuberculated. Ears hairy; claws long. Confined to 
the Ethiopian region. 

Dendromys1—A. small Rodent, with the habits of a Dormouse, 
characterised by its grooved incisors, slender form, and long scaly 
tail, which ‘is sparsely haired. Two other Murines described as 
Steatomys? and Lophuromys® are referred to this subfamily. The 
first is of plump form, with a rather short and thickly haired 
tail, and grooved incisors. The latter resembles Steatomys in form, 
but has fine flattened bristles instead of fur, and plain incisors. 

Subfamily Cricetine.—Molars #, tuberculate and rooted, with 
the tubercles of the upper ones arranged in two longitudinal rows 
(Fig. 206, B). This subfamily has an almost 
cosmopolitan distribution, and appears to include 
the most generalised members of the family, from 
which the more specialised J/urinw have been 
evolved. 

Cricetus.sA—According to the arrangement pro- 
posed by Mr. O. Thomas ® this genus is taken to 
include both the Hamsters of the Old World 
(Cricetus proper) and the white-footed or Vesper 
Mice (Hesperomys) of the New. Cheek-pouches 
are frequently present, and may be very large. Fre. 203.—Left upper 
The first molar (Fig. 206, 6) generally has six molars of Mus (1) and 
tubercles. The tail may be very short. ne 

This large and unwieldy genus may be divided into a number of 
groups or subgenera. The typical group includes the Hamsters of - 
the Old World, characterised by the large size of their cheek-pouches, 
the walls of which are connected with muscles arising from the 
lumbar vertebre. The tail is remarkable for its shortness. The 
best-known species is C. frumenturius, inhabiting Europe and Northern 
Asia. The American forms, which range over the whole of that 
continent, comprise a number of subgenera, of which the following 
are the most important. Lhipidomys, including Dormouse-like 
forms with long tails and a dentition like that of the typical 
group; Oryzomys, represented by Murine species; Culoniys, with 
short tail and Hamster-like body ; / esperimus, with only five tuber- 
cles on the first molar; Onychomys, in which the tail is extremely 


1 Andrew Smith, S. African Quart. Journ. vol. ii. p. 158 (1834). 
? Peters, Reise n. Mossambigque, vol. i. p. 162 (1852). 
3 Peters, Monatsber. Ah, Berlin, 1874, p. 234. 
+ Cuvier, Regre Animal, vol. i. p. 198 (1817). 
° Proc. Zool. Soc, 1888, p. 138. 


464 RODENTIA 


short and Hamster-like, and the form is Arvicoline ; Scapteromys, of 
Murine form with a long and hairy tail; Phyllotis, with a shorter 
tail ; Habrothria, an Arvicoline group, with a short and thinly haired 
tail; and Oxymycterus, distinguished from the preceding by having 
a nail instead of a claw on the pollex. With regard to the dis- 
tribution of these forms Mr. Thomas! remarks that in South 
America as we proceed southwards there is a general tendency “to 
a disappearance of the tropical and northern Mouse- and Dormouse- 
like subgenera Rhipidomys, Vesperimus, and Oryzomys, with the 
appearance and increase of the Vole- and Hamster-like Habrothrix 
and Calomys—a change that is curiously paralleled in the Old World 
by the gradual supercession of Mus and Myoxus in favour of Arvicola 
and Cricetus as we go northwards from tropical to temperate and 
arctic regions.” One species has spines in the fur. 

Remains of Cricefus are abundant in the Pleistocene cavern- 
deposits of Brazil, where a number of the forms are referable to 
existing species; the genus is also represented in the Miocene of 
North America and Europe, the species from the former area having 
been described as Eomys, and those from the latter as Cricetodon. 

Holochilus? (Nectomys).—The Rats of this genus are allied to 
the American forms of Cricetus, but have the third upper molars 
proportionately larger and the skull more stoutly built. This 
genus is confined to Brazil, and contains about six species, some of 
which are the largest indigenous Rats of America. Two species are 
aquatic in their habits, and have short webs between the toes of 
their hind feet. 

Sigmodon® differs from Cricetus in the pattern of the molar 
teeth. It contains one species only, the Rice-Rat, S. hispidus, 
ranging from the United States to Ecuador. 

Lhithrodon,* and Ochetodon2—These are more or less like 
Cricetus, but with grooved upper incisors. The first is a South- 
American genus, and contains five Rat-like species, one from 
Venezuela, another from Peru, and the other three from Patagonia. 
The second consists of three North American mice, of about the 
size and proportions of the English Wood-Mouse (Mus sylvaticus). 

Neotoma.6—A peculiar North American genus, in which the 
teeth simulate the prismatic appearance of those of the _4rvicoline. 
There are four species known as Wood-Rats, all of about the size 
of Mus decumanus ; one of them (N. cinerea) having a tail almost as 
bushy as a Squirrel’s while the other three have ordinary scaly 


Baris tale + Proc. Zool. Soc. 1884, p. 451. 
* Brandt, Mém. Acad. Imp. St. Pétersbourg, sér. 8, vol. iii. p. 428 (1835). 
* Say and Ord, Journ. cad. Philad. vol. iv. p. 352 (1825). 
+ Waterhouse, Proc. Zool. Soc. 1837, p. 29. 
° Coues, Proc. Acad. Philad. 1874, p. 184. 
6 Say and Ord, Journ. Acad. Philad vol iv, p. 846 (1825). 


MURIDE 465 


Fossil remains of Neotoma from cavern-deposits in Pennsylvania 
are not improbably referable to the existing Florida Rat (WV. 
flridana). Paciculus, from the Miocene of the United States, is 
regarded as an allied extinct genus with enamel-folds to the molars. 

Hypogeomys1—This and the following genera are confined 
to Madagascar, where they are the sole representatives of the 
Rodentia. Hypogeomys is a very peculiar form of large size, with 
long ears, feet, and tail. There is only one species, H. antimena, a 
fawn-coloured Rat about 9 inches long. 

Nesomys.2—Contains two species of long-haired Rats, more or 
less rufous in colour, about the size of the Brown Rat. 

Brachytarsomys.2—Represented only by B. albicauda, a pretty 
velvety-haired fawn-coloured Rat, with short feet and a long tail. 

Hallomys.t—The only species (H. audeberti) is very like a 
Nesomys, but has much longer hind feet. 

Eliurus.5—Represented by one small Dormouse-like species, 
characterised by its nearly naked and short ears, and long tail, of 
which the proximal third is scaly, and the remainder covered 
with long hair. The pollex is rudimental, but the hallux well 
developed. 

Subfamily Arvieolinze.—Molars usually imperfectly rooted or 
rootless, and composed of two longitudinal rows of triangular 
prisms placed alternately 
(Fig. 207). Tail moderate 
or short. Common to the 
Palearctic and Nearctic 
regions. 

The Voles, as the members 
of this group are commonly 
termed, are so closely con- 
nected with the Cricetines 
that they may be regarded 
merely as a branch of that 
subfamily which has attained 
a peculiarly specialised type 


ai Fic. 207.—Upper (A) and lower (B) molars of the 
of molar dentition. The Water-Vole (Arvicola amphibius). 


Voles are externally dis- 

tinguished, as a rule, from true Rats and Mice by their more 
clumsy and heavy build and less graceful movements ; by the small 
size of their eyes, the bluntness of the muzzle, the small ears, and 
the shorter limbs and tail. 


1 Grandidier, Rev. and Mag. Zool. 1869, p. 388. 
2 Peters, Sitzber. Ges. Nat. Freunde, 1870, p. 54 (1871). 
3 Giinther, Proc. Zool. Soc. 1875, p. 79. 
4 Jentink, Notes Leyd. Mus. vol. i. p. 107, note 27 (1879). 
5 Milne-Edwards, Ann. Sci. Nat. sér. 6, vol. xx. art. 1, bis, p. 1 (1886). 
30 


466 RODENTIA 


Phenacomys—A North American genus distinguished by its 
rooted molars, and thus connecting the typical forms with 
Cricetines like Neotoma. Several species have been described by 
Dr. C. H. Merriam. 

Arvicola2—The type genus Arvicola has rootless molars, and 
naked soles to the feet. It includes over forty species inhabiting 
Europe, North America, and Asia, a few species entering into the 
northern limits of the Oriental region in India. Three species of 
the genus are found in the British Isles, of which the following 
account is given by Mr. O. Thomas :— 

The common Water-Vole (4. amphibius) is as large as the Brown 
Rat. Its fur is long, soft, and thick, of a uniform grizzled brown 
all over, except when, as is not uncommon, it is wholly black. The 
tail is about half the length of its head and body, and the hind feet 
are unusually long and powerful, although not webbed, and have 
five rounded pads on their lower surfaces. Its molar teeth (see 
Fig. 207) present the following number of prismatic spaces :—in 
the upper jaw the first, or anterior, has 5, the second 4, and the 
third 4, of which the last is very irregular in shape, and is 
sometimes itself divided into two, making 5 in all; in the lower 
jaw the first has 7 spaces, of which the 3 anterior are generally not 
fully separated from one another, the second has 5, and the third 
3. These numbers for the different teeth are taken as the 
characters of the subgenus Paludicola of Dr. Blasius, by whom this 
method of subdividing the genus was first introduced. The Water- 
Vole is one of the commonest English mammals, and is perhaps the 
most often actually seen of all, owing to its diurnal habits. It 
frequents rivers and streams, burrowing deeply into their banks, 
and in this way often causing considerable damage. Its food 
consists almost wholly of water-weeds, rushes, and other vegetable 
substances, but, like so many other Rodents, it will also occasionally 
eat animal food, in the shape of insects, mice, or young birds. 
The female during the warm season of the year has three or four 
litters, each of from two to seven young. The range of the 
Water-Vole extends over the whole of Europe and North Asia, 
from England to China, but it is not found in Ireland. The common 
Field-Vole, or short-tailed Field-Mouse (4. agrestis), representing 
the subgenus Agricola, is about the size of a House-Mouse, but 
with a short stumpy body, and a tail only about one third the 
length of the head and body combined. Its hind feet have six 
pads on their inferior surfaces. The colour is dull grizzled brown 
above, and grayish-white below. Its molar teeth have respectively 
5, 5, and 6 prismatic spaces above, and 9, 5, and 3 below. The 


1 Merriam, Fauna of North America, No. 2, p. 28 (1889). 


* Lacépéde, Mém. de ’Institut, vol. ili. p. 495 (1801). Many writers employ 
the earlier name Jicrotus for the true Voles. 


MURIDA 467 


Field-Vole is one of the commonest of our smaller mammals, and 
frequents fields, woods, and gardens in enormous numbers, often 
doing very considerable damage in the latter, owing to its fondness 
for garden produce of all kinds. It is spread over the whole of 
Great Britain from the Hebrides southwards. Abroad its range 
extends from Finland to North Italy and from France and Spain 
to Russia. The Bank-Vole (4. glareolus) resembles in size and 
general appearance the common Field-Vole, but may be dis- 
tinguished by its more or less rusty or rufous-coloured back, its 
larger ears, and the relatively longer tail, which attains to about 
half the length of the head and body. Its molar teeth present 
characters so different from those of all other Voles as to have 
caused it to be regarded as belonging to an entirely distinct genus, 
for which the name of Evotomys has been used. Their chief 
distinction lies in the fact that, unlike those of all other Voles, 
their pulp-cavities close up in adult life, and they form distinct 
roots, more’ resembling those of the ordinary Rats and Mice. 
The enamel-spaces of these teeth number respectively 5, 4, and 
5 above, and 7, 3, and 3 below. The habits of this species are 
in every way similar to those of the Field-Vole. Its range in 
Great Britain extends northwards to Morayshire, beyond which it 
has not yet been observed. It is also found all along the north 
temperate zone from France to China, and is replaced in North 
America by a closely allied animal known as A. gapperi. It is 
probable, however, that both 4. gappert and A. glareolus are only 
southern climatic offshoots of a still more northern species, the 
A. rutilus of Northern Europe, Siberia, and Arctic America. 

Fossil remains of Arvicola are common in European Pleistocene 
deposits, and they have also been obtained from the Upper 
Pliocene of the Norwich Crag. 

Synaptomys..—Represented by one North American species, 
having grooved upper incisors, skull and molars like those of 
Myodes, with the external characters of Arvicola. 

Myodes.2— Distinguished from Arvicola by the more clumsy 
build, convex obtuse head, extremely short and Rabbit-like tail, 
short ears, small feet, the soles of which are furred, elongated claws, 
and thick fur, as well as by the breadth and massiveness of the 
skull, in which the zygomatic arch has a laminar expansion and 
the palate a peculiar contour; while the root of the lower incisor 
does not extend behind the last molar, the upper incisors are 
bevelled, and not grooved, and the molars have a characteristic 
pattern, which cannot be well explained without a figure. 

The Lemmings, as the members of the genus are commonly 
called, are represented by the Norwegian Lemming (J/. lemmus, Fig. 
1 Baird, Mamm. North America, pp. xliv. 558 (1857). 

2 Pallas, Zoogr. Rosso-Asiat. vol. i. p. 173 (1811). 


468 RODENTIA 


208), and the North American M. obensis. Different individuals of 
the Norwegian Lemming vary considerably both in size and colour, 
but its usual length is about 5 inches, and its soft fur yellowish 
brown, marked with spots of dark brown and black. It has a 
short, rounded head, obtuse muzzle, small bead-like eyes, and short 
rounded ears, nearly concealed by the fur. The tail is very short. 
The feet are small, each with five claws, those of the fore feet 
strongest, and fitted for scratching and digging. The usual dwell- 


4 
0 


Fic. 208.—The Lemming (Myodes lemmus). 


ing place of the Lemmings is in the highlands or fells of the great 
central mountain chain of Norway and Sweden, from the southern 
branches of the Langfjeldene in Christiansand-stift to the North 
Cape and the Varangerfjord. South of the Arctic circle they are, 
under ordinary circumstances, exclusively confined to the plateaus 
covered with dwarf birch and juniper above the conifer region, 
though in Tromsé-amt and in Finmarken they occur in all suitable 
localities down to the level of the sea. The nest is formed under a 
tussock of grass or a stone, constructed of short dry straws, and 
usually lined with hair. The number of young in each nest is 
generally five, sometimes only three, but occasionally seven or eight, 
and at least two broods are produced annually. Their food is 


MURIDE 469 


entirely vegetable, especially grass-roots and stalks, shoots of the 
dwarf birch, reindeer-lichens, and mosses, in search of which they 
form, in winter, long galleries through the turf or under the snow. 
They are restless, courageous, and pugnacious little animals. When 
suddenly disturbed, instead of trying to escape they will sit upright, 
with their back against a stone or other coign of vantage, hissing 
and showing fight in a very determined manner (Fig. 208). 

The circumstance which has given more popular interest to the 
Lemming ‘than to a host of other species of the same order of 
animals is that certain districts of the cultivated lands of Norway 
and Sweden, where in ordinary circumstances they are quite un- 
known, are occasionally and at very uncertain intervals, varying 
from five to twenty or more years, literally overrun by an army of 
these little creatures, which steadily and slowly advance, always in 
the same direction, and regardless of all obstacles, swimming across 
streams and even lakes of several miles in breadth, and committing 
considerable devastation on their line of march by the quantity of 
food they consume. In their turn they are pursued and harassed 
by crowds of beasts and birds of prey, as bears, wolves, foxes, dogs, 


-,. Wild cats, stoats, weasels, eagles, hawks, and owls, and never spared 


by man; even the domestic animals not usually predaceous, as 
cattle, goats, and reindeer, are said to join in the destruction, 
stamping them to the ground with their feet, and even eating their 
bodies. Numbers also die from diseases apparently produced from 
overcrowding. None ever return by the course by which they 
came, and the onward march of the survivors never ceases until they 
reach the sea, into which they plunge, and swimming onwards in 
the same direction as before perish in the waves. These extra- 
ordinary and sudden appearances of vast bodies of Lemmings, and 
their singular habit of persistently pursuing the same onward course 
of migration, have given rise to various speculations, from the 
ancient belief of the Norwegian peasants, shared in by Olaus 
Magnus, that they fall down from the clouds, to the almost equally 
untenable hypothesis, ingeniously maintained by the late Mr. W. 
D. Crotch, that they are acting in these migrations in obedience to 
an instinct inherited from vastly ancient times, and are still seeking 
the congenial home in a supposed submerged Atlantis, to which 
their ancestors of the Miocene period were wont to resort when 
driven from their ordinary dwelling-places by crowding or scarcity 
of food. The principal really ascertained facts regarding these 
migrations seem to be as follows. When any combination of cir- 
cumstances has occasioned an increase in the numbers of the 
Lemmings in their ordinary dwelling-places, impelled by the rest- 
less or migratory instinct possessed in a less developed degree by 
so many of their congeners, a movement takes place at the edge of 
the elevated plateau, and a migration towards the lower-lying land 


470 RODENTIA 


begins. The whole body moves forward slowly, always advancing 
in the same general direction in which they originally started, but 
following more or less the course of the great valleys. They only 
travel by night; and, staying in congenial places for considerable 
periods, with unaccustomed abundance of provender, notwith- 
standing all the destructive influences to which they are exposed, 
they multiply excessively during their journey, having families still 
more numerous and more frequently than in their usual homes. 
The progress may last from one to three years, according to the 
route taken, and the distance to be traversed until the sea-coast 
is reached, which in a country so surrounded by water as the 
Scandinavian peninsula must be the ultimate goal of such a journey. 
This may be either the Atlantic or the Gulf of Bothnia, according 
as the migration has commenced from the west or the east side of 
the central elevated plateau. Those that finally perish in the sea, 
committing what appears to be a voluntary suicide, are only acting 
under the same blind impulse which has led them previously to 
cross smaller pieces of water with safety. 

Cuniculus.'—Cranial and incisive characters those of Myodes, 
in the main, but the molars more of an Arvicoline type, the first 
upper one differing from that of all other members of the family in 
having seven prisms. Externally of the general shape of Myodes, 
but distinguished by the absence of external ears, the shortness and 
dense furring of the feet, the obsolete pollex with rudimentary 
nail, and the great length of the two middle claws of the manus. 
Represented by one species, the Banded Lemming (C. torquatus), of 
the Arctic region. 

Remains of both C. torquatus and Myodes lemmus occur in British 
Pleistocene deposits. 

Fiber.?—Closely allied to Arvicola, both externally and in cranial 
and dental characters, but with the tail nearly as long as the body 
(apart from the head), compressed, nearly naked, and reticulate. 
Feet incompletely webbed, and the whole body adapted for a 
thoroughly aquatic life. 

The Musk-Rat or Musquash (F. zibethicus, Fig. 209) is the only 
representative of this genus, and the largest member of the sub- 
family, the head and body being about 12 inches in length. It is 
rather a heavily built animal, with a broad head, no distinct neck, 
and short limbs; the eyes are small, and the ears project very little 
beyond the fur. The fore limbs have four toes and a rudimentary 
thumb, all with claws; the hind limbs are larger, with five distinct 
toes, united by short webs at their bases. The tail is laterally 
compressed, nearly naked, and scaly. The hair much resembles 
that of a beaver, but is shorter; it consists of a thick soft under- 

? Wagler, Isis, 1832, p. 1220. 
° Cuvier, Légons d’ Anatomie Compar. tab. 1 (1800). 


MURIDE 471 


fur, interspered with longer stiff, glistening hairs, which overlie and 
conceal the former on the upper surface and sides of the body. 
The general colour is dark umber-brown, almost black on the back 
and gray below. The tail and naked parts of the feet are black. 
The musky odour from which it derives its name is due to the 
secretion of a large gland situated in the inguinal region, and present 
in both sexes. 

The Musk-Rat is peculiar to America, being extensively distri- 
buted in suitable localities in the northern part of the continent, 
extending from the Atlantic to the Pacific, and from the Rio Grande 


Fia. 209.—The Musk-Rat (Fiber zibethicus.) 


to the barren grounds bordering the Arctic Seas. It is aquatic in 
its habits, living on the shores of lakes and rivers, swimming and 
diving with great facility, feeding on the roots, stems, and leaves of 
water-plants, or on fruits and vegetables which grow near the 
margin of the streams it inhabits. Musk-Rats are most active at 
night, spending the greater part of the day concealed in their 
burrows dug out of the bank, consisting of a chamber with numerous 
passages, all of which open under the surface of the water. For 
winter quarters they build more elaborate houses of conical or 
dome-like form, composed of sedges, grasses, and similar materials 
plastered together with mud. As their fur is an important article 
of commerce, large numbers are annually killed, being either trapped 
or speared at the mouths of their holes. 

The skull of the Musk-Rat is shown in Fig. 203 (p. 459) ; its 
structure is essentially Arvicoline, but the squamosals are greatly 


472 RODENTIA 


expanded, with a corresponding reduction of the parietal and inter- 
parietal, and the interorbital constriction of the frontals attains its 
greatest development. Fossil remains of Fiber occur in the North 
American Pleistocene. 

Neofiber.1—This genus, while agreeing with /vber in the characters 
of the skull and teeth, differs by the cylindrical tail, and the normal 
form of the feet, in which the toes are not bent laterally at an angle 
with the sole. The single species NV. allent, commonly known as 
the Round-tailed Musk-Rat, is found in Florida, and is much less 
completely aquatic in its habits than Fiber. Its colour is brown 
above, and silvery-white mixed with rufous below, the sides of the 
body gradually shading from brown to rufous, the forehead and 
’ the tip of the nose are black, while the tail is rufous mingled with 
black. 

Subfamily Siphneinee. — Includes two genera of Mole-like 
Rodents with an Arvicoline dentition, but with the body thoroughly 


Fic. 210.—Siphneus armandi. (From Milne-Edwards.) 


adapted for a subterranean life, the limbs and tail being very short, 
and the external ears rudimentary. Both are Palearctic. 

Ellobius.2—The Russian £. talpinus, the typical representative 
of the genus, has short claws, and comes nearest to the Arvicoline. 
E. fuscocapilius is from Afghanistan. 

Stphneus.s—This genus (Fig. 210) includes species inhabiting 
Northern and Central Asia, and is characterised by the great length 
of the claws of the manus. Remains of an existing species occur 


1 True, Proc. U.S. Nat. Mus. vol. vii. p. 170 (1884). 
* Fischer, Zoognosia, vol. iii. p. 72 (1814). 
* Brants, Het. Geslact der Muizen, p. 20 (1827). 


MURIDE 473 


in the Pleistocene of the Altai, while an extinct one has been 
described from the Pliocene of North China. 

Subfamily Deomyinse. — Represented only by the under- 
mentioned genus, in which the bituberculate anterior and tricusp- 
idate middle ridge of the first upper molar presents a condition 
intermediate between that obtaining in the ('ricetine and that of 
the Murine. 

Deomys.A—Externally as in aus. Pollex with a narrow nail ; 
hind feet elongate. Infraorbital vacuity of skull triangular, not 
narrowed below. Upper incisors with a pair of minute grooves. 
First upper molar with seven distinct tubercles, of which three are 
placed on the middle ridge, and two on each of the others. One 
species, J. ferrugineus, from the Lower Congo, an animal about the 
size of the Common Mouse. 

Subfamily Murine. — Molars rooted and tuberculated ; those 
of the upper jaw with three longitudinal rows of tubercles (Fig. 
206, 1). 

This group includes the true Rats and Mice, and may be 
regarded as more 
specialised than 
the Cricetina. 
All the members 
of the group 
closely resemble 
one another, and 
are light and 
active, with large 
ears, bright eyes, 
and long and 
sealy tails. Their 
coloration, in 
conformity with 
the fossorial and 
nocturnal habits 
of most of the 
forms, is sombre, 
and their move- 
ments are re- 
markably agile 
and graceful. 


* . Fra, 211.—The Australian Brown-footed Rat (Mus ficsei pes). 
Mus.2—Incisors ‘After: Gould. 


narrow, without 
grooves. Structure of molars as in Fig. 206, 4 (p. 463). Incisive 
foramina of skull long ; coronoid process of mandible well developed. 
1 Q. Thomas, Proe. Zool. Soc. 1888, p. 130. 
2 Linn, Syst. Vat, 12th ed. vol. i, p. 79 (1766). 


474 RODENTIA 


Ears and eyes large; muzzle naked at the extremity. Fur soft, in 
some cases intermingled with spines. Pollex with a short nail in 
place of a claw. No cheek-pouches. Tail long, nearly naked, 
with rings of overlapping scales. Vertebre: C7, D13,L6,S 4, 
C 26-32. 

This genus is the largest in the whole mammalian class, com- 
prising not less than 130 species, ranging over the whole of 
the Old World, with the noteworthy exception of Madagascar. 
On the whole, the species are more numerous in tropical than 
in temperate regions, and very few occur in cold countries. 
Many of the species living in warm climates have flattened spines 
mingled with the fur; these spines being shed in winter, when a 
warmer covering is necessary, and replaced by hair. Five species 
occur in England, which are briefly noticed below; and it may be 
observed that none of the species are much larger than Jf. decumanus 
or smaller than J. minutus. As a rule the habits of the species 
are similar to those of the English forms, but a few are arboreal, 
while others again, like the one represented in Fig. 211, are 
aquatic. The earliest known representatives of the genus (exclud- 
ing -Leanthomys gaudryi of the Lower Pliocene Pikermi beds of Attica) 
occur in the Pleistocene of Europe. 

The Brown or Norway Rat (Jf decumanus) is a heavily built 
animal, growing to § or 9 
inches in length, with a 
bluff rounded head, small 
ears (Fig. 212, 4), and a 
comparatively short tail, 
which is always shorter 
than the head and body 
combined, and generally 
not longer than the body 
alone. The colour is a 
uniform grayish - brown 
above and white below, 
the ears, feet, and tail being 
flesh coloured. Black 
varieties, which are often 
mistaken for true Black 
Rats, are by no means rare, 
but the differences in size 
Fic. 212.—a, Head of Brown Rat (M. decumanus). and proportions form a 

B, Head of Black Rat (Mus rattus). ready means of distinguish- 


; ing the two. The Brown 
Rat is believed to be a native of Western China, where a race 


(Vv. humiliatus) has been discovered so like it as to be practically 
indistinguishable. Both this, and the next species agree in their 


MURIDE 475 


predaceous habits, omnivorous diet, and great fecundity. They 
bear four or five times in the year from four to ten blind and 
naked young, which are in their turn able to breed at an age of 
about six months; the time of gestation being about twenty 
days. 

The Black Rat (Jf rattus) is a smaller and more lightly built 
species, generally not more than 7 inches in length, with a slender 
head (Fig. 212, B), large ears, and a thin tail of about 8 or 9 
inches in length. The colour is usually a glossy bluish-black, some- 
what lighter below; but in the tropical variety described as MM. 
alecandrinus the general colour is gray or rufous, and the belly 
white. The disposition of the Black Rat is milder than that of 
AM. decumanus, and the white and pied rats kept as pets mostly belong 
to this species. In many localities where it was formerly abundant 
it has been entirely superseded by Af: decumanus, but it is said that 
in some parts of Germany it has been lately reasserting itself. 

M. musculus, the Common House-Mouse, is, like the Brown Rat, 
originally a native of Asia, whence it has spread to all the inhabited 
parts of the globe. Its habits and appearance are too well known 
to need any description. 

M. sylvaticus, the Wood or Long-tailed Field-Mouse, is very 
common in many parts of England, often taking to barns and out- 
houses for shelter during the winter. It is of about the same size 
and proportions as J/. musculus, but of a bright reddish-gray colour, 
with a pure white belly. 

M. minutus, the Harvest-Mouse, is the smallest of the European 
Mice, seldom exceeding 24 or 3 inches in length. It is of a 
yellowish-red colour, with comparatively short ears and tail. It 
lives entirely away from human habitations, generally dwelling in 
grass or corn-fields, where it builds a globular nest of dried grass of 
the size of a cricket-ball, in which the young are nurtured. 

Nesocia.1—General characters those of Mus, but the incisors 
and molars very much wider, and the tubercles of the latter more 
connected by transverse ridges, thus producing a laminated type 
of structure. 

This genus has been placed by some writers in a distinct sub- 
family with Phileomys, but Mr. O. Thomas regards it as so closely 
allied to Mus that even its generic separation may be open to 
question. It comprises several species, mostly spread over Southern 
Asia, ranging from Palestine to Formosa, and from Kashmir to 
Ceylon, but N. scudlyi is found in Turkestan. The great Indian 
Bandicoot-Rat (NV. bandicota) is the largest representative of the 
subfamily, often exceeding a foot in length. N. bengalensis is 
remarkable for possessing no less than eighteen mamme. Fossil 
remains of Nesocia occur in the Pleistocene of Madras and in the 


1 Gray, Ann. Mag. Nat. Hist. vol. x. p. 264 (1842). Amended from Nesokia. 


476 RODENTIA 


Pliocene. of Northern India; those from the first-named deposits 
being referable to existing species. 

Golunda.1—Like Mus, but with a distinct groove down the front 
of the upper incisors. There are only three species, one from 
Western India, one from West Africa, and the other frofi Eastern 
Africa. : 

Uromys.2—Differs from Mus in having the scales of thetttdil:nstt 
overlapping, but set edge to edge, so as to form a sort of mosaic 
work. There are about six species of Uromys, spread over the 
northern part of the Australian region from the Aru Islands to 
Queensland. 

Chiruromys.23—Externally like Mus, but with the terminal 
portion of the tail without scales above, quite naked, transversely 
wrinkled, and prehensile. Scales of remainder*oftzilsmore or less 
pentagonal, and arranged in oblique diagonal series. Supraorbital 
vacuity of skull without projecting plate in external wall. In- 
cisive foramina short and narrow; auditory bulla small. Upper 
molars very complex, with the tubercles (of which there are eleven 
in the first tooth) low, and distinctly arranged in transverse rows. 
Known only by C. forbesi, from mountains in New Guinea, which 
must be regarded as a specialised form very similar in outward 
appearance to Uromys cervinipes. 

Hapalotis.*—Hind limbs elongated. Incisive foramina very 
large. No coronoid process to the mandible. This genus is con- 
fined to Australia, where there are about fifteen species known. 
They are pretty little animals, with long ears and tail, and in many 
respects resemble the Jerbogs; whose place they seem to take in 
the sandy Australian deserts. Remains of H. albipes occur in the 
Pleistocene of New South Wales. 

Mastacomys.°—Like Mus, but with the molars remarkably 
broadened, and with only four mamme. The single species of the 
genus is as yet only known from Tasmania, though it has been 
found fossil in New South Wales; it is somewhat similar in size 
and general appearance to the English Water-Vole, but has much 
longer and softer fur. 

Acanthomys.-—Fur almost entirely composed of flattened spines. 
Teeth and skull as in Mus, but the coronoid process of mandible 
very small. There are six species of Spiny-Mice known, all of 
about the size of the Common Mouse. They are found in Syria, 


Gray, Charlesworth’s Mag. Nat. Hist. vol. i. p. 586 (1837). Syn. Pelomys, 
Peters (1852). 

? Peters, Monatsber. Ak. Berlin, 1867, p. 343. 

3 0. Thomas, Proc. Zool. Soc. 1888, p. 237. 

* Lichtenstein, Darst. new. Stéiugethiere, pt. iv. pl. 29 (1829). 

° 0. Thomas, Ann. Mag. Nat. Hist. ser. 5, vol. ix. p. 413 (1882). 

5 Geoffroy, Ann. Set. Nat. sér, 2, vol. x. p. 126 (1840). Acomys. 


SPALACIDE 477 


Palestine, and Eastern Africa as far south as Mozambique. 4. 
dimidiatus presents the appearance of a little Hedgehog when its 
spines are erected; it inhabits the stony deserts of Arabia Petraea 
and Palestine, and feeds on bulbs. A fossil Mouse (4. gaudryi) 
referred to this genus occurs in the Lower Pliocene of Attica. 

Echinothria.1—A very remarkable rat with an extremely elong- 
ated muzzle, all the bones of the face being much produced. The 
incisors are faintly grooved. The only species is £. leucura, an 
animal of about the size of the Brown Rat, with its fur thickly 
mixed with spines. It is found in Celebes.: 

Typhlomys.*—This genus is represented by a single species from 
China, which resembles a House-Mouse in size and general appear- 
ance, but has smaller ears, while the eyes are so reduced in size as 
to be totally concealed by the long eyelashes. 

Cricetomys* and Saccostomus..—These two African genera have 
been—from the presence of cheek-pouches—usually placed in the 
neighbourhood of Cricetus, but their molars are of the Murine type. 
Cricetomys is said to have grooved upper incisors, and is represented 
only by C. gambianus. There are two species of Saccostomus. 

Pithechirus.—A small Rodent from Sumatra and Java described 
under this name is a true Mouse, having nothing to do with 
Chiropodomys, to which it has been compared. 


Family SPALACIDA. 


Mole-like forms, with very small or rudimentary eyes and ear- 
conchs, large claws, and short or/rudimentary tail. Form cylin- 
drical. Incisors large; premolars present or absent; molars rooted, 
with re-entering enamel-folds ; palate narrow. 

Subfamily Spalacine.— Angular part of the mandible arising 
from the lower edge of the socket of the lower incisor. No pre- 
molars. . 

Spalav.°—Represented by the great Mole-Rat (S. typhlus) of 
South-Eastern Europe, in which the eyes are completely covered by 
the skin. 

Ehizomys..—Eyes uncovered, although very minute; small 
naked ear-conchs ; and a short partially hairy tail. Includes 
several species from Northern India, Tibet, China, Burma, Malaya, 
and Eastern Africa. A fossil species occurs in the Pliocene Siwaliks 
of Northern India. 

1 Gray, Proc. Zool. Soc. 1867, p. 599. Amended from Echimys. 
2 Milne-Edwards, Budi. Soc. Philom. sér. 6, vol. xi. p. 9 (1877). 
3 Waterhouse, Proc. Zool. Soc. 1840, p. 2. 
+ Peters, Monatsber. Ak. Berlin, 1846, p. 258. 
5 Giildenstaidt, Nov. Comment. Petrop. vol. xiv. art. i. p. 409 (1770). 
8 Gray, Proc. Zool. Soc. 1830, p. 95. 


478 RODENTIA 


Subfamily Bathyergine.—Angular part of the mandible arising 
from the side of the socket of the lower incisor. Premolars absent 
or present. Confined to the Ethiopian region. 

Bathyergus.1.—Upper incisors strongly grooved; p 4, m 3; no 
ear-conchs ; very powerful claws. One species (B. maritimus), from 
South Africa, attaining a length of about 10 inches. 

Georychus? and Alyoscalops.s—-Upper incisors without grooves. 
Georychus, with some half dozen species, generally has p +; Myo- 
scalops, with one species, usually has p 3, and the second toe of the 
foot is the longest. In Georychus the premolar may be wanting, 
and some examples of Myoscalops have only two teeth of this 
series. 

Heterocephalus.s—Small and nearly naked forms, with small 
head, small eyes, no ear-conchs, moderately long tail, and powerful 


fore feet provided with a pair of large pads; p 2, m =. Two 
species. These very remarkable little Rodents are regarded by 
Mr. O. Thomas as very closely allied to Georychus, but specialised, 
and, so to speak, somewhat degraded for a purely subterranean life, 
for which their hairless body is peculiarly adapted. They are 


found in Somali-land, where they burrow in the sandy soil. 


Family GEOMYID&. 5 


Terrestrial or fossorial forms, with large cheek-pouches opening 
on the cheeks outside the mouth. Squamosal much expanded, 
and the jugal extending forwards to the lachrymal. P21; molars 
rooted or rootless, with transverse lamine. Nearctic and Neo- 
tropical regions. 

Subfamily Geomyine.—lIncisors broad ; mastoid not appearing 
on the top of the skull; eyes small; ear-conch rudimentary ; limbs 
short, subequal. Habits fossorial. 

Geomys.’—U pper incisors deeply grooved. The common North 
American Pouched-Rat or “Pocket-Gopher ” (@. bursarius) inhabits 
the plains of the Mississippi and lives in burrows. Several other 
species are recognised from the Southern United States, Mexico, 
and Central America. The genus is represented in the Pleistocene 
and Pliocene of the United States. 

Thomomys.'—Upper incisors plain. Represented by two species, 


1 Nlliger, Prodromus Syst. Mamm. p. 86 (1811). 

® Tlliger,, Zoe. cit. p. 87. 

2 0. Thomas, Proc. Zool. Soc. 1890, p. 448 = Heliophobius ; Peters, Monatsber. 
Ak. Berlin, 1846, p, 243.—Preoceupied. 

* Riippel, Mus, Senkend. vol. i. Siugeth. p. 99 (1834). 

5 Including the Saccomyide of Coues. 

® Rafinesque, Amer. Monthly Mag. vol. ii. p. 45 (1817). 

7 Wied, Nova Acta Ac. Ces. Leop.-Car. vol. xix. pt. i, p. 383 (1839). 


DIPODIDE 479 


with numerous varieties found all over Canada and North America 
west of the Rocky Mountains. Remains referred to an existing 
species occur in the Pliocene of Oregon.  Lntoptychus, from the 
Miocene of the United States, is an allied genus, with broad incisors 
and rootless molars. 

Subfamily Heteromyine.—Incisors narrow; mastoid appearing 
largely on the top of the skull; eyes and ears moderate or large ; 
hind limbs and tail elongated. Habits terrestrial. 

Dipodomys.1This genus is characterised by the rootless molars. 
It is best known by D. phillipsi, the Kangaroo-Rat of the desert 
regions east of the Rocky Mountains, having habits like those of 
the Jerboas. The typical forms have four toes in the pes; but in 
others, which it has been proposed to separate as Dipodops, there 
are five: D. ordi and D. agilis belong to the latter group. 

Perognathus? and Heteromys.2—In both these genera, which are 
represented by species of very small size, the molars are rooted ; 
the latter being distinguished by the presence of flattened spines 
mingled with the fur, and having species ranging into South 
America. According to Dr. C. H. Merriam the forms described as 
Cricetodipus are not separable from Perognathus ; while Dr. Coues 
considers that Saccomys was founded upon a species of Heteromys. 
Pleurolichus, from the Miocene of the United States, is regarded as 
an extinct genus allied to Heteromys. 


Family DIpopID&. 


Terrestrial forms usually with four upper cheek-teeth, and typi- 
cally with the following characters. Incisors compressed; molars 
with transverse enamel-folds ; infraorbital vacuity of skull (Fig. 7, 
p. 37) large and rounded ; jugal ascending in front to the lachry- 
mal; and the mastoid part of the auditory bulla usually very large. 

Subfamily Sminthinee.—Molars rooted ; » 3, m 3. Skull with 
the infraorbital vacuity widest below, and the incisive palatal 
foramina long. Limbs short. Paleearctic. 

Sminthus.s— Represented by the Rat-like S. vagans from Northern 
Europe and Asia, in which the ears are rather long and pointed, the 
tail is covered with short hairs and nearly as long as the body, 
while the molars present a somewhat complicated pattern. This 
genus has generally been regarded as an aberrant member of the 
Muride, but was transferred in 1887 to the present family by 
Dr. H. Winge. 


1 Gray, dan. Mag. Nat. Hist. vol. vii. p. 521 (1840). 
2 Wied, Nova Acta Ac. Ces, Leop.-Car, vol. xix. pt. i. p. 869 (1839). 
3 Desmarest, Mammalogie, p. 318 (1820). 
+ Keyserling und Blasius, Wirbelthiere Ewrop. p. 38 (1840). 


480 RODENTIA 


Subfamily Zapodinze.—Molars rooted ; p 1, m 2; cervical ver- 
tebre free ; hind limbs elongated ; metatarsals separate ; hind feet 
with five digits. Nearctic region. 

Zapus.i—The American Jumping-Mouse (Z. hudsonianus) ex- 
tends over almost the whole North-American continent from Labra- 
dor to Mexico. 

Subfamily Dipodinze.— Molars rooted ; p oxy m %; cervical 
vertebre more or less ankylosed; hind limbs elongated; metatarsals 
united ; hind feet with only three functional digits. Palearctic 
and Ethiopian regions. 

This subfamily includes the true Jerboas, and contains three 
genera: Dipus? with three toes, and Alactaga* and Platycercomys * 
with five, the outer two not reaching to the ground. The latter is 
distinguished by the absence of premolars, and comprises many 
species extending from Siberia to Nubia. 

Remains of the existing Alactaga decumana® occur in the Pleisto- 
cene of Germany, and those of Zapus hudsonianus in the corresponding 
strata of the United States. Platycercomys has been recorded from 
the Pleistocene of Northern Asia. 

Subfamily Pedetinze.—Molars rootless ; cervical vertebre free ; 
hind limbs elongated; metatarsals separate ; hind feet with four 
digits. Vertebre: C7, D12,L7,83, C30. Ethiopian region. 

Pedetes,® the Cape Jumping-Hare (P. caffer), by far the largest 
species of the family, extends from Mozambique and Angola to the 
Cape of Good Hope. 


Section HYSTRICOMORPHA. 


Skull (Fig. 213) with a stout zygomatic arch; jugal not sup- 
ported below by a continuation of the maxillary zygomatic process ; 
infraorbital vacuity large ; mandible with the angular part arising 
from the outer side of the bony socket of the lower incisor. 
Clavicles perfect or imperfect ; fibula distinct. One premolar in 
each jaw. 


Family OcTODONTIDE. 


Clavicles complete. Skull with long incisive foramina extend- 
ing into the maxille ; and usually an inferior angle to the Jugal. 
Molars with external and internal enamel-folds; p 3, except in 
Ctenodactylus. Mamme placed high up on the sides of the body. 
Confined to the Ethiopian and Neotropical regions, with the excep- 


1 Coues, Bull. U.S. Geol. Surv. Terrs. ser. 2, No. 5, p. 253 (1873). Syn. 
Jaculus, Wagler. 2 Gmelin, Syst. Nat., vol. i. p. 157 (1788). 

3 F. Cuvier, Proc. Zool. Soc. 1836, p. 141. 

* Brandt, Bull. Ac. St. Pétersbouwrg, 1844, p. 209. 

5 = A, jaculus, Auct. _ © Mliger, Prodromus Syst. Mamm. p. 81 (1811). 


OCTODONTIDZ 481 


tion of one species of Echinomys which ranges into Central America. 
Habits mostly terrestrial, but occasionally fossorial or natatorial. 

Subfamily Ctenodactylines. —NMolars semi-rooted ; jugal as in 
Dipodide ; the two inner toes of the hind feet with a horny comb 
and rigid bristles. Ethiopian region. 

Ctenodactylus.1i— Represented only by C. gundi from North 
Africa, on the borders of the Sahara. Has no premolars ; each foot 
has four digits; the hind limbs are rather longer than the fore; the 
ears small; and the tail reduced toa stump. This animal is about 
the size of the Water-Vole, and dwells on rocky ground, its habits 


Fic, 213.—Skull of Hydrocherus capybara (reduced). 


being diurnal. The peculiar comb-like inner toes are employed for 
dressing the fur. 

Pectinator.2—Closely allied to the preceding, but with a minute 
premolar in each jaw ; and a moderately long and bushy tail. One 
species (P. speket), from Somali-land. 

Subfamily Octodontinze.—Molars semi-rooted or rootless, with 
simple enamel-folds ; fur soft. There are some six existing genera, 
including Rat-like species, all of which are South American, except 
Petromys, which is Ethiopian. 

Octodon.2—Upper and lower molars alike ears moderate ; tail 
of medium length and tufted. Vertebre: C7, D12,L7, 84, C 
25. Typically represented by C. cumingi of Chili and Peru, with 
other species from Chili and Bolivia.’ They live in large com- 
munities. 

1 Gray, Spicilegia Zoologica, p. 10 (1830). 
2 Blyth, Journ. As. Soc. Bengal, vol. xxxiv. p. 294 (1855). 
3 Bennett, Proc. Zool. Soc. 1832, p. 46. 
31 


482 RODENTIA 


Habrocoma..—Lower molars more complex than the upper ; 
ears large ; and fur extremely soft. Two Bolivian species. 

Schizodon.*— One species, inhabiting elevated spots in the 
Southern Andes, and characterised by the enamel-folds of the upper 
molars meeting in the middle line. The external characters are 
much the same as in Ctenomys, but the ears are larger and the claws 
shorter. 

Ctenomys.?—Incisors broad ; molars rootless, with kidney-shaped 
crowns; last molar small and cylindrical; eyes and ears very 
small; claws larger than the toes. Some four species. Fossil 
remains are common in the Pleistocene of Buenos Ayres and the 
cavern-deposits of Brazil. Habits fossorial. 

Spalacopus.*—Represented by two Chilian species, distinguished 
from the preceding genus by the rudimentary ears. These rodents 
store up magazines of food in their burrows. 

Petromys..—The South African P. typicus is closely allied to 
Spalacopus, but differs by its harsh fur, the shortness of the pollex, 
and the somewhat bushy tail. The teeth are semi-rooted, with 
single inner and outer enamel-folds, nearly meeting in the middle. 

Subfamily Eehinomyinze.—Molars semi-rooted or rootless, with 
deep and curved enamel-folds ; fur more or less harsh, frequently 
mixed with spines; tail generally long. One Ethiopian genus, and 
the remaining nine or so Neotropical. Many of the species are 
of large size, some being arboreal and others aquatic. _ 

Myopotamus.6—Incisors very large; molars with two internal 
and two external enamel-folds in the upper, and three internal and 
one external in the lower jaw, last molar the largest ; ears moder- 
ate ; tail about two-thirds the length of the head and body, scaly, 
and sparsely haired; hind feet webbed; five digits. Vertebre : 
C7,D13,L6,8 4,C 25. The well-known Coypu (JZ coypu), the 
only existing representative of this genus, is one of the largest 
living members of the order, and attains a length of about 2 feet. 
It is common in South America, living in burrows near water, and 
feeding on aquatic plants. Fossil remains of the genus occur in the 
caverns of Brazil, as well as in the Tertiaries of Argentina. 

Capromys.'—This genus comprises arboreal forms from the West 
Indies allied to the Coypu, but, according to Dr. G. E. Dobson, 
showing signs of affinity with the Hystricide. The incisors are 
smaller than in the Coypu, and the upper molars hare one internal 


1 Waterhouse, Proc. Zool. Soc. 1837, p. 80. Amended from dracon, 
* Waterhouse, Proc. Zool. Soc. 1841, p. 91. 
° De Blainville, Bull. Soc. Philom. 1826, p. 62. 
+ Wagler, ibid. p. 1219. 
5 Andrew Smith, 8S. African Quart. Journ. vol. ii. p. 2 (1831). 
® Geoffroy, Ann. du Muséum, vol. vi. p. 81 (1805). 
7 Desmarest, J/ém. Soc. d’ Hist. Nat. vol. i. p. 44 (1822). 


OCTODONTID.E 483 


and two external enamel-folds; the ears are comparatively small ; 
the tail usually of considerable length, and the general form some- 
what Rat-like. The typical C. pilorides is somewhat smaller than 
the Coypu, and is confined to Cuba; it is remarkable for the 
sub-division of the lobes of the liver into a number of lobules. 
C. brachyurus and C. prehensilis are also confined to Cuba. In 
Jamaica the genus is represented by C. melanurus, which is somewhat 
smaller than a Rabbit, and has no secondary lobulation of the liver.! 

Aulacodus.°-—Upper incisors with three deep grooves; molars 
as in Capromys. Fur very harsh; tail moderate, sparsely haired ; 
manus with rudimentary pollex, and small fifth digit; pes with no 
hallux, and rudimental fifth digit. One species (4. swinderianus), 
from Western and Southern Africa, which attains a length of nearly 
2 feet, and dwells in burrows. 

Plagiodon.2—Allied to Capromys, but with the enamel-folds of 
the molars very complex, and forming a kind of zig-zag pattern in 
those of the upper jaw. Represented only by P. edium of Hayti 
and Jamaica. 

Loncheres* and Echinomys.6—These genera include small South 
American species, in most of which flattened lanceolate spikes are 
mingled with the fur. The majority of the species occur in Guiana 
and Brazil, but one species of Hchinomys has been recorded from 
Central America. Fossil remains of both genera occur in the 
cavern-deposits of Brazil. 

Mesomys..—This genus resembles Loncheres externally, but the 
pollex has a short curved claw, and there are no spines in the fur. 

Dactylomys..—A Brazilian genus presenting the following dis- 
tinctive features. Ears short; tail long and scaly ; pollex minute ; 
third and fourth digits of manus elongated, with short convex nails. 
Incisors flat; molars divided into two lobes, each of which has 
a single enamel-fold. Represented by two species, D. typus and 
D. amblyonyx, both of which seem to be rare and but little known. 
In the elongation of some of the digits Dactylomys recalls Chiromys 
among the Primates. 

Cercomys..—This South American genus is usually placed near 
Carterodon, from which it is readily distinguished by the pointed 
muzzle and the plain incisors. 


1 For description and anatomy of this species see Dobson, Proc. Zool. Sov. 
1884, p. 233. 

2 Temminck, Monographies des Mammiferes, vol. i. p. 245 (1827). 

3 Cuvier, Ann. Se’. Nat, sér, 2, vol. vi. p. 847 (1836). Amended. 

4 Tlliger, Prodromus Syst. Mamm. p. 90 (1811). 

5 Desmarest, Nouv. Dict. d’ Hist. Nat. vol. x. p. 45 (1817). Amended from 
Echimys. 8 Wagner, Wiegmann’s Archiv, 1845, pt. 2, p. 145. 

7 Geoffroy, dun. Sei. Nat. sér. 2, vol. x. p. 126 (1838). 

8 F. Cuvier, Iammiferes, 6ine livr. (1829). 


484 RODENTIA 


Carterodon1—This genus, which was originally described upon 
the evidence of skulls from the Brazil caves, but subsequently found 
living, is readily distinguished by the broad and grooved incisors. 
The upper molars have one inner and two outer enamel-folds ; 
those of the lower jaw being the reverse of this. 

Fossil Forms.—Remains of the existing genus Loncheres occur in 
the Brazilian cave-deposits, which also yield the extinct Dicolpomys. 
A large number of fossil Octodontide from the Tertiaries of South 
America have been described under many generic names, but it 
will be sufficient to mention that Phloramys and Pithanotomys are 
considered to be allied to Ctenomys ; while Jlvrenia, Orthomys, and 
Trilodon show affinity to Myopotamus. Pellegrinia, from the Pleisto- 
cene of Sicily, may be allied both to Ctenodactylus and Octodon. 


Family THERIDOMYIDA. 


This extinct family, which is represented in the Tertiaries of 
Europe and the United States, comprises several genera of com- 
paratively small Rodents, which are regarded by Dr. Schlosser as 
nearly related to the Octodontide, although connected by Archwomys 
with the Chinchillide. The dental formula is the same as in the 
Octodontide. In the typical genus Theridomys, from the Lower 
Miocene and Upper Eocene of Europe, the molars are rooted, and 
have three or four re-entering enamel-folds, which form isolated 
discs on the worn crowns. Syllophodus, from the Miocene of the 
United States, is closely allied. Protechinomys and Trechomys are 
genera from the Phosphorites of Central France with rooted molars ; 
while in dArchwomys of the same deposits the molars are rootless, 


with the enamel-folds dividing their crowns into lamine, as in the 
Chinchillas. 


Family HystrRicip 2. 


Build stout. Limbs subequal. A number of long and stout 
spines in the integument. Facial portion of skull short and broad, 
and the jugal without an inferior angle. Molars with external and 
internal enamel-folds ; completely or partly rooted. 

Subfamily Synetherinze.—Molars rooted ; clavicles complete ; 
upper lip not cleft; soles tuberculated; pollex absent; four mamme ; 
tail generally prehensile; spines mixed with long hairs. This group 
is confined to America, all the forms except one being arboreal, 
and their habits less strictly nocturnal than in the next subfamily. 
There are three genera. 

Erethizon.2—Represented by the common Canadian Porcupine 


1 Waterhouse, Nat, Hist. of Mami. vol. ii, p- 351 (1848). 
> 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 <Arctoidea. This division is founded 
mainly on characters exhibited by the base of the skull, but is 
corroborated by the structure of other parts.1 The presence or 
absence of a bridge of bone, covering the external carotid artery in 
a part of its course by the side of the alisphenoid bone, and enclosing 
the “alisphenoid canal” (see Fig. 8, p. 38), a character to which the 
late Mr. H. N. Turner first drew attention, might seem unimportant 
at first sight, but it is curiously constant in certain groups, which 
we have other reasons, derived often from a combination of less 


1 See Flower, ‘On the Value of the Characters of the Base of the Cranium 
in the Classification of the Order Curnivora,” Prov. Zool. Soc. 1869, p. 4; Mivart, 
“On the Classification and Distribution of the Elurcidea,” ibid. 1882, pp. 135 
and 459; see also The Cat, an Introduction to the Study of Backboned Animals, 
especially Mammals, vy the same author, 1881. 


-ELUROIDEA 501 


easily definable characters, to regard as natural. It is therefore 
generally mentioned in the following family definitions. 

It must, however, be stated that while the arrangement is a 
convenient one as regards the existing Carnivores, it will not hold 
good when the fossil forms are included. Thus there is ample 
evidence to show that the Dogs and Bears were formerly so inti- 
mately connected that in a paleontological classification the Canide 
cannot be satisfactorily separated from the Urside ; while in another 
direction the Cunide were closely allied to the ancestral Viverride. 
The most important objection against this classification is, however, 
the apparent intimate connection exhibited by fossil forms between 
the Viverride and the JJustelide, which, so far as the present evi- 
dence goes, tends to show that the latter are derived from the 
former. If this be eventually fully proved, it would seem to 
indicate that the Arctoidea are not a natural group; and that the 
resemblances between the Urside and Justelide have been independ- 
ently acquired, in the course of the descent of the one family from 
a Canoid, and of the other from a Viverroid stock. 


Section JELUROIDEA. 


The Atluroidea or Cat-like Carnivores include the Felide, 
Fiverride, Proteleide,and Hyenide. 
The existing representatives of 
this section present the following 
common features. Auditory bulla 
(Fig. 222) much dilated, rounded 
smooth, thin-walled, and (except 
in the Hyenide) divided into two 
chambers by a septum. Bony 
auditory meatus short.  Par- 
occipital process applied to, and 
spread over the hinder part of 
the bulla (Fig. 222). Mastoid 
process never very salient, and 
often obsolete. Carotid canal 
(Fig. 8, p. 38, car) small, some- 
times very inconspicuous. Con- 
dyloid and glenoid foramina con- 
cealed or wanting. Czcum small, 


rarely absent. Os penis generally 
small and irregular (large in 
Cryptoprocta). | Cowper’s glands 
present ; prostate distinctly lobed. 
Some details of the anatomy of 


Fig. 222.—Lett side of the palatal aspect of 
the cranium and mandible of the Suricate (Suri- 
cata tetradactyla). c, Carotid foramen ; J, fissure 
in floor of auditory meatus. From Mivart, 
Proc. Zool. Soc. 1882, p. 184, 


the soft parts will be found under the head of Genefta. 


502 CARNIVORA 


Family FELIDZ. 


In all the forms, both recent and fossil, which can be included 
in this family the canines are strongly developed, there are never 
more than one upper and two lower molars, and the three lower 
incisors are placed in the same horizontal line. With one exception, 
the humerus has an entepicondylar foramen. 

The following characters are common to all the existing 
members. True molars reduced to one above and below, that of 
the upper jaw very small and transversely extended. Only two 
inferior premolars. Upper carnassial with three lobes to the 
blade ; lower without talon or inner cusp. Auditory bulla not ex- 
ternally constricted. No alisphenoid canal. Carotid canal very 
minute. Digits 5-4. Dorsal vertebre 13. 

Felis.1—The whole structure of the animals of this genus ex- 
hibits the Carnivorous type in its fullest perfection. Dentition: 
i 3,¢4, p 3, m+; total 30. A distinctly cusped inner tubercle 
to the upper car- 
nassial. Claws com- 
pletely —_retractile. 
The upper anterior 
premolar (p. 2), al- 
ways small, and may 
be absent without 
any other modifica- 
tion in the dental 
or other structures. 
Such a variation 
should not therefore 
be considered as 
of generic import- 
ance. Incisors very 
small. Canines 
; large, strong, slightly 
recurved, with trenchant edges and sharp points, and placed wide 
apart (Fig. 223). Premolars compressed and sharp pointed. The 
most posterior in the upper jaw (the carnassial), a very large tooth, 
consisting of a sub-compressed blade, divided into three unequal 
lobes supported by two roots, with a very small inner tubercle 
placed near the front end of the tooth and supported by a distinct 
root (Fig. 220). The upper true molar a very small tubercular 
tooth placed more or less transversely at the inner side of the 
hinder end of the last. In the lower jaw the true molar (carnassial) 
reduced to the blade alone, which is very large, trenchant, and 


Fic, 223.—Frout view of skull of Lion (Felis leo). 


? Linn, Syst. Nat. 12th ed. vol. i. p. 60 (1766). 


“FELIDA 503 


much compressed, divided into two subequal lobes. Occasionally 
it has a rudimentary talon, but never an inner cusp. The skull 
is generally short and rounded, though proportionally more elon- 
gated in the larger forms. The facial portion is especially short 
and broad, and the zygomatic arches are very wide and strong. 
The auditory bulle are large, rounded, and smooth. Vertebre: 
C 7, D 13,L 7, S 3, C 13-29.  Clavicles better developed 
than in other Carnivora, but not articulating with either the 
scapule or sternum. Limbs digitigrade. Anterior feet with 
five toes, the third and fourth nearly equal and longest, the 
second slightly and the fifth considerably shorter; the pollex 
still shorter, not reaching as far as the metacarpo-phalangeal 
articulation of the second. Hind feet with only four toes. The 
third and fourth the longest, the second and fifth somewhat shorter 
and nearly equal ; the hallux represented only by the rudimentary 
metatarsal bone. The claws all very large, strongly curved, com- 
pressed, very sharp, and exhibiting the retractile condition in the 
highest degree. The tail varies greatly in length, being in some a 
mere stump, in others nearly as long as the body. Ears of moderate 
size, more or less triangular and pointed. Eyes rather large. Iris 
very mobile, and with a pupillary aperture which contracts under 
the influence of light in some species to a narrow vertical slit, in 
others to an oval, and in some to a circular aperture. Tongue 
thickly covered with sharp-pointed, recurved horny papille. Czecum 
small and simple. 

As in structure so in habits, the Cats may be considered the 
most specialised of all the Carnivora. All the known members of 
the genus feed, in the natural state, almost exclusively on warm- 
‘blooded animals which they have themselves killed. One Indian 
species (F. viverrina) preys on fish and even (it is said) on freshwater 
molluscs. Unlike the Dogs, they never associate in packs, and 
rarely hunt their prey in open ground, but from some place of con- 
cealment wait until the unsuspecting victim comes within reach, or 
with noiseless and stealthy tread, crouching close to the ground for 
concealment, approach near enough to make the fatal spring. In 
this manner they frequently attack and kill animals considerably 
exceeding their own size. They are mostly nocturnal, and the 
greater number, especially the smaller species, more or less arboreal. 
None are aquatic, and all take to the water with reluctance, though 
some may habitually haunt the banks of rivers or pools, because 
they more easily obtain their prey in such situations. 

The numerous species of the genus are very widely diffused over 
the greater part of the habitable world, though most abundant in 
the warm latitudes of both hemispheres. No species are, however, 
found in the Australian region, or in Madagascar. Although the Old- 
World and New-World Cats (except perhaps the Northern Lynx) 


504 CARNIVORA 


are all specifically distinct, no common structural character has been 
pointed out by which the former can be separated from the latter. 
On the contrary, most of the minor groups into which the genus 
has been divided have representatives in both hemispheres. 

Notwithstanding the considerable diversity in external appear- 
ance and size between different members of this extensive genus, 
the structural differences are but slight, and so variously combined 
in different species that the numerous attempts hitherto made to 
subdivide it are all unsatisfactory and artificial, The principal 
differences are to be found in the form of the cranium, especially 
of the nasal and adjoining bones, the completeness of the bony orbit 
posteriorly, the development of the first upper premolar and of the 
inner tubercle of the upper carnassial, the length of the tail, the form 
of the pupil, and the condition and coloration of the fur, especially 
the presence or absence of tufts or pencils of hair on the external 
ears. Writing in 1881 Professor Mivart+ gave the number of 
existing species of Felis as 48, but by Mr. Blanford’s reduction of 
the number of Indian species? the list may now be diminished to 
some 41. The following account is chiefly devoted to some of the 
more important and better known species. 

A. Old World Species—The Lion (Ff. leo, Fig. 224) has been 
well known to man from the earliest historic times. Its geographi- 
cal habitat made it familiar to all the races among whom human 
civilisation took its origin, and its strongly marked physical and 
moral characteristics have rendered it proverbial, perhaps to an 
exaggerated degree, and have in all ages afforded favourite types 
for poetry, art, and heraldry. The literature of the ancient Hebrews — 
abounds in allusions to the Lion ; and the almost incredible numbers 
that are stated to have been provided for exhibition and destruction 
in the Roman amphitheatres (as many as six hundred on a single 
occasion by Pompey, for example) show how abundant these 
animals must have been within accessible distance of the capital of 
the world. 

The geographical range of the Lion was once far more extensive 
than at present, even within the historic period covering the whole 
of Africa, the south of Asia, including Syria, Arabia, Asia Minor, 
Persia, and the greater part of Northern and Central India, and also 
the south-eastern portion of Europe, as shown by the well-known 
story told by Herodotus of the attacks by Lions on the Camels which 
carried the baggage of the army of Xerxes on its march through 
the country of the Ponians in Macedonia. The very circum- 
stantial account of that historian shows that the animal in his time 
ranged through the country south of the Balkans, through Rou- 
mania to the west of the River Carasu, and through Thessaly as far 

1 The Cat, pp. 392-426 (1881). 
* Fauna of British India, ‘ Mammalia,” pp. 56-90 (1888). 


PEEIDE 505 


south as the Gulf of Lepanto and the Isthmus of Corinth, having 
as its western boundary the River Potamo and the Pindus mountains. 
The whole of the evidence relating to the existence of Lions in 
Europe, and to their retreat from that continent shortly before the 
commencement of the Christian era, has been collected in the article 

1 “Felis spelea” in Boyd Dawkins and Sandford’s British Pleisto- 
cené Mammalia (1868). Fossil remains attest a still wider range, as 
it is shown in the same work that there is absolutely no osteologi- 


Fig. 224.—Lion and Lioness, after a drawing by Wolf in Elliot's Monograph of the Frida. 


cal or dental character by which the well-known Cave Lion (F. 
xprlea), so abundantly found in cave-deposits of the Pleistocene age 
in Western Europe, can be distinguished from the existing F. ko. 
At the present day the Lion is found in localities suitable to its 
habits, and where not exterminated (as it probably was in Europe) 
by the encroachments of man, throughout Africa from Algeria to 
the Cape Colony, and in Mesopotamia, Persia, and some parts of 
the north-west of India. According to Blanford,} Lions are still 
very numerous in the reedy swamps bordering the Tigris and 
Euphrates, and also occur on the west flanks of the Zagros moun- 
tains and the oak-clad ranges near Shiraz, to which they are 
1 Zoology and Geology or Eastern Persia (1878. 


506 ; CARNIVORA 


attracted by the immense herds of swine which feed on the acorns. 
The Lion nowhere exists in the table-land of Persia, nor is it found 
in Baluchistan. In India, where it is verging on extinction, it 
appears now to be confined to parts of Kattywar and Rajputana, 
though within the present century its range extended through the 
north-west part of India, from Bahawalpur and Sind to at 
least the Jumna (about Delhi), southward as far as Khandesh, and 
in Central India through the Saugor and Narbada territories, 
Bundelkund, and as far east as Palamau. It was extirpated in 
Harriana about 1824. One was killed at Rhyli, in the Dumaoh 
district, Saugor and Narbada territories, so late as in the cold 
season of 1847-48 ; and one was shot in 1810 near Kot-Deji, Sind.! 

The great variations in external characters which different Lions 
present, especially in the colour and the amount of mane, has given 
rise to the idea that there are several species, or at all events dis- 
tinct varieties peculiar to different localities. It was at one time 
supposed, on the authority of Captain Walter Smee, that the Lion 
of Gujerat differed essentially from that of Africa in the absence of 
a mane, but subsequent evidence has not supported this view, which 
was probably founded upon young specimens having been mistaken 
for adults. Lions from that district as well as from Babylonia, 
which have lived in the gardens of the London Zoological Society, 
have had as fully developed manes as any other of the species. 
Mr. F. C. Selous* has shown that in South Africa the so-called 
Black-maned Lion and others with yellow scanty manes are found, 
not only in the same locality, but even among individuals of the 
same parentage. 

The Lion belongs to a well-defined group, containing the largest 
members of the genus, and differing from the others in the well- 
marked character that the anterior cornu of the hyoid arch is but 
little ossified, so that this arch is connected with the cranium by a 
long ligament, instead of by a continuous chain of bones, and by 
the less important one that the pupil of the eye, when contracted, 
is a circular hole, instead of a vertical slit as in the cat. The Lion 
agrees with the Tiger and the Leopard in these respects, but differs 
from them in its uniform style of colouring, and from all the other 
Felide in the arrangement of its hairy covering ; thus the hair of the 
top of the head, chin, and neck, as far back as the shoulder, is not 
only very much longer, but also differently disposed from the hair 
elsewhere, being erect or directed forwards, and so constituting the 
characteristic ornament called the mane. There is also a tuft of 
elongated hairs at the end of the tail, one upon each elbow, and 
in most lions a copious fringe along the middle line of the under 

1 See Blanford, Fawne of British India, “Mammalia,” p. 57 (1888). 
2 Transactions of the Zoological Society, vol. i. p. 165 (1835). 
8 A Hunter's Wanderings in Africa, 1881, p. 258. 


FELIDA 597 


surface of the body, wanting, however, in some examples.! It must, 
however, be observed that these characters are peculiar to the adults 
of the male sex only, and that young lions show indications of 
the darker stripes and mottlings so characteristic of the greater 
number of the members of the genus. 

The usual colour of the adult is yellowish-brown, but it may 
vary from a deep red or chestnut brown to an almost silver gray. 
The mane, as well as the long hair of the other parts of the body, 
sometimes scarcely differs from the general colour, but it is usually 
darker and not unfrequently nearly black. The mane begins to 
grow when the animal is about three years old, and is fully de- 
veloped at five or six. 

In size the Lion is only equalled or exceeded by the Tiger 
among the existing Felidw;, though both species present great 
variations, the largest specimens of the latter appear to surpass the 
largest Lions. A full-sized South African Lion, according to Selous, 
measures slightly less than 10 feet from nose to tip of tail, follow- 
ing the curves of the body. Harris gives 10 feet 6 inches, of which 
the tail occupies 3 feet. The Lioness is about a foot less. The 
tongue, like that of the other species of the genus, is long and flat, 
and remarkable for the development of the papille of the anterior 
part of the dorsal surface, which (except near the edge) are modified 
so as to resemble long, compressed, recurved, horny spines or claws; 
these, near the middle line, attaining the length of one-fifth of an 
inch. They give the part of the tongue on which they occur the 
appearance and feel of a coarse rasp, and serve the purpose of such 
an instrument in cleaning the flesh from the bones of the animals 
on which the Lions feed. 

The habits of the Lion in a state of nature are fairly. well known 
from the united observations of numerous travellers and sportsmen 
who have explored those districts of the African continent in which 
it is still common. It lives chiefly in sandy plains and rocky places 
interspersed with dense thorn-thickets, or frequents the low bushes 
and tall rank grass and reeds that grow along the sides of streams 
and near the springs where it es in wait for the larger herbivorous 
animals on which it feeds. Although it is occasionally seen abroad 
during the day, especially in wild and desolate regions, where it is 
subject to but little molestation, the night is, as in the case of so 
many other predaceous animals, the period of its greatest activity. 
It is then that its characteristic roar is chiefly heard, as thus graphi- 
cally described by Gordon Cumming :— 


1 Mr. Selous, whose opportunities for obtaining evidence upon this subject 
were very large, says that in the region of South Africa, between the Zambesi 
and the Limpopo rivers, he never saw a lion with any long hair under the body, 
and that the manes of the wild lions of that district are far inferior in develop- 
ment to those commonly seen in menageries in Europe. 


508 CARNIVORA 


“One of the most striking things connected with the Lion is 
his voice, which is extremely grand and peculiarly striking. It 
consists at times of a low, deep moaning, repeated five or six times, 
ending in faintly audible sighs ; at other times he startles the forest 
with loud, deep-toned, solemn roars, repeated in quick succession, 
each increasing in loudness to the third or fourth, when his voice 
dies away in five or six low muffled sounds very much resembling 
distant thunder. At times, and not unfrequently, a troop may be 
heard roaring in concert, one assuming the lead, and two, three, or 
four more regularly taking up their parts, like persons singing a 
catch. Like our Scottish stags at the rutting season, they roar 
loudest in cold frosty nights; but on no occasions are their voices 
to be heard in such perfection, or so intensely powerful, as when 
two or three troops of strange Lions approach a fountain to drink 
at the same time. When this occurs, every member of each troop 
sounds a bold roar of defiance at the opposite parties; and when 
one roars, all roar together, and each seems to vie with his comrades 
in the intensity and power of his voice. The power and grandeur 
of these nocturnal concerts are inconceivably striking and pleasing to 
the hunter’s ear.” 

“The usual pace of a Lion,” C. J. Andersson? says, “is a walk, 
and, though apparently rather slow, yet, from the great length of 
his body, he is able to get over a good deal of ground in a short 
time. Occasionally he trots, when his speed is not inconsiderable. 
His gallop—or rather succession of bounds—is, for a short distance, 
very fast—nearly or quite equal to that of a horse. Indeed, unless 
the steed has somewhat the start when the beast charges, it will be 
puzzled to escape. Many instances are on record of horsemen who 
have incautiously approached too near to the Lion, prior to firing, 
who have been pulled down by him before they could get out of 
harm’s way. Happily, however, the beast soon tires of the exertion 
of galloping, and unless his first rush succeeds he, for the most part, 
soon halts and beats a retreat.” “The Lion, as with other members 
of the feline family,” the same writer tells us, “seldom attacks his 
prey openly, unless compelled by extreme hunger. For the most part 
he steals upon it in the manner of a cat, or ambushes himself near 
to the water or a pathway frequented by game. At such times he 
lies crouched upon his belly in a thicket until the animal approaches 
sufficiently near, when, with one prodigious bound, he pounces upon 
it. In most cases he is successful, but should his intended victim 
escape, as at times happens, from his having miscalculated the 
distance, he may make a second or even a third bound, which, 
however, usually prove fruitless, or he returns disconcerted to his 
hiding-place, there to wait for another opportunity.” His food con- 
sists of all the larger herbivorous animals of the country in which 

1 The Lion and the Elephant, 1873, p. 19. 


FELIDA 509 


he resides-—buffaloes, antelopes, zebras, giraffes, or even young 
elephants or rhinoceroses, though the adults of these latter he dare 
not attack. In cultivated districts the cattle, sheep, and even human 
inhabitants are never safe from his nocturnal ravages. He appears, 
however, as a general rule, only to kill when hungry or attacked, 
and not for the mere pleasure of killing, as with some other car- 
nivorous animals. Moreover, he by no means limits himself to 
animals of his own killing, but, according to Selous, often prefers 
eating game that has been killed by man, even when not very fresh, 
to taking the trouble to catch an animal himself. All books of 
African travel and sport abound with stories, many of which are 
apparently well authenticated, of the lion’s prodigious strength, as 
exemplified by his being able to drag off a whole ox in his mouth 
to a long distance, even leaping fences and dykes with it. 

The Lion appears to be monogamous, a single male and female 
continuing attached to each other irrespectively of the pairing 
season. At all events the Lion remains with the Lioness while the 
cubs are young and helpless, and assists in providing her and them 
with food, and in educating them in the art of providing for them- 
selves. The number of cubs at a birth is from two to four, usually 
three. They are said to remain with their parents till they are 
about three years old. The following account by an eye-witness 
gives a good idea of Lion family life! :— 

“T once had the pleasure of, unobserved myself, watching a 
lion family feeding. JI was encamped on the Black Umfolosi in 
Zululand, and towards evening, walking out, about half a mile 
from camp, I saw a herd of zebra galloping across me, and when 
they were nearly 200 yards off, I saw a yellow body flash towards 
the leader, and saw him fall beneath the lion’s weight. There 
was a tall tree about 60 yards from the place, and anxious to see 
what went on, I stalked up to it, while the lion was still too much 
occupied to look about him, and climbed up. He had by this time 
quite killed the beautifully striped animal, but instead of proceed- 
ing to eat it, he got up and roared vigorously, until there was an 
answer, and in a few minutes a lioness, accompanied by four 
whelps, came trotting up from the same direction as the zebra, 
which no doubt she had been to drive towards her husband. 
They formed a fine picture as they all stood round the carcase, 
the whelps tearing it and biting it, but unable to get through the 
tough skin. Then the lion lay down, and the lioness driving her 
offspring before her did the same four or five yards off, upon which 
he got up, and, commencing to eat, had soon finished a hind leg, 
retiring a few yards on one side as soon as he had done so. The 
lioness came up next and tore the carcase to shreds, bolting huge 


1 Hon. W. H. Drummond, The Large Game and Natural History of South 
and South-East Africa, 1875, p. 278. 


510 CARNIVORA 


mouthfuls, but not objecting to the whelps eating as much as they 
could find. There was a good deal of snarling and quarrelling 
among these young lions, and occasionally a stand-up fight for a 
minute, but their mother did not take any notice of them, except 
to give them a smart blow with her paw if they got in her way. 
. . . There was now little left of the zebra but a few bones, which 
hundreds of vultures were circling round waiting to pick, while 
almost an equal number hopped awkwardly about on the ground 
within 50 or 60 yards of it, and the whole lion family walked 
quietly away, the lioness leading, and the lion, often turning his 
head to see that they were not followed, bringing up the rear.” 

Though not strictly gregarious, Lions appear to be sociable 
towards their own species, and often are found in small troops, 
sometimes consisting of a pair of old Lions, with their nearly full- 
grown cubs, but occasionally of adults of the same sex ; and there 
seems to be good evidence that several Lions will associate together 
for the purpose of hunting upon a preconcerted plan. As might 
be supposed, their natural ferocity and powerful armature are 
sometimes turned upon one another; combats, often mortal, occur 
among male Lions under the influence of jealousy ; and Andersson 
relates an instance of a quarrel between a hungry Lion and Lioness 
over the carcase of an Antelope which they had just killed, and 
which did not seem sufficient for the appetite of both, ending in 
the Lion not only killing, but even devouring his mate. Old Lions, 
whose teeth have become injured with constant wear, often become 
“man-eaters,” finding their easiest means of obtaining a subsistence 
in lurking in the neighbourhood of villages, and dashing into the 
tents at night and carrying off one of the sleeping inmates. Lions 
differ from most of the smaller Felide in never climbing trees ; 
indeed, as mentioned before, they are rarely found in forests. 

With regard to the character of the Lion, those who have had 
opportunities of observing it in its native haunts differ greatly. 
The exaggerated accounts of early writers as to its courage, 
nobility, and magnanimity have led to a reaction, which causes 
some modern authors to speak of it in language quite the reverse, 
and to accuse it of positive cowardice and all kinds of meanness. 
Livingstone goes so far as to say, “‘ Nothing that I ever learned of 
the lion could lead me to attribute to it either the ferocious or 
noble character ascribed to it elsewhere,” and he adds that its roar 
is not distinguishable from that of the ostrich. Of course these 
different estimates depend to a great extent upon the particular 
standard of the writer, and also upon the circumstance that 
Lions, like other animals, undoubtedly show considerable individual 
differences in character, and behave differently under varying cir- 
cumstances. They are certainly not so reckless as‘ to be entirely 
devoid of the instinct of self-preservation, and if one, perhaps 


FELIDE S11 


satiated with a good meal the night before, unexpectedly disturbed 
in the day-time, will occasionally retreat when confronted, even by 
an unarmed man, that is scarcely a reason for assigning cowardice 
as one of the characteristics of the species. The latest authority, 
Selous, while never denying the daring courage of the Lion when 
hungry or provoked, and vindicating the awe-inspiring character of 
the roar of several Lions in unison, when heard at close quarters, 
as the grandest sound in nature, says with regard to its outward 
aspect :— 

“Tt has always appeared to me that the word ‘majestic’ is 
singularly inapplicable to the lion in its wild state, as when seen 
by daylight he always has a stealthy furtive look that entirely 
does away with the idea of majesty. To look majestic a lion 
should hold his head high. This he seldom does. When walking 
he holds it low, lower than the line of his back, and it is only 
when he first becomes aware of the presence of man that he some- 
times raises his head and takes a look at the intruder, usually 
lowering it immediately, and trotting away with a growl. When 
at bay, standing with open mouth and glaring eyes, holding his 
head low between his shoulders, and keeping up a continuous low 
growling, twitching his tail the while from side to side, no animal 
can look more unpleasant than a lion; but there is then nothing 
majestic or noble in his appearance.” 

Notwithstanding this evidently truthful description of the 
animal when seen under what may be called unfavourable circum- 
stances, no one with an eye for beauty can contemplate the form 
of a fine specimen of a Lion, at all events in a state of repose, even 
though in the confinement of a menagerie, without being impressed 
with the feeling that it is a grand and noble-looking animal. 

The Tiger (F. tigris) is so closely related to the Lion that it is 
chiefly by external characters that the two species are distinguished. 
There are, however, slight distinctions in the proportionate size of 
the lower teeth, the general form of the cranium, and the relative 
length of the nasal bones and ascending processes of the maxillaries 
by which the skull of the Lion and Tiger can be easily discriminated 
by the practised observer. 

Although examples of both species present considerable varia- 
tions in size, and reliance cannot always be placed upon alleged 
dimensions, especially when taken from skins stripped from the 
body, it seems well ascertained that the length of the largest-sized 
Bengal Tiger may exceed that of any Lion. According to Mr. W. 
T. Blanford,! adult males measure from 5} to 64 feet from the 
nose to the root of the tail; the tail itself measuring some 3 feet 
in length. Measured along the curves of the head and back to the 
tip of the tail, males usually give a length of from 9 to 10 feet, 

1 Fauna of British India, ‘‘ Mammalia,” p. 59 (1888). 


512 CARNIVORA 


but some specimens reach to 12 feet. The female is somewhat 
smaller, and has a lighter and narrower head. The Tiger has no 
mane, but in old males the hair of the cheeks is rather long and 
spreading. The ground colour of the upper and outer parts of the 
head, body, limbs, and tail is a bright rufous fawn, and these parts 
ave beautifully marked with transverse stripes of a dark, almost 
black colour. The markings vary much in different individuals, 
and even on the two sides of the same individual. The under 
parts of the body, the inside of the limbs, the cheeks, and a large 
spot over each eye are nearly white. The Tigers which inhabit 


Via, 225.—The Tiger (Felis tigris). 


hotter regions, as Bengal and the south Asiatic islands, have shorter 
and smoother hair, and are more richly coloured and distinctly 
striped than those of Northern China and Siberia, in which the fur 
is longer, softer, and lighter coloured. 

The Tiger is exclusively Asiatic, but has a very wide range in 
that continent, having been found in almost all suitable localities 
south of a line drawn from the river Euphrates, passing along the 
southern shores of the Caspian and Sea of Aral by Lake Baikal to 
the Sea of Okhotsk. Its most northern range is the territory 
of the Amur, its most southern the islands of Sumatra, Java, and 
Bali. Westward it reaches to Turkish Georgia and eastward to 
the island of Saghalin. It is absent, however, from the great 
elevated plateau of Central Asia, nor does it inhabit Ceylon, 


FELIDAS 513 


Borneo, or the other islands of the Indo-Malayan Archipelago, 
except those above mentioned. Its absence from Ceylon leads 
Mr. Blanford to conclude that the Tiger has only recently migrated 
into Southern India. 

The principal food of the Tiger in India is cattle, deer, wild hog, 
and pea-fowl, and occasionally human beings. The regular * man- 
eater” is generally au old Tiger whose vigour is passed, and whose 
teoth are worn and defective: it takes up its abode in the ucigh- 
bourhood of a village, the population of which it finds an easier 
prey than the larger or wilder animals named above. Though 
chietly affecting grassy plains or swamps, it is also found in forests, 
and seems to be fond of haunting the neighbourhood of old ruins. 
As arule, Tigers do not climb trees: but when pressed by fear, as 
during an inundation, they have been known to do so. They take 
to the water readily and are good swimmers. The Tigers of the 
Sundarbans (Ganges delta) continually swim from one island to the 
other to change their hunting-grounds for deer. The following 
oxtract on the habits of the Tiger is taken from Sir J. Fayrer’s 
Royal Liger of Bengal (A879) :— 

“The tigress gives birth to from two to five, even six cubs; 
but three is a frequent number. She is a most attectionate and 
attached mother, and generally guards and trains her young with 
tho most watehful solicitude. They remain with her until nearly 
full grown, or about the second year, when they ave able to lall for 
themselves and begin life on their own account, Whilst they 
remain with her she is peculiarly vicious and aggressive, defending 
them with the greatest courage and energy, and when robbed of 
them is terrible in her rage: but she has been known to desert 
them when pressed, and even to cat them when starved. As soon 
as they begin to require other food than her milk, she kills for 
them, teaching them te do so for themselves by practising on small 
animals, such as deer and young calves or pigs. At these times 
she is wanton and extravagant in her eruelty, killing apparently 
for the gratification of her ferocious and bloodthirsty nature, and 
pethaps to excite and instruct the young ones, and it is not until 
they are thoroughly capable of killing their own food that she 
separates from them. The young tigers are fav more destructive 
than the old. They will kill three or four cows at a time, whilst 
the older and more experienced rarely kill more than one, and this 
at intervals of from three or four days to a week. For this pur- 
pose the tiger will leave its retreat in the dense jungle, proceed to 
the neighbourhood of a village or gowrie, where cattle feed. and 
during the night will steal on and strike down a bullock, drag it 
into a secluded place, and then remain near the ‘marrie, or 
“kill, for several days, until it has eaten it, when it will proceed 
in search of a further supply, and, having found good hunting 


Qe 
care) 


514 CARNIVORA 


ground in the vieinity of a village or gowrio, continuo its ravages, 
destroying one or two cows or bufliloes a week. It is very fond of 
the ordinary domestic cattle, which in the plains of India are 
generally weak, balf-starved, under-sized: creatures. One of these 
is easily struck down and carried or dragged off, ‘The smaller 
buffaloes are also easily disposed of ; but the buffalo bulls, and 
especially the wild ones, are formidable antagonists, and have 
often been known to beat the Tiger off, and even to wound him 
seriously.” 

In many districts of India the number of ‘Tigers has been very 
considerably diminished of late years. In somo other conntrios 
they appear, however, to be on the inerease ; thus according to 
one of the administration reports of Java laid before the Duteh 
Chambors, portions of that island are being depopulated through 
Tigers. In 1882 the population of a villago in the south-west of 
the Bantam province was removed and transferred to an islind olf 
the coast in consequence of the trouble caused to the people by 
Tigers. These animals have now become an intolerable pest in 
parts of the same province. The total population is about 600,000, 
and, in [887, sixty-one were killed by ‘Tigers, and in consequence 
of the dread existing among the people, it has been proposed to 
deport the inhabitants of the villages most. threatened to other 
parts of the country whore Tigers are not so common, and where 
they can pursue their agricultural occupations with a yreater 
degree of security. At present they foar to go anywhere near 
the borders of the forest. The people scem disinclined, or they 
lack the means and courage, to attack and destroy their cnemy, 
although considorable rewards aro offered by Government for the 
destruction of beasts of proy. In 1888 the reward for killing a 
Royal Tiger was raised to two hundred florins. It appears also that 
the immunity of the Tiger is in part due to superstition, for it is 
considered wrong to kill one unless he attacks first. or othorwise 
does injury 

The Leopard (2. pardus, Fig. 226), although belouging to the 
same restricted group as the two preceding species, is distinguished 
from both by its inferior size, and its coloration. The animal 
now commonly known as the Leopard was called Pard (adpdos and 
mapdudts) or Panther (rdvOyp) by tho ancients. Leopard (deo-pardus) 
is a later term, originally applied, it is believed, to the Chota or 
Hunting Leopard, upon the supposition that it was a creature 
intermediate between the Lion and the trne Pard. If so it) has 
heen completely transferred to the more common species, and 
though in this sense a porfectly unnecessary and unmenning term, 
has gradually superseded those by which this was originally known. 
Pard, so commonly used by Elizabethan authors, is now nearly 
obsolete in the Knglish language, and Panther has either become 


FELIDAE 515 


synonymous with Leopard, or is used vaguely for any similar large _ 
feline animal, even the Puma of America. 

Owing to their extensive geographical range, and the great 
variations, both in size, form, and coloration to which Leopards are 
subject, zoologists have scarcely decided whether all the forms 
popularly referred to this animal should be regarded as specifically 
alike, or whether they should constitute several distinct species, 
but the prevailing opinion is in favour of the former view. The 


Fic. 226.—The Levpard (Felis pardus). 


attempts to separate a larger and more robust variety, under the 
name of Panther, from a smaller and more graceful form, to which 
the term Leopard might properly be restricted, have failed, owing 
to the existence of intermediate conditions which cannot he assigned 
definitely to either one or the other form.’ The most marked 
anatomical difference yet noted in different varieties of leopard is 
in the length of the tail as compared with that of the body, even 
the number of the caudal vertebre showing variation, though within 
what limits, and whether correlated with other characters, has not 
yet been clearly ascertained. The fur of those specimens which 
inhabit the most northern confines of its range of distribution, as 


1 See W. T. Blanford, Fauna of British India, ‘‘Mammalia,” p. 69 (1888). 


516 CARNIVORA 


North China, is longer and softer, and the markings are con- 
sequently less distinct than on those from more congenial climates, 
and the well-marked variation thus produced has given rise to the 
idea of specific distinction. 

The size of different individuals, as before said, varies greatly, 
the head and body usually measuring from 3} to 44 feet in length, 
and the tail from 21 to 3 feet, but specimens have been met with 
which fall short of or exceed these limits. The ground colour of 
the fur varies from a pale fawn to a rufous buff, graduating into a 
pure white on the under parts and inside of the limbs. This is 
spotted over with dark brown or black ; the spots on the back and 
sides being arranged in rosettes or broken rings, which vary greatly 
in size and distinctness in different individuals, but are without the 
central spot seen in those of the Jaguar. The spots on the under 
parts and limbs are simple and blacker than those on the other parts 
of the body. The bases of the ears behind are black, the tips buff. 
The upper side of the tail is buff, spotted with broken rings like 
the back, its under surface white with simple spots. The hair of 
the cubs is longer than that of the adults, its ground colour less 
bright, and its spots less distinct. Perfectly black Leopards, which, 
however, in certain lights show the characteristic markings on the 
fur, are not uncommon. These appear to be examples of melanism, 
occurring as individual variations, sometimes in one cub out of a 
litter of which the rest are normally coloured, and therefore not 
indicating a distinct race, much less a species. These are met 
with chiefly in Southern Asia. We are not aware of any recorded 
case from Africa, though there seems no reason why they should 
not occur. 

In habits the Leopard resembles the other large Cat-like animals, 
yielding to none in the ferocity and bloodthirstiness of its dis- 
position. It is exceedingly quick and active in its movements, but 
seizes its prey by waiting in ambush or stealthily approaching to 
within springing distance, when it suddenly rushes upon it and 
tears it to the ground with its powerful claws and teeth. It preys 
upon almost any animal it can overcome, such as antelopes, deer, 
sheep, goats, monkeys, peafowls, and is said to have a special liking 
for dogs. It not unfrequently attacks human beings in India, 
chiefly children and old women, but instances have been known of 
a Leopard becoming a regular “man-eater.” When favourable 
opportunities occur, it often kills many more victims than it can 
devour at once, apparently to gratify its propensity for killing, or 
only for the sake of their fresh blood. It generally inhabits woody 
districts, and can climb high trees with facility if necessary for its 
safety when hunted, but usually lives on or near the ground, among 
rocks, bushes, and roots and low branches of large trees. . 

The present geographical range of the Leopard is very extensive, 


FELID.E 517 


as it is met with in various suitable localities, where not too much 
interfered with by human cultivation, throughout the greater part 
of Africa from Algeria to the Cape Colony, and through the whole 
of the South of Asia from Palestine to China, including all India 
south of the Himalaya, and the islands of Ceylon, Java, Sumatra, 
and Borneo. Fossil bones and teeth, indistinguishable from those 
of existing Leopards, have been found in cave-deposits of Pleisto- 
cene age in Spain, France, Germany, and England. The evidence 
of the former existence of the Leopard in England is described at 
length by Boyd Dawkins and Sanford in their Biifish Pleistocene 
Mammalia 

The Ounce, or Snow Leopard (F. unci), inhabits the highlands 
of Central Asia, from the lofty mountains of Tibet to the southern 
parts of Siberia, at altitudes of from 9000 to 18,000 feet above the 
sea. It is about the size of the common Leopard, but lighter in 
colour, with longer fur, less distinct spots, and a long thick tail. 
Its skull differs in shape from that of all the other Felid ; the 
facial portion being very broad, the nasal bones especially being 
wide and depressed, and the zygomatic arches very strong and 
deep. The Clouded Tiger (F. nebulssa) is a beantifully marked 
species, with elongated head 
and body, long tail, and rather 
short limbs. The canine teeth 
are proportionally longer than 
in any existing member of 
the genus. It is thoroughly 
arboreal, and is found in the 
forests of South-East Asia and 
the islands of Sumatra, Java, 
Borneo, and Formosa. 
F. serval, the Serval, from 
South Africa, is yellow with 
black spots, and has a short 
tail and large ears. Numer- 
ous smaller species called Tiger 
Cats and Wild Cats, of which 


the Oriental F. marmorata wae eas : a 
(Fig. 227) isa good example, Fic. 237.—The Marbled Cat (Felis marmorata). 
From Blanford, Mammalia of British India, p. 74, 


are found throughout the 
warmer parts of Asia and 
Africa. The Wild Cat of Europe, F. cafus, still inhabits the 
mountainous and wooded parts of Great Britain. 

The Catire Cat (F. caffra*), of Africa and Southern Asia, was the 
species held in veneration by the ancient Ezyptians, and immense 
1 Monographs of the Paleontographica! Society, 1872. 

2 syn. F. imacrocelis. ° Syn. F, maniculata and ealigata. 


after Elliot. 


518 CARNITORA 


numbers of its mummified remains havo recently been found in 
Egypt, whence they have been imported in large quantities to this 
country for manure. This species is generally regarded as the 
main ancestral stock from which the European Domestic Cat has 
been derived ; one of the arguments in support of this opinion being 
that the whole of the sole of the hind foot of /. caffra is black, and 
that the same feature obtains in the darker varicties of the Domestic 
Cat; while in F. catus there ave only spots of black upon this 
portion of the limb. Remains of tho Caflre Cat occur in the 
Pleistocene cave-deposits at Gibraltar, The Indian 2. rubigiinose is 
the smallest species of Cat. , 

The Caracal or Persian Lynx (2. earaeal) is an animal about 
the size of a fox, of slender build, with a moderately long tail, 
reaching down to the heels. It is of a uniform vinous or bright 
fulvous brown colour above, and is paler, sometimes almost. white, 
beneath. It is quite or almost entirely unspotted. The tail has a 
black tip, and the ears are black externally, long, upright, pointed, 
and surmounted by a pencil of fine black hairs. It inhabits Central 
and North-West India, Persia, Arabia, Syria, and the greater part 
of Africa. 

The true Lynxes comprise various species or varieties found 
in the northern and temperate regions of both the Old and New 
World, all larger than the truo Wild Cats, with long limbs, short 
stumpy tail, ears tufted at the tip, and pupil of the eye linear when 
contracted, Their fur is generally long and soft, varying, however, 
according to season and locality, and always longish upon the 
cheeks. Their colour is always light brown or gray, and generally 
more or less spotted with a darker shade. The naked pads of the 
feet are more or less covered by the haiv that grows between them. 
The skull and skeleton do not diffor markedly from those of the 
other cats, but the small anterior upper premolar tooth found in 
many other species is usually wanting ; and the lower carnassial has a 
rudimental talon, Their habits ave exactly those of the other Wild 
Cats, and they are execeded by none in the untiameble savageness 
of their disposition. They capture their prey in the samo manner, 
either lying in wait, or noisclessly stealing within reach, and then 
making « sudden rush or spring upon it. Their food cousists of 
any mammals or Iirds which they can overpower. In inhabited 
countries they commit extensive ravages upon sheep, lambs, and 
poultry. Lynxes genorally frequent. rocky places and forests, being 
active climbers, and passing much of their time among the brauches 
of the trees. Their skins are of considerable commercial value. 

Zoologists ave by no mews agreed at present as to the specific 
distinctions, if any really exist, between the various modifieations 
of this group. As many as cight species ave sometimes recognised, 
four belonging to the Old and four to the New World. The former 


FELIDA 519 


are /’. lynx, of Scandinavia, Russia, Northern Asia, and till lately 
the forest regions of Central Europe ; though not an inhabitant of 
Britain during the historic period, its remains have been found in 
cave-deposits of Pleistocene age; F. cervaria, Siberia; PF. pardina, 
Turkey, Greece, Sicily, Sardinia, and Spain; and F”. isabellina, Tibet. 
The American varieties are F. canadensis, the most northern species, 
and F. rufa, the American Wild Cat or Bay Lynx, extensively dis- 
tributed from the Atlantic to the Pacific throughout nearly the 
whole latitude of the United States, but replaced in Texas and 


Via. 228.— European Lynx (felis lynx). Frou a drawing by Wolf in Elliot’s 
Monograph of the Felide. 


southern California by J. iueulats, and in northern Oregon and 
Washington territory by I. fasciata. 

In both cases, as might be supposed, specimens obtained from 
the more southern climates are shorter in their fur, more brightly 
coloured, and more distinctly spotted than those from colder regions. 
When only a few individuals of each most markedly different form 
are examined the distinctions are sufficiently evident. The occur- 
rence, however, of transitional or intermediate forms makes it 
extremely difficult to draw the line between the different varieties 
or species, or to assign definite characters by which they can be 
separated. Wherefore it is best at present to accept the so-called 
species as only provisional, and wait until more abundant materials, 
with fuller knowledge of the localities from which they are derived, 


520 CARNIVORA 


and of the variations due to age, sex, season, and climate, have 
been more carefully studied. We shall then probably come to the 
conclusion that all or nearly all the existing forms of northern 
Lynxes, whether American or Eurasian, belong to what may fairly 
be called a species, which is becoming by degrees differentiated into 
several more or less strongly marked local varieties. Mr. W. T. 
Blanford has indeed shown that the Tibetan Lynx (JF. isabellina) 
is inseparable from F. lyn ; the specimens from Gilgit being inter- 
mediate in colour between the typical forms of the two races. 
On the other hand, from the evidence of cranial characters, Professor 
Mivart is disposed to regard F. pardina as a valid species. 


Fic, 220.—The Puma (Felis concolor). 


B. New World Species—The Puma or Couguar (£. concolor, Fig. 
229), commonly called “Panther” in the United States, is about 
the size of a Leopard, but of an uniform brown colour. It usually 
measures from nose to root of tail about 40 inches, the tail being 
rather more than half that length. The head is rather small com: 
pared with that of other Cats and has no mane. The ears are large 
and rounded. The tail is cylindrical, with some bushy elongation 
of the hairs near the end, but not forming a distinct tuft as in the 
Lion. The general colour of all the upper parts and sides of the 
adult is a tawny yellowish-brown, sometimes having a gray or 
silvery shade, but in some individuals dark or inclining to red. 
The lower parts of the body, inner surface of the limbs, the 
throat, chin, and upper lip are dirty white; the outside of the ears, 


FELIDE 521 


particularly at their base, and a patch on each side of the muzzle 
black ; the end of the tail dusky. The young are, when born, 
spotted with dusky brown and the tail ringed ; these markings 
gradually fading, and quite disappearing before the animal becomes 
full-grown. 

The Puma has an exceedingly wide range of geographical 
distribution, extending over a hundred degrees of latitude, from 
Canada in the north to Patagonia in the south, and was formerly 
pretty generally diffused in suitable localities from the Atlantic to 
the Pacific Ocean, but the advances of civilisation have in recent 
years considerably curtailed the extent of the districts which it 
inhabits. In Central America it is still common in the dense forests 
which clothe the mountain ranges as high as 8000 or 9000 feet 
above the sea-level, where the hideous sound of its howling is 
said to be almost continuously heard at night during the breeding 
season. Though an expert climber, it is by no means confined to 
wooded districts, being frequently found in scrub and reeds along 
the banks of rivers, and even in the open pampas and prairies. Its 
habits much resemble those of the rest of the group to which it 
belongs; and, like the Leopard, when it happens to come within 
reach of an abundant and easy prey, as the sheep or calves of an 
outlying farming station, it kills far more than it can eat, either 
for the sake of the blood only or to gratify its propensity for 
destruction. It rarely attacks man, and, when pursued, escapes if 
possible by ascending lofty trees. Several instances have occurred 
of Pumas becoming tame in captivity. Edmund Kean, the cele- 
brated actor, had one which followed him about like a dog. When 
caressed they express their pleasure by purring like a domestic 
cat. ; 

F. onca, the Jaguar, is a larger and more powerful animal than 
the last, and more resembles the Leopard in its colours. It also is 
found in both North and South America, but with less extensive 
range, reaching northwards only as far as Texas, and southwards 
nearly to Patagonia. It climbs as well as the Puma, and preys to 
a great extent upon monkeys. Several allied smaller elegantly 
spotted forms inhabiting the intratropical regions of America are 
commonly included under the name of Ocelot or Tiger Cat, though 
zoologists are still undecided whether under this designation several 
distinct species have not been confused, or whether all the Ocelots 
are to be referred to a single species (/. pardalis) showing great 
individual or racial variation. Their fur has always a tawny yellow 
or reddish-gray ground colour, and is marked with black spots, 
aggregated in streaks and blotches, or in elongated rings enclosing 
an area which is rather darker than the general ground colour. 
They range through the wooded parts of tropical America, from 
Arkansas in the north as far south as Paraguay, and in their habits 


522 CARNIVORA 


resemble the other smaller members of the Cat tribe, being ready 
climbers and exceedingly bloodthirsty. 

LF. yagquavund’, rather larger than the Domestic Cat, with an 
elongated head and body, and of a uniform brownish-gray colour, 
ranges from Matamoras to Paraguay. . eyrw is a small Cat, very 
Musteline in form, having an elongated head, body, and tail, and 
short limbs, and is also of a uniform light reddish-brown colour. 
It is a native of South America and Mexico. F. pajeros is the 
Pampas Cat. The American Lynxes have been already noticed 
with those of the Old World. 

C. Fossil Species—It has been already incidentally mentioned 


Fic. 230.—The Ocelot (Felis pardalis). 


that several of the existing species of Felis, such as the Lion 
Leopard and Caffre Cat, are met with in a fossil condition in the 
European Pleistocene deposits, and it may be added that the Pardine 
Lynx has left its remains in the cavern-deposits of Gibraltar. The 
caves of Brazil have yielded remains of the Jaguar and Ocelot ; 
while the Puma is found in the Pleistocene of the United States, 
Existing species now inhabiting India are met with in cayern- 
deposits in Madras. In the Pliocene Siwaliks of Northern India 
the huge extinct F. cristutu shows characters connecting it both 
with the Tiger and the Jaguar ; and the same deposits also contains 
the remains of a small species of the size of F. bengalensis. In 
Europe numerous species occur in the Upper and Lower Pliocene, 


FELIDA 523 


some of which were as large as a Leopard. /. atrom and F. augusta, 
of the Pliocene of the United States, were of the dimensions of the 
Lion. 

Cyncelurus.i—The Cheeta or Hunting Leopard (C. jubatus) is dis- 
tinguished from the other Felide by the inner tubercle of the upper 
carnassial, though supported by a distinct root, having no salient 
cusp upon it; by the tubercular molar being more in a line with 
the other teeth; and by the claws being smaller, less curved, and 
less completely retractile, owing to the feebler development of the 
elastic ligaments. The skull is short and high, with the frontal 
region broad and elevated in consequence of the large development 
of the frontal air-sinuses. The head is small and round, the body 
light, the limbs and tail long. Its colour is pale yellowish-brown 
with small black spots. The Cheeta is less savage and more 
easily tamed than most of the Cats. In Asia it has been trained 
for the chase of the Antelope. It has rather an extensive geo- 
graphical range from the Cape of Good Hope, throughout Africa 
and the south-western parts of Asia, as far as Southern India. 

Extinct Generu—A number of forms are gradually becoming 
known, especially through the researches of American palzonto- 
logists, which, though evidently animals of the same general type, 
and therefore to be placed in or near the family Felidw, depart so 
much in various details of structure that they must be referred to 
different genera. As one of the points in which Felis manifests its 
specialisation is the reduction of the number of the molar series of 
teeth, with concomitant shortening of the jaws, it might be 
supposed that in the earlier and perhaps ancestral forms these 
teeth would be more numerous and approach more nearly to the 
primitive or typical number of the heterodont mammals, viz. seven 
on each side. This is actually the case. Similarly we find that 
many of these forms exhibit a less specialised structure of the teeth 
themselves, as is shown by the absence of the anterior lobe of the 
upper carnassial, and the retention of the hind talon in the corre- 
sponding lower tooth. Again, some of them have an alisphenoid 
canal in the skull; while the femur may have a third trochanter, 
and the claws be very imperfectly retractile. 

An extremely generalised form is the small Proclurus, from the 
Upper Eocene and Lower Miocene, with p 4, m 4, an alisphenoid 
canal, and a third trochanter to the femur. Divictis, of the North 
American Miocene, is a larger allied form, with p 3, m 4; the upper 
carnassial having no anterior lobe, and the ungual phalanges being 
devoid of bony sheaths. The characters of the base of the skull, 
and the form and relations of the astragalus, differ very consider- 
ably from Felis. Pseudelurus, from the French Miocene, is another 
very generalised Feline, in which there may be either three or four 


1 Wagler, Syst. Amphib. ete. p. 30 (1830). 


524 CARNIVORA 


premolars, and the lower carnassial may retain its inner cusp. 
-Elurictis, of the French Phosphorites, with j =— m a together 
with several American Miocene genera, such as Vimravus (p 3, 
m 4), drchelurus (p aap m 4), Pogonodon (p 3, m +), and Hoplo- 
phoneus (p cS, m 1), approach more closely to the modern Cats, 
although many or all of them retain the alisphenoid canal, and have 
not yet developed the anterior lobe to the upper carnassial, or lost 
the talon to the lower one. Hoplophoneus has a descending flange 
to the mandible ; and its scapholunar bone has a line indicating its 
dual origin; while the femur still retains the third trochanter, 
of which all traces are lost in the modern Cats. 

On the other hand, some of the extinct Felide show a most 
remarkable tendency towards a specialisation not occurring in any 
of the surviving members of the family, viz. an enormous develop- 
ment of the upper canines, with which is usually associated an 
expansion downwards and flattening of the anterior part of the 
ramus of the lower jaw, on the outer side of which the canine lies, 
when the mouth is closed. In JJacheredus neogeus, the Sabre- 
toothed Tiger, from the caves of Brazil and also from Pleistocene 
deposits near Buenos Ayres, an animal about the size of a Tiger, 
these teeth are 7 inches in length, greatly compressed, and finely 
serrated on the trenchant anterior edges. Similar serrations are 
seen on a much fainter scale in the unworn teeth of modern Tigers. 
Many modifications of this commonly-called “ machzrodont ” type 
have been met with both in the Old and New World. In JL 
cultrudens, of the Upper Pliocene of Italy and France, the upper 
canine is long and narrow, with smooth cutting edges: the smaller 
form described as JL meganthereon being apparently the female 
of this species. I. crenatidens, of the same deposits, is distinguished 
by the shorter and broader upper canine, in which both edges are 
strongly serrated; the same feature occurring in the closely allied 
or identical IV. latidens of the English cavern-deposits. The Italian 
Pliocene form described as JV. nestianus has serrations only on the 
hinder edge of the upper canine, and the third lower premolar 
is separated by a long interval from the fourth, 1 necator, 
of the Pleistocene of South America, is remarkable as being the 
only member of the family in which the humerus has no ente- 
picondylar foramen. A very remarkable form, Eusmilus, from the 
Upper Eocene Phosphorites of Central France, differs from all other 
known Felines in having only two pairs of incisors in the lower jay, 
and a small canine separated by a very long diastema from the 
cheek-teeth, which consist only of one premolar and one sectorial 
true molar. The lower jaw is enormously expanded towards the 
symphysis to protect the large upper canines. This animal then, 
although of Eocene age, appears to form the culminating develop- 


VIVERRIDE 525 


ment of the sabre-toothed or macherodont dentition, the most 
specially carnivorous type of structure known. 

Other species of Alacherodus are found in the Pliocene de- 
posits of Europe and Asia. The accompany- 
ing woodcut exhibits the last two upper teeth 
of the Indian Jf siralensis, from which it will 
be seen that the inner tubercle of the carnassial 
is much reduced in size, while the molar is 
very minute. 


Fumily VIVERRIDZ. 


Premolars $ or 4. Molars + or 3. Upper 
carnassial usually without an anterior lobe, and Sp ee 
the lower one with a well-developed talon; of the tet upper carn 
second lower incisor (as in all the following 24 molar of Mack 
families) raised above the level of the first and “"”*"** 
third. Auditory bulla externally constricted, and divided by a 
septum. An alisphenoid canal (with very rare exceptions). Carotid 
canal distinct as a groove on the side of the bulla. Humerus 
usually with an entepicondylar foramen. Digits usually 5-5, but 
sometimes the pollex or hallux or both may be wanting. Dorsal 
vertebre 13 or 14. Limited in distribution to the Old World. 

The subfamily Cryptoproctine contains the single genus C'rypto- 
prota  Dentition: i 3, ¢ 4, p 4+, m 4+; total 36. The teeth 
generally closely resemble those of the Felide. The first premolar 
of both jaws is very minute and early deciduous. The upper 
carnassial has a very small inner tubercle, quite at the anterior part 
of the tooth. The true molar is very small and placed transverselv. 
The lower carnassial has a large trenchant bilobed blade, and a 
very minute talon, but no inner cusp. Skull generally like that of 
Felis, but proportionately longer and narrower. Orbit widely open 
behind. Vertebre: C 7, D 13, L 7,8 3, C 29. Body elongated. 
Limbs moderate in size. Feet subplantigrade ; five well-developed 
toes on each, with sharp, compressed, retractile claws. Ears 
moderate. Tail long and cylindrical. 

The only known species, C. feror, the “ Foussa” of the Malagasy, 
is peculiar to Madagascar, being the largest carnivorous animal in 
the island. It is about twice the size of the common Cat (5 feet 
from nose to end of tail), with short close fur of nearly uniform 
pale brown. Little is as yet known of its habits, except that it is 
nocturnal, frequently attacks and carries off goats, and especially 
kids, and shows great ferocity when wounded, on which account it 
is much dreaded by the natives. 

The remaining numerous specific and generic modifications found 


1 Bennett, Trans. Zool. Soc. vol. i. p. 137 (1833). 


526 CARNIVORA 


in the existing animals belonging to this family seem to arrange 
themselves mainly into two tolerably distinct groups, distinguish- 
able by the characters of the auditory bulla and neighbouring parts 
of the base of the skull, and by the structure of the feet. The one 
form has the genus /iverra or Civet Cats for its most typical repre- 
sentative, and the other Herpestes or the Ichneumons. 

Subfamily Viverrine.—Auditory bulla oval, or rather conical, 
broad and truncated and not everted behind, narrow in front ancl 
more or less compressed at the sides. The outer or anterior 
chamber very small and flat. The meatus with scarcely any 
inferior lip, its orifice being close to the tympanic ring. Par- 
occipital process triangular, its apex projecting slightly beyond the 
bulla. Claws strongly curved and more or less retractile. Perineal 
scent-glands generally present. 

This subfamily includes both Ethiopian and Oriental forms, but 
the former are the more numerous. 

The typical section, which includes five yenera, has the follow- 
ing characters. Dentition: 7 3,¢ 4, p 4, m 3 (4 in Prionodon) - 
total 40. Skull elongated ; facial portion small and compressed. 
Orbits well-defined but incomplete behind. Teeth always sectorial, 
never very small. Vertebre: C 7, D 13, L 7 (or D 14,L 86), 
5 3, C 22-30. Body elongated and compressed. Head pointed in 
front; ears rather small. Extremities short. Feet small and 
rounded. ‘Toes short, five on each foot. First toe both on fore 
and hind feet much shorter than the others. Palms and soles 
covered with hair, except the pads of the feet and toes, and in 
some species a narrow central line on the under side of the sole, 
extending backwards nearly to the heel. Tail moderate or long ; 
usually marked with dark and light rings. A pair of large glandular 
follicles situated on the perineum (in both sexes), and secreting in 
most species an oily substance of a peculiarly penetrating odour, 

The numerous species of this section form a large series, the 
two extremes of which differ considerably, but the several genera 
into which they may be divided blend so into one another that it is 
difficult to differentiate them sharply. 

All the animals of this section are, for their size, extremely 
active, fierce, and rapacious. They feed. chiefly on small mammals 
and birds. 

Virerra.—This includes the largest species. The teeth (Fig. 
232) are stouter and less compressed than in the other genera ; the 
second uppermolar being especially larger. The auditory bulla smaller 
and more pointed in front. Body shorter and stouter: limbs 
longer ; tail shorter, tapering. Under side of tarsus completely 
covered with hair. Claws longer and less retractile. Fur rather 

_ long and loose, and in the middle line of the neck and back usually 


4 Linn. Syst. Vat. 12th ed. vol. i. p. 63 (1766). 


VIVERRIDE 527 


elongated so as to form a sort of crest or mane; neck with a black 
gorget. Pupil circular when contracted. Perineal glands greatly 
developed. These characters apply especially to VY. cwetta, the 
African Civet, or “Civet-Cat” as it is commonly called, an animal 
rather larger than a common Fox, and an inhabitant of intra- 
tropical Africa. V. zibetha, the Indian Civet, of about equal size, 
inhabits Bengal, China, the Malay Peninsula, and adjoining islands. 
V. tangalunga, from Java, Sumatra, Borneo, and the Philippines, 


ee ee 


Fic. 232.—The left upper dentition of the Indian Civet (Viverra zibetha). From the 
Pulwontologia Indica. 


and V7. megaspila, from Burma, are smaller but nearly allied 
animals; the latter being more distinctly spotted than either of the 
others. From these species and the next-the civet of commerce, 
once so much admired as a perfume in England, and still largely 
used in the East, is obtained. The animals are kept in cages, and 
the odoriferous secretion collected from the interior of the perineal 
follicles with a spoon or spatula. 

The Rasse or Lesser Indian Civet (V. malaccensis) may be re- 
garded as the representative of a distinct group of Viwerra, although 
often referred to a separate genus (Viverricula). The size of this 
animal is smaller than in the typical group, the build is slighter, the 
muzzle finer, the claws sharper and more curved, and there is no 
erectile mane along the back. Generally there is an alisphenoid 
canal in the skull ; and the anterior chamber of the auditory bulla is 
much more inflated than the hinder one, so that the apparent length 
of the whole bulla is increased. This species is found over the 
greater part of India, and extends to the Malay Peninsula and 
Southern China. 

Large species of Viverra occur in the Pleistocene and Pliocene of 
India, and also in the Pliocene of France, which approximate in 
some characters of the dentition to the extinct genus Jctitherium, 
mentioned at the end of the family. Species of this genus have 
also been described from the Miocene and Upper Eocene of Europe. 
The Lower Miocene /”. antigua has an alisphenoid canal, and all the 
other cranial characters of the typical forms. 

Fossa..—The Fossa of Madagascar comes so close to the Rasse 


1 Gray, Proc. Zool. Soc. 1864, p. 518. 


528 CARNIVORA 


that its right to generic distinction seems doubtful. There is, 
however, no scent-pouch. The limbs are slender; and there are 
two small bare spots on the sole of the hind foot, above the 
plantar pads. There is no dark line along the back; the throat 
gorget of Viverra is absent; and in the tail the spots only tend to 
form rings, which are not complete. The skull has an alisphenoid 
canal, and a large bulla as in the typical group of Viverra. 
Genetta..—The Genettes are smaller animals, with more elon- 
gated and slender bodies, and shorter limbs than the Civets. Skull 
elongated and narrow. Auditory bulla large, elongated, rounded 


Fic. 233.—The Common Genet (Genetta vulgaris). 


at both ends. Teeth compressed and sharp pointed. The inner 
side of the third upper premolar has a tubercle not present in the 
previous genus, and the talon of the lower carnassial is larger. 
Pupil contracting to a linear aperture. Tail long, slender. Fur short 
and soft, spotted or cloudy. Under side of the tarso-metatarsus 
with a narrow longitudinal bald streak. No pouch for storing the 
secretion of the scent-gland. G. vulgaris, the common Genet 
(Fig. 233), is found in France south of the river Loire, Spain, 
South-Western Asia, and Africa from Barbary to the Cape. 
G. felina, senegalensis, tigrina, and pardalis are other named species, 
all African in habitat. 

A few details (taken from Professor Mivart’s memoirs on the 


* Cuvier, Régne-Animal, vol. i. p. 156 (1817). 


PIFERRID£E 


‘an 


29 


-Eluroidea) of the anatomy of the soft parts of the Genet may be 
ziven as illustration of these parts in the Carnivora generally, and 
of this family and genus im particular. The salivary glands are 
shown in Fig. 19 (p. 56), and these conform to the general type 
prevalent in the 
-Eluroidea. Thus 
there is a distinct 
zygomatic gland; 
the parotid with 
ig (Steno’s) duct 
is well developed : 
and there is a 
small = submaxil- 
lary gland) The 
stomach (Fig. 
234), while con- 
forming to the 
simple type char- 


acteristie of the 


Fic. 234 
ee . a pyloric valve: >, 
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 <te 
a f The puctll of the eve when 
zroup, and vertic- 
ons are required 


ameree cf su oh sinuses in 
: st members of the rst 


in the © hare bat more observa 


s character can be ah iv relied upon. The form and 
cf the tail is citen used ior the purposes of classification, 
ts characters do met comeide with those of the evarium. Since 
; wth American Canidz have 222 lonz bushy cals of 
Foxes and the sswis of Wolves. Takiez into aceount various 
combinations of 7hese and other minor characters, the species may 
be arranged in the folowing ereuns, which some authors have 
considered as of generic importance. 

A. faesii or Lupine —Thke typical sroup. or Canis proper, 
contains the larzest members of the zenus. the true Wolves of the 
northern parcs ‘of both Old and New Worlds (C. Imus, ere. the 

Jackals of Seuthern Asia and Africa (C. aureus, m =, ete.), and 
the various breeds of the domestic Doz (C. familiarizs. The wue 
Wolves are (excluding some varieties of the domescie Duz: the 
sargest members of the genus. and have a wide seosraphical mance. 
extending over nearly =e whole of Europe and Asia. and North 
America from Greenland to Mexico. but they are net found in 
South America or Africa, being replaced in bork of these e 
by various species of Jackals and Foxes. As mich: be eel 
i iS eXtensive ranze. and the varied ch oi the climatic 
conditions of the countries they inhabit. they presen: great diversi- 
ties of ce. length and thickness of tur, and colorazion, although 
resembling each ‘other in 21] Impertant structural char . These 
differences have given rise to a supposed uilrighctiy of species, 
a by the names of C. taps. CL dycaen (Centra! Europe). 
CL laniger and CL niger Ties. id pullipes (India), CL actigenieli 
Co neriis, Co meciogus, ete. of North America, but it is very doubt- 
ful whether some of these ought to be distinzuished as other than 
local varieties. Mr. W. T. Blanford, in his recent work on the 
mammals of India, regards the :wo forms from Tibet mentioned 
above as inseparable from C. jupus. In North America there is 
a very distinct smaller species. called the Corote or Prairie Wolf 
(Cc s): and perhaps the Japanese Wolf (C. hedej lex) may also 
be distine:. ‘sk: hough. except for its smaller size and shorter ! leas : 
is scarcely distinguishable from the common species Thos 
generally discribut red throughout the pele aia the bss 

Y Prov Zc #, Doe Fx 2s os Jackals, W-: 
aud Foxes: @ es eine the Catz da (Dsel. 


CANIDE 549 


Wolf (C. pallipes), which is rather smaller and slighter than C. Jupus, 
is not found in Ceylon, nor in Burma and Siam. The ordinary 
colour of the Common Wolf is a yellowish or fulvous gray, but 
specimens have been met with almost pure white and others entirely 
black. In northern countries the fur is longer and thicker, and the 
animal generally larger and more powerful than in the southern por- 
tion of its range; this being especially the case with the Tibetan 
races. The habits of the Wolf are similar everywhere, and it is still, 
and has been from time immemorial, especially known to man in all 
the countries it inhabits as the devastator of his flocks of sheep. They 
do not catch their prey by lying in ambush, or stealing up close to 
it and making a sudden spring as the Cat tribe do, but by fairly 
running it down in open chase, which their speed and remarkable 
endurance enable them to do; and usually, except during summer, 
when the young families of cubs are being separately provided for 
by their parents, they assemble in troops or packs, and by their 
combined and persevering efforts are able to overpower and kill 
even such great animals as the American Bison. It is singular that 
such closely allied species as the Domestic Dog and the Arctic Fox 
are among the favourite prey of Wolves, and, as is well known, 
children and even full-grown people are not unfrequently the 
objects of their attack when pressed by hunger. Notwithstanding 
the proverbial ferocity of the Wolf in a wild state, many instances 
are recorded of animals taken when quite young becoming perfectly 
tame and attached to the person who has brought them up, when 
they exhibit many of the ways of a Dog. They can, however, 
rarely be trusted by strangers. 

The history of the Wolf in the British Isles and its gradual 
extirpation has been thoroughly investigated by Mr. J. E. Harting 
in his work on Eatinct British Animals, from which the following 
account is abridged: To judge by the osteological remains which 
the researches of geologists have brought to light, there was per- 
haps scarcely a county in England or Wales in which, at one time 
or another, wolves did not abound, while in Scotland and Ireland 
they must have been still more numerous. The fossil remains 
which have been discovered in Britain are not larger than, nor in 
any way to be distinguished from, those of European wolves of the 
present day. Wolf-hunting was a favourite pursuit of the ancient 
Britons as well as of the Anglo-Saxons. In Athelstan’s reign these 
animals abounded to such an extent in Yorkshire that a retreat was 
built by one Acehorn, at Flixton, near Filey, wherein travellers 
might seek refuge if attacked by them. As is well known, great 
efforts were made by King Edgar to reduce the number of wolves 
in the country, but, notwithstanding the annual tribute of 300 
skins paid to him during several years by the king of Wales, he 
was not altogether so successful as has been commonly imagined. 


550 CARNIVORA 


In the reign of Henry III the number of wolves in some parts of 
the country was sufficient to induce the king to make grants of land 
to various individuals upon the express condition of their taking 
measures to destroy these animals wherever they could be found. 
In Edward IT’s time the king’s forest of the Peak, in Derbyshire, 
is especially mentioned as infested with wolves, and it was not 
until the reign of Henry VII (1485-1509) that wolves appear to 
have become finally extinct in England. This, however, is rather 
a matter of inference from the cessation of all mention of them in 
local records than from any definite evidence of their extirpation. 
Their last retreat was probably in the desolate wolds of Yorkshire. 
In Scotland, as might be supposed from the nature of the country, 
the wolf maintained its hold for a much longer period. There is a 
well-known story of the last of the race being killed by Sir Ewen 
Cameron of Lochiel in 1680, but there is evidence of wolves having 
survived in Sutherlandshire and other parts into the following 
century (perhaps as late as 1743), though the date of their final 
extinction cannot be accurately fixed. In Ireland, in Cromwell’s 
time, wolves were particularly troublesome, and said to be increas- 
ing in numbers, so that special measures were taken for their 
destruction, such as the offering of large rewards for their heads, 
and the prohibition (in 1652) of the exportation of “ wolf-dogs,” the 
large dogs used for hunting the-wolves. The active measures 
taken then and later reduced their numbers greatly, so that 
towards the end of the century they became scarce, but, as in the 
case of the sister island, the date of their final disappearance cannot 
now be ascertained. It has been placed, upon the evidence of 
somewhat doubtful traditions, as late as 1766. 

Remains of C. /upus are common in the European Pleistocene ; 
while the Indian Pliocene C. cauéleyi, of which the upper teeth 
are shown in Fig. 251, was probably the ancestor of C. pallipes. 
C. neschersensis, of the Upper Pliocene of France, was a smaller 
extinct Wolf. A lower jaw from the French Pleistocene, described 
under the name of Lycorus, has only three premolars, but evidently 
belongs to the Wolf. 

The Jackals are smaller than the Wolves, with the bushy tail 
about one-third the length of the head and body, and the car- 
nassials relatively shorter as compared with the tubercular molars. 
The Common Jackal (C. awreus, Fig. 252) has a very wide distri- 
bution, ranging from South-Eastern Europe through South-Western 
Asia to India and Burma, and also occurring in Northern Africa ; 
being replaced in the Ethiopian region by closely allied species. 
Remains indistinguishable from C. aureus occur in the Pliocene 
Siwaliks of Northern India. Jackals hunt at night in packs, 
uttering the piercing cries so well known to all who have resided 
in countries where these animals are found. 


CANIDA SSE 


The origin of the Domestic Dog, with its numerous breeds, 
has been, the subject of much controversy. Some naturalists 
believe it to be a distinct species, descended from one that no 
longer exists in a wild state; others have sought to find its pro- 
genitors in some one of the wild or feral races, either of true Dogs, 
Wolves, or Jackals; while others again believe that it is derived 
from the mingling of two or more wild species or races. It 
was probably the earliest animal domesticated by man, and few if 
any other species have undergone such an extraordinary amount of 
variation in size, form, and proportion of limbs, ears, and tail— 


avin 


HN 


\\\ 
AN 


al i iG 
yen 


Fic. 252.—The Jackal (Canis aureus). 


a 


variations which have been perpetuated and increased by careful 
selective breeding. The Dingo or Australian Dog is met with wild, 
and also as the domestic companion of the aboriginal people. Dogs 
were also in the possession of the natives of New Zealand and other 
islands of the Pacific, where no placental mammals exist naturally, 
on their discovery by Europeans in the last century. 

The second group includes the wild Dogs of the south-east of 
Asia, described as Cyon, and distinguished by slight modifications 
as C. rutilans, C. dukhunensis, and C. javanicus, and differing from 
the above in wanting the small last lower tubercular molar. This 
difference reduces the number of the teeth to the same as in 
Viverra, and is precisely paralleled by some of the species of the 
extinct genus Cynodictis mentioned below. The muzzle is shorter 


a32 CARNIVORA 


than in other species, and the facial profile is slizhtly convex 
instead of concave. The mamme are also 12 or 14 instead of <ke 
normal 10: while there is long hair between the foot-pads) Wild 
Dozs inhabit not only the whole of the Oriental region, but extend 
into Central Asia as far north as the Altai and Amurland (C. a.pinus.. 
, dukhunensis ranges from the fcrest regions of peninsular India to 
Gilgit and Western Tibet, where it must inhabit open country. In 
their zeneral form, and more especially the shoriness of the legs. 
chese animals come nearer to the Jackals than to the Wolves. They 
hunt their prey in packs. Remains of species of this group occur 
in the cavern-deposits of the Continent, and have been described 
under the name of C. curcparus. 

A group tor which the name Ly:alopez has been proposed com- 
prises certain South American Canidex, distinguished from Canis 
proper by their longer tails and Fox-like aspect :—C. eavicrircrus, 
C. brasiliensis, C. melampus, Co retulus, Cl julricaudus, C. azare. C. 
margellanicus, CL ariscus. “The last three have been further separated 
(under the name of PsuJalezz) on account of slizht diterences in 
the relative size of the molar teeth, and of their “pupil being ellip- 
tical when contracted. _Vyefvreutes (one species. C. preeyonides, from 
Japan and North-East Asia) has no claims to generic distinction but 
such as are founded upon its long loose fur, short ears, and short 
bushy tail, vee give it some superficial resemblance to a Raccoon. 

BL dood or Pulpine Serics.—The Mulpine group (T.[pes; 
includes the t true Foxes, of which there are numerous varieties and 
species, spread over North America, Eurasia, and Africa, which 
have been described under the names of (. rulpes (Modpes 0! ae 
the common Fox of saa Ss ; C. niloticus, adustus, and rarieg tis 
Africa: C. fareseens, moniznus, benaalensis, japonicus, eorsie, ‘Asia : 
(. Tulrus, macrurus, velox, North America. Mr. Bl: oe 1 con- 
cludes, however, that the Asiatic (. darzseons and C. monianus, and 
very probably the North American Cross Fox (C. fulrus) are merely 
local races of €. rules, distinguished by certain peculiarities ot 
coloration. The English Fox. measures about 2 feet in length 
exclusive of the tail, which is about a foot long. Its fur is of a 
reddish-brown colour above, and more or less white beneath: the 
back of the ears and the fore part of the limbs are black, and the 
tip of irs bushy tail is white. Its long, sharp muzzle, erect pointed 
ears, and sharp eve. zive it the well- known appearance of sagacity 
and cunning. The Fox is a solitary animal, inhabiting a burrow, 
which it either excavates for itself, or obtains by ejecting the 
badger or the rabbit. So averse, indeed. is the Fox to dig for 
itself, that when foiled in its attempts to dispossess the badger, it 
has been known to take up its quarters with the latter, and it can be 
induced to make its home in artificial burrows constructed of stone 

l Fauna of British India, ‘- Mammalia.” pp. 153. 154 (155s.. 


CANIDAE 553 


and earth for the purpose of facilitating the operation of digging 
out the cubs. The Fox also occurs in woods, and even in the open 
country without burrows, lying in its “cover” by day and stealing 
forth at night in search of its prey. Remains of the Common 
Fox occur not unfrequently in the Pleistocene deposits of Europe. 
The Indian C. bengalensis is a very much smaller and well-marked 
species. ' 

The tail of the above forms is clothed with soft fur and long 
hair, uniformly mixed; from them Baird distinguishes, under the 
name of Urocyon, other species which have a concealed erect mane 
of stiff hairs along the upper line of the tail. These have also a 
shorter muzzle and a wide space between the temporal crests ; they 
are C. virginianus and C-. littoralis, both from North America. The 
Arctic Fox (C. lagopus, genus Leucocyon, Gray) has the tail very full 
and bushy and the soles of the feet densely furred below. Its 
colour changes according to season from bluish-gray to pure white. 

Certain small elegant African Foxes (C. zerda, famelicus, and 
chama), with very large ears and corresponding large auditory 
bulle, have been separated under the name of Mennecus, and are 
commonly known as Fennecs. 

The earliest undoubted occurrence of the genus Canis seems to 
be in the Upper Miocene of Switzerland, where it is represented 
by the Fox-like C. eningensis. In the Upper Pliocene of France 
C. megamastoides is said to be allied to the Foxes and Jackals, but 
with some signs of affinity to the extinct Cynodictis. In the Pliocene 
Siwaliks of India there occurs C’ curvipalatus, of the size of a small 
Fox, which appears to have certain resemblances to Otocyon. 

Lycaon1—This genus resembles in most of its characters the 
Dogs of the Lupine series, but the teeth are rather more massive 
and rounded, the skull is shorter and broader, and there are but 
four toes on each limb, as in Hywna. The one species, L. pictus, the 
Cape Hunting Dog (Fig. 253) from South and East Africa, is very 
distinct externally from all the other Canide. It is nearly as large 
as a Mastiff, with large, broadly ovate erect ears, and singularly 
coloured, being not only variable in different individuals, but un- 
symmetrically marked with large spots of white, yellow, and black. 
It presents some curious superficial resemblances to Hyena crocuta, 
perhaps a case of mimetic analogy. It hunts its prey in large 
packs. <A lower jaw from a cave-deposit in Glamorganshire, which 
‘agrees with that of the existing form in the presence of an anterior 
cusp to the last lower premolar, has been made the type of a dis- 
tinct species (L. anglicus). 

Icticyon.2—The Bush-Dog (JZ. venaticus), from Guiana and Brazil, 
is a species about the size of a Fox, with close hair, and short legs 
1 Brookes, Grifith’s Animal Kingdom, vol. v. p. 151 (1827). 

2 Lund, XK. Danks. Vid. Selsk. Afhand. vol. xi. p. 62 (1845). 


eek CARNIVORA 


and tail, distinguished from all other Dogs by the reduction of the 
molar teeth to 1, and their comparatively small size. The lower 
carnassial is also characterised by the loss of the inner cusp of the 
blade, and the secant form of its hind talon; both these features 
indicating a specialised type. Remains of the Bush-Dog are found 
in the Pleistocene cavern-deposits of Brazil, and were originally 
described under the name of Sjesthos. 

Otocyon.1—Dentition: i 3,¢ 4, p 4+, m Sort total 46 or 48. 
The molar teeth are thus in excess of any other living heterodont © 


Fic. 253.—The Cape Hunting Dog (Lycaon. pictus). 


mammal. They have the same general characters as in Cis, 
with very pointed cusps. The lower carnassial shows little of its 
typical characters, having five cusps on the surface; these can, 
however, be identified as the inner cusp, the two greatly reduced 
and obliquely placed lobes of the blade, and two cusps on the talon. 
The skull generally resembles that of the smaller Foxes, particu- 
larly the Fennecs. The auditory bulle are very large. The hinder 
edge of the mandible has a very peculiar form, owing to the 
great development of an expanded, compressed, and somewhat 
inverted subangular process. Vertebre: C7,D13,L 7,83, C 22. 
Ears very large. Limbs rather long. Toes 5-4. One species, 
O. megalotis, from South Africa, rather smaller than a common Fox. 

Professor Huxley looks upon this as the least differentiated or 
most primitive existing form of the family, regarding the presence 


> 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 <x a 

33 imbs. Pipe se bree fom 
irieu the Middle Miocene of France. 
which, so fir as its teeth are concerned. 
connects mphicwet with the Ursa 
genus Hpenarcius a> 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<roups have probably had a common orizin. 

he Crecdonta as a whole are characterised by the small size 
of the brain, the absence of a single differentiated carnassial tooth, 
and the triangular form or secant character of their upper molars. 
In the carpus the scaphoid and lunar were usually distinct ; the 
femur has a third trochanter; the upper or tibial surface cf the 
astragalus usually wants the zroove found in modern Carniroves : 
and the feet were plantigrade. The curicus resemblance of che 
molars of many of these forms to those of the Marsupials may 
indicate a genetic relationship between the two groups: but, on the 
other hand. the presence of a full set of milk-teeth and the absence 
of palatal vacuities. or of an inflection <i the angle of the mandible. 
sharply distinguishes them from that order. Space permits of a 
notice only of the more interesting forms. 

AHyenodentidz.—This iamily is taken to include some of the 
more specialised types. such as the European and American 
Hucaodon and Oshyrna and the European Pteredei. In Hyenedon 
Fig. 2¢s) the dental formula is i =. » 2. p 4. m 2: the fourth 
premolar above and the first true molar below being formed upon 
the ~ carnassial” plan, tut the teeth behind these, instead of being 
taberculated as in all existing Carnivora, repeat the characters of 
the carnassial. and also increase in size, especially in the lower jaw. 
trom before backwards. The last lower molar differs from the 
two preceding teeth. and is very like the carnassial of Felis, The 
scaphoid and lunar of the carpus were fused tozether. Some species, 
as H. lepiorhynchus, were as large as a Wolf, while cthers did not 
exceed a Fox in size. Pferedin is readily distinguished by having 
wv 3, bv the larger size of the inner tubereles of the upper molars, 
and the similarity in the form of the three lower molars. In some 
species there were only two upper incisors. and the first lower pre- 
molar may be wanting. Chyrna is a specialised form with i =. 
¢ fy; P 4, m 2, and a very long mandibular svmphrsis. 

Provirerridt.—The European and American genus Prorirerra 
(Cynchyriiedan or Supoluphus) may be regarded as representing a 
second family. The dental formula in this genus is the typical i 3, 
cf. p 4. m 3, the upper molars have a large inner tubercle, while 
the lower molars are differentiated into a blade and talon, the 


CREODONTA 609 


blade having a large inner cusp. The upper teeth closely resemble 
the molars of Dasywrus, while the lower molars are like the lower 
carnassial of Cynodictis and Viverra; and thus indicate how the 
Creodonts may have passed into the true Carnivores through the 
extinct Afiacide. 

Arctocyonide and Mesonychide.—The first of these families is 
represented by Arctocyon primevus, one of the oldest known Tertiary 
mammals, from the lowest Eocene beds of La Fere, department of 
Aisne, France, and also by other species from corresponding beds 
at Rheims. The dental formula is 7 3, ¢ 4, p an m 3. The upper 
molars (Fig. 279) are tritubercular, with an incipient postero- 
internal column (hypocone) ; 
the lower are quadrituber- 
cular; and the premolars 
simple. The typical species 
was of large size, but the 
two of which the teeth are 
figured were considerably Fic. 279.—The three right upper molars of Arcto- 
smaller. In the American cyon duelt (a), and the second of A. gervaisi (b); from 


Mesonyx the dental formula the Lowest Eocene of Rheims. pr, protocone ; pa, 
paracone ; me, metacone; hy, hypocone; ml, meta- 


was the typical one, the jaws conule; pl, paraconule. (From Osborn.) 
were comparatively short, the 


mandibular symphysis was elongated, the cheek-teeth were of 
simple structure, and resembled the premolars of many of the true 
Carnivora, and the astragalus had a grooved tibial surface and 
distinct distal facets for the cuboid and navicular, resembling in the 
latter respect the corresponding bone of a Perissodactyle Ungulate. 
The terminal phalanges had deeply fissured extremities, and are said 
to be more like those of Rodents than true Carnivores. A/esonyx 
ossifragus was larger than a Grizzly Bear. Amblyctonus, of the same 
deposits, differs by the smooth tibial face of the astragalus and the 
development of an anterior cusp to the lower molars. 


39 


CHAPTER XII 
THE ORDER INSECTIVORA 


THE Insectivora comprise a number of comparatively small mam- 
mals, generally of terrestrial, although rarely of arboreal or aquatic 
habits, and presenting the following common features. They are 
unguiculate, and have plantigrade or subplantigrade, and generally 
pentadactylate feet, in which the pollex and hallux are not oppos- 
able to the other digits. They are diphyodont and heterodont, and 
the teeth are rooted. The molars are studded with sharp cusps, 
the crowns of the upper molars being either quadrangular or trian- 
gular; there are never less than two incisors in either side of the 
mandible ; and in many cases the incisors, canines, and anterior 
premolars are not clearly differentiated from one another (Fig. 280) ; 
the canines being 
usually weak. 
Clavicles are pre- 
sent, except in 
Potamogale. The 
body is clothed 
with fur or pro- 
tected by an 
armature of 
spines; the 


Fie. 280.—Right lateral aspect of the anterior portion of the 5% . 
cranium of Lrinaceus collaris. Enlarged. (From Dobson, Proc. Zool. tes tes are in- 
Soc. 1881, p. 403.): eninal or placed 


near the kidneys, 
and are not received into a scrotum; the penis is pendent or sus- 
pended from the wall of the abdomen; the uterus is two-horned 
and with or without a distinct corpus uteri; the placenta is dis- 
coidal and deciduate ; and the smooth cerebral hemispheres do not 
extend backwards over the cerebellum (Fig. 281). The projee- 
tion of the muzzle far beyond the extremity of the lower jaw is a 
very general feature. The humerus generally has an entepicondylar 


INSECTIVORA 611 


foramen. Certain forms, such as Talpa and Guleopithecus, are unique 
among mammals in having ossified intercentra in the dorso-lumbar 
region of the vertebral column. 
Representatives of this order are found throughout the temperate 
and tropical parts of both hemispheres ; 
(except South America and Australia), 
and exhibit much variety both in 
organisation and in habits. With the 
exception of the Tupaiide, all are noc- 
turnal; the greater number are cursorial, 
but some (Talpa, Chrysochloris, Oryzorictes) 
are fossorial ; some (Potamogule, Necto- 
gale, Myogale) are natatorial, and a few 
(Tupatide) arboreal; while the species 
of the aberrant genus Guleopithecus glide 
through the air like the Flying Squirrels. 
To the great majority the term insecti- 
vorous is strictly applicable, Galeopithecus 
alone being phytophagous ; while Pota- 
mogale is said to feed on fish, and the 
different species of Moles live chiefly on 
worms. The general organisation of the 
Insectivora indicates a very low type, 
and were it not for the specialised , Fio- 28. Upper surface of the 
character of their placentation and the Geel, Paes sta, oa. 
tendency to lose the differentiated char- 
acters of the anterior teeth they might be regarded as closely 
allied to the ancestral type of many of the heterodont mammals. 
The strongly marked distinction of the canines from the incisors 
and anterior premolars in the Mesozoic and most of the Tertiary 
mammals (excepting some of the Ungulates) points, however, very 
decidedly to the conclusion that the want of definition between 
these teeth in many of the modern Insectivora is an acquired 
feature. Fossil forms apparently indicate a relationship on the 
one hand with the Creodont Carnivora, and on the other with 
the Lemuroid Primates ; indeed it is in some instances impossible 
to say whether'extinct genera are really Insectivores or Lemuroids. 
In most Insectivora the cranial cavity is of small relative size, 
and in none is the brain-case elevated to any considerable extent 
above the facial line. The facial part of the skull is generally 
much produced, and the premaxillary and nasal bones are well 
developed. The zygomatic arch is usually slender or deficient, the 
latter being the case in most of the species; and postorbital pro- 
cesses of the frontals are found only in the Galcopithecide, Tupaiide, 
and Macroscelidide. The number of dorsal vertebre varies from 13 
in Talpa to 19 in Centetes,; that of the lumbar from 3 in Chryso- 


612 INSECTIVORA 


chloris to 6 in Talpa and S22: ; and of the caudal from the rudi- 
mentary series of $ in Centetes to the £0 or more of Mieregale. Not 
less variable are the characters of the vertebrae themselves: the 
spinou: processes often being very lonz in one and shert in another 
species of the same genus. In the + ricide and Mu-gale the neural 
arches of the cervical vertebre are very slender. In the Swristiat 
and Gymnura the four anterior vertebre develop large single hrpa- 


Mvysgale, and Tulpa small oval ossicles are found on the inferior 
surfaces of the lumbar interspaces. In LEvinaceus, owing to the 
thickness of the neural cord in the cervical region and its abrupt 
termination, the diameter of the neural canal in the cervical and 
first two dorsal vertebre greatly exceeds that of any of the succeed- 
ing vertebre. The sternum is variable, but generally narrow, 
bilobate in front, and divided into segments. The pectoral girdle 
presents some remarkable adaptive modifications, most fully ex- 
pressed in July, having relation to the use cf the jore limbs in 
burrowing; but in the Golden Moles (Cirvscchloris) the forearm 
and manus alone become specially modified ior this purpose. In 
Galespithecus and Muacrescelides the bones of the forearm (radius 
and ulna) are distally united. The manus has generally five dizizs, 
but in Fé:nchocyon and in one species of Orvzrictzs the pollex is 
wanting, while in the true Moles it is extremely modified. The 
femur has, in most species, a prominent ridge below the greater 
trochanter representing a third trochanter. In Galepithecus, Zupaia, 
Centetes, Hemicentetes, Friculus, and Sslzncdon the tibia and fibula 
are distinct, but in all the other genera more or less united 
together. The pes usually possesses five digits (rarely four by 
reduction of the hallux); and in some forms, as in the leaping 
species (Macreseelides, Rhunehecvon), the tarsal bones are greatly 
elongated. The form of the pelvis. and especially of the sym- 
physis pubis, varies within certain limits: and these differences 
have been proposed by Leche as a basis for the classification cf the 
families. Thus in the Gulespithecida, Tupaiide, and Maercse-lidide 
there is a long symphysis; in the Erinaccidx, Ceat:tide, and P<tame- 
galide the symphysis is shorz: and in the S-ncidz, Talpide, and 
Chrusochloride there is none. 

Space does not admit of attempting a sketch of the modifica- 
tions of the muscular system, which will be found fully described 
in Dr. Dobson's Wenearaph, referred to in the biblicgraphy. As to 
the nervous system, it has been already mentioned that the brain 
throughout the order presents a low type of organisation; in none 
ct the members do the cerebral hemispheres present any trace of 
convolutions, nor do they extend backwards so as to cover the 
cerebellum, while the olfactory lobes are large and project in front, 


INSECTIVORA 613 


and the corpus callosum is short and thin. In the Hedgehogs 
(Erinaceus) the spinal column ends abruptly opposite the third or 
fourth dorsal vertebra in a slender filament, and the dorsal and 
lumbar nerves, given off in front of this point, are carried back- 
wards in two compressed bundles occupying the suddenly narrowed 
spinal canal as far as the sacrum. 

Owing to the similarity in the character of the food, the truly 
insectivorous species, forming more than nine-tenths of the order, 
present little variety in the structure of their digestive organs. 
Except in Guleopithecus the stomach is a simple, thin-walled sac ; 
but in some, as in Centetes and allied genera, the pyloric and 
cesophageal openings are very close together. The intestinal canal 
has much the same calibre throughout, and varies from three (in 
the Shrews) to twelve times (in the Hedgehogs) the length of the 
head and body. In the arboreal genera, Galeopithecus and Tupaio, 
as well as in the Macroscelidide, all of which probably feed in 
part on vegetable substances, most of the species possess a cecum. 
The liver is deeply divided into lobes, the right and left lateral 
being cut off by deep fissures ; and both the caudate and Spigelian 
lobes being generally well developed. The gall-bladder, which is 
usually large and globular, is placed on the middle of the posterior 
surface of the right central lobe. 

In most of the members of the order (Soricide, Centetide, Chryso- 
chloride) the penis is capable of being more or less completely 
retracted within the fold of integument surrounding the anus; in 
some (Galeopithecid, Tulpide) it is pendent in front of the anus ; 
while in others (Mucroscelidide, Hrinaceide, Solenodontide) it is 
carried forwards and suspended from the abdominal wall. In the 
subfamily Centetine and Chrysochloris the testes lie immediately 
behind the kidneys, but in others more or less within the pelvis. 
During the rutting season they become greatly enlarged, forming 
protrusions in the inguinal region. Except in Lhynchocyon the 
uterine cornua are long and open into a short corpus uteri, which 
in many species (Soricide, Talpide, Centetide, Chrysochloride) is not 
separated from the vagina by a distinct os uteri. With the 
exception of Galeopithecus all Insectivora appear to be multiparous, 
the number of young at a birth varying from two to eight in 
Evrinaceus, and from twelve to twenty in Centetes. The position 
of the mammary glands and the number of the teats vary greatly. 
Thus in Galeopithecus there are two pairs of axillary teats, and in 
Solenodon a single post-inguinal pair ; but in most species they range 
from the thorax to the abdomen, varying from two pairs in Gymnura 
to twelve in Centetes. In Chrysochloris the thoracic and inguinal teats 
are lodged in deep cup-shaped depressions. 

Odoriferous glands exist in many species. In most Shrews 
these glands occur on the sides of the body at a short distance 


614 INSECTIVORA 


behind the axilla, and their exudation is probably protective, since 
few carnivorous animals will eat the dead bodies of these creatures. 
In both species of Gymnura and in Potumogule large pouches are 
situated on either side of the rectum and discharge their secretions 
by ducts, opening in the first-named genus in front of, and in the 
latter within the margin of the anus. In Centetes the ducts of 
similarly situated racemose glands open by pores at the bottom of 
deep pits placed at either side of the anus. 

The integument is thin, but in many species is lined by a 
muscular coat, which is probably more developed in the Hedge- 
hogs (Lrinaceide) than in any other mammal. In this family 
and the Centetide most of the species are protected by spines 
implanted in the panniculus carnosus muscle, and more or less 
replacing the fur of the upper surface of the body. 

The order is usually divided into two suborders, but the very 
aberrant genus which constitutes the first might well be raised to 
ordinal rank. It has little in common with the true Insectivora, 
but as it certainly belongs to no other of the recognised mammalian 
orders it is retained among them chiefly to avoid the inconvenience 
of increasing the number of ordinal divisions for the sake of a 
single isolated form. 


Suborder DERMOPTERA. 


Upper and lower incisors compressed, multicuspidate, the lower 
deeply pectinated; fore and hind limbs connected by a broad 
integumentary expansion forming a parachute. 


Family GALEOPITHECID®. 


In addition to the characters given under the head of the sub- 
order it may be mentioned that the orbit is nearly surrounded by 
bone, the zygomatic arches are well developed, the tympanic forms 
a bulla, the ulna is distally united with the radius, the tibia and 
fibula are distinct, the pubic symphysis is long, the penis is pendent, 
the testes are received into inguinal pouches, the mamme are 
axillary, the uterus is two-horned, and there is a large cecum. 

Galeopithecus \—Dentition : 1 3,c¢4, p 2, m2; total 34. Second 
upper incisor and canine with two roots. Two species—G@. vlans 
and G. philippinensis. The former, which is distinguished from the 
latter by the form of the upper incisors, has a total length of nearly 
2 feet. The long and slender limbs are connected by a broad 
integumentary expansion extending outwards from the sides of the 
neck and body, and forming also a web between the fingers and 
toes as far as the base of the claws (Fig. 282); the hind limbs are 


1 Pallas, deta Acad, Sei. Imp. Petropolis, vol. iv. pt. 1, p. 208 (1780). 


GALEOPITHECID.£ 615 


further connected by a similar expansion passing outwards along 
the back of the feet to the base of the claws, and, inwardly, involy- 
ing the long tail to the tip, forming a true interfemoral membrane, 
as in the Bats. 

The two species of Flying Lemurs, as the representatives of this 
genus are commonly but erroneously called, live in the forests of the 


Fic, 282.—Feet of Galeopithecus philippinensis. 


Malay Peninsula, Sumatra, Borneo, and the Philippine Islands, where 
they feed chiefly on the leaves and fruits of trees. Their habits are 
nocturnal, and during the daytime they cling to the trunks or limbs 
of trees, head downwards, in a state of repose. With the approach 
of night their season of activity commences, when they may be 
seen gliding from tree to tree supported on their cutaneous 
parachute, and they have been observed to traverse in this way a 
space of 70 yards with a descent of only about one in five. 
Galcopithecus was referred by some of the older zoologists and 
anatomists to the Bats, and by others to the Lemurs, but Professor 
Peters’s view, that it belongs to neither of these orders, and should 
be considered an aberrant Insectivore, has been very generally 
accepted, although, as mentioned above, the association is by no 
means a close one. Besides differing from the Bats in the form of 
the anterior limbs and of the double-rooted outer incisor and canine, 
it also contrasts strongly with them in the presence of a large sac- 
culated cecum, and in the great length of the colon, which is so 
remarkably short in all the Chiroptera. From the Lemurs, on the 
other hand, the form of the brain, the characters of the teeth, the 
structure of the skull, and the deciduate discoidal placenta com- 
pletely separate it. In a recent elaborate memoir on the myology 
and affinities of Galeopithecus Dr. Leche! considers that we have in 
this genus an indication of the mode in which the Insectivora were 
modified into the Chiroptera, although it is completely off the direct 


1 Ueber die Stiugethiergattung Galeopitheeus. Sv. Ak. Handi, vol. xxi. pt. xi. 
(1886). 


616 INSECTIVORA 


line of descent. The deeply pectinated crowns of the lower incisors 
of Galcopithecus are quite unique in the class, and the only approach 
to the double-rooted canine, except in Hrinaceus and Talpa, is found 
among the Marsupials in Perameles, where the root of the canine is 


grooved. 


Suborder INSECTIVORA VERA. 


Upper and lower incisors conical, unicuspidate or with basal 
cusps only, the lower not pectinated ; limbs free, formed for 


terrestrial progression. : 
The following table gives a key to the distinctive characters of 


the existing families :— 


I. Upper molars broad, multicuspidate, with more or less well-defined 
W-shaped crowns. 
A. Symphysis pubis long ; generally a caecum ; cerebral cavity 
comparatively large. 
a, Orbit encircled by bone ; metatarsus moderate ; arboreal. 
Tupaude. 
b. Orbit not encircled by bone ; metatarsus greatly elongated ; 
terrestrial. Macroscelidide. 
B. Symphysis pubis short or none; no cacum ; cerebral cavity 
small ; skull without postorbital processes. 
a, First and second upper molars with a central fifth cusp. 
a’. Tympanic annular, not forming a bulla. Erinaceide. 
b. No central fifth cusp to upper molars. 
a’. Tympanic annular, not forming a bulla; no zygomatic 
arch. Soricide. 
b’. Tympanic forming a bulla; zygomatic arch developed. 
Talpide. 
Il. Upper molars narrow, with V-shaped crowns. 
a. Tympanic annular, not forming a bulla; zygomatic arch 
imperfect. 
a’, No clavicles. Potamogalide. 
b”. Clavicles well developed. 
a”. Skull constricted between the orbits; penis sus- 
pended. Solenodontide. 
b”. Skull not constricted ; penis pendent, retractile. 
Centetide. 
¥. Tympanic forming a bulla ; zygomatic arch well developed. 
Chrysochloride. 


The second section, in which the molars are of the primitive 
tritubercular type, should probably be regarded as containing the 
most generalised representatives of the order ; and it is noteworthy 
that the whole of them are confined to Africa, Madagascar, and 
the West Indies, whereas most of the first section are widely 
distributed over the Palzarctic and Oriental regions. None of the 


TUPAUDE 617 


existing families of the second section are known ina, fossil condition, 
although it is suggested that the extinct Leptictide includes allied 


types. 
Family TUPAUD.E. 


Skull with comparatively large brain-case, orbit surrounded by 
bone, well-developed zygomatic arch, perforated jugal, and a tympanic 


Pia. 288.—The Pentailed Tree-Shrew (Ptilocercus low’). From Gray, Proc. Zool. Soc. 1848. 
$ natural size. 


bulla. Upper molar broad, with cusps arranged ina W. Pubic 
symphysis long; radius and ulna, and tibia and fibula separate ; 
metatarsus only slightly longer than tarsus. Usually a short ecxecum. 
Habits arboreal and diurnal. Confined to the Oriental region. 
Tupaiu.—Dentition : i 2, ¢ 4, p 8, m 35 total 38. Feet naked 
beneath, the sole furnished with projecting pads; claws moderate, 
curved, and sharp; head pointed; ears rounded; tail bushy, 
distichous, with short hair below. The Tree-Shrews, of which there 
are some nine species, are found in India, Burma, the Malay 
Peninsula, the Nicobars, Sumatra, Java, and Borneo. The species 
closely resemble one another, differing chiefly in size and in the 
1 Rafiles, Trans, Linn. Soc. vol, xiii, p. 256 (1822). 


618 INSECTIVORA 


colour and length of the fur. Their general appearance is very 
Squirrel-like. Their food consists of insects and fruit, which they 
usually seek in the trees, but also occasionally on the ground. 
When feeding they often sit on their haunches, holding the food, 
after the manner of Squirrels, between their fore-paws. 

Pttlocercus..— Represented only by the Pentailed Tree-Shrew 
(P. lowi, Fig. 283) of Borneo, in which the tail is of extraordinary 
length, with the proximal two-thirds naked, and the remaining third 
furnished witha bilateral fringe of long hairs, from which the genus 
takes its name. 

Extinct Genera.—An Insectivore from the Middle Miocene of 
France, described as Lantanotherium, is said to be nearly allied to 
Tupua. The genus Parasorex, from strata of similar age, has the 
dental formula i 3, ¢ 4, p 4, m 3, and is regarded as connecting the 
present with the following family. 


Family MACROSCELIDID. 


Skull with comparatively large brain-case, strong zygomatic 
arch, a tympanic bulla, orbit surrounded by bone, imperforate 
jugal, and usually no postorbital process. Molars broad, with 
four cusps arranged in a W. Pubic symphysis long; proximal end 
of tibia and fibula united; radius and ulna united or separate ; 
metatarsus much longer than tarsus. A large cecum. Habits 
terrestrial, saltatorial, and nocturnal. The family is confined to 
Africa. 

Alacrascelides.°-—Dentition : ¢ 3, ¢ 4, p 4, m x3 ; total 40 or 42. 
Distal extremity of radius and ulna united. Five digits in manus, 
and five or four in pes. This genus, which is taken to include 
Petrodromus, comprises ten species widely distributed throughout the 
African continent. All are closely related, resembling one another 
in general form, and even in the colour of the fur. They fall into 
two groups, distinguished by the presence or absence of a small 
third lower molar.* JZ. tetradactylus (Fig. 284), the type of the 
genus Petrodromus, differs from all the other species in the absence 
of the hallux, and of the third lower molar. These animals are 
commonly known as Jumping Shrews, and, like the following 
genus, have the muzzle much produced. 

Lthynchocyon.A—Dentition : i 3, ¢ 4, p 4, m 2; total 36. Upper 
incisor frequently shed in the adult. Radius and ulna distinct ; 


1 Gray, Proc. Zool. Soc. 1848, p. 23. ° Andrew Smith, S. African Quart. 
Journ, vol. ii. No. 1, p. 64 (1833). 

° The above correct formula of the dentition of this family has been recently 
worked out by 0. Thomas, Proc. Zool. Soc. 1890, pp. 445, 446. 

4 Peters, Bericht k. preuss. Ak. Wiss. 1847, p. 36. 


ERINACEIDE 619 


hind limbs relatively shorter, and proboscis longer than in the type 
genus ; four digits in each foot. Four closely allied species have 
been described from East Africa. The head and body of the type 


Fig. 284.—Maeroscelides tetradactylus. x4. (From Peters, Reise nach Mossambique.) 


species measures about & inches in length; and the long tail is 
covered with a ringed skin, sparsely haired. Its habits are fossorial. 


Family ERtS ACEID.£. 


Skull with a small brain-case ; no postorbital process ; slender 
and occasionally imperfect zygomatic arch, and an annular tympanic, 
which does not form a bulla. Upper molars with four principal 
cusps and a small central median cusp. Acromion of scapula bifid ; 
pubic symphysis short; radius and ulna free, but tibia and fibula 
united proximally. No cecum; penis carried forward and sus- 
pended from the wall of the abdomen. Habits terrestrial. Found 
in the Palearctic, Ethiopian, and Oriental regions. 

Subfamily Gymnurine.—Palate completely ossified ; pelvis 
very narrow ; fur without spines. 

Gymnura.—Dentition: ¢ 3, ¢ 4, p 4, m 3; total 44. This 
genus, if Hylomys is rightly included, is represented by the two 

1 Horsfield and Vigors, Zool. Journ. vol. iii, p. 246 (1828). 


620 INSECTIVORA 


species, G. raffest and G. suilla, from the Malay Penisula and Indian 
Archipelago. The former has the appearance of a large Rat with a 
long tail and head and projecting mobile snout ; the latter, which is 
much smaller, with a short tail and small third upper premolar, has 
long been known under the name of Hylomys suillus, and classed 
with the Tupatide. Both species present a very generalised type 
of dentition, in this respect occupying an almost central position in 
the order. G. suilla is represented in Mount Kina-Balu, Borneo, by 
a variety characterised by the presence of a dark dorsal streak. 
Many zoologists prefer to retain Hylomys as a distinct genus. 

Subfamily Erinaceine.—Palate imperfectly ossified; pelvis 
wide ; fur with spines. 

Erinaceus.\—Dentition : i 3, ¢4,p 3, m3; total 36. The first pair 
of upper incisors (Fig. 285) are considerably larger than the others, 
and are widely 
separated from one 
another in the 
middle line; the 
canine is very simi- 
lar to the third in- 
cisor ; and, except 
in E. europeus (Fig. 
285), each of these 
\ teeth is inserted by 
\i two distinct roots 
(Fig. 280, p. 610). 

Fia. 285.—Right lateral aspect of the anterior portion of the The first lower in- 
Dobson, Pro, Zoek oe. 1851,p- 408) Te Pm cisor is large and 
proclivous. The 

number of vertebra is C 7, D 15, L 6, 8 3, © 11. 

The Hedgehogs comprise nearly twenty species, distributed 
throughout Europe, Africa, and the greater part of Asia, but not 
found in Madagascar, Ceylon, Burma, Siam, the Malay Peninsula, 
or Australia. All the species resemble one another in the armature 
of spines investing the upper surface and sides of the body ; and 
all possess the power of rolling themselves up into the form of 
a ball, protected on all sides by the strong spines; the dorsal 
integument being brought downwards and inwards over the head 
and tail, so as to include the limbs also, by the action of special 
muscles. The common Hedghog (£. europeus) is the most aberrant 
species, differing from all the rest in the peculiarly shaped and 
single-rooted third upper incisor and canine (Fig. 285), and in its 
very coarse, harsh fur. The dentition of the long-eared North 
Indian form, £. collaris (Fig. 280), may be considered characteristic 
of all the other species, the only important. differences being found 

* Linn, Syst. Nat. 12th ed. vol. i. p. 75 (1766). 


SORICIDAL 623 


in the variable size and position of the second upper premolar, 
which is very small, external, and deciduous in the Indian 
E. micropus and pictus. The former species, limited to South India, 
is further distinguished by the absence of the jugal bone. Of the 
African species, E. diadematus, with long frontal spines, is probably 
the commonest; while EF. albiventris has been made the type of a 
separate genus on account of the total absence of the hallux. 

The well-known European species feeds on insects, worms, slugs, 
mice, rats, lizards, snakes, etc., as well as on eggs, fruit, and roots. 
It hibernates during the winter. The young are usually produced 
in July or August in litters of not more than four, but there may 
be a second litter in October; and the period of gestation is be- 
lieved not to exceed a month. The Indian, and probably also the 
African species, do not hibernate. 

The existing 7. ewropeus dates from the Pleistocene period, and 
extinct species of the genus are found in the Upper and Middle 
Miocene of the Continent. 

Extinct Genera.—The French Lower Miocene genus, Paleoerin- 
aceus, appears to be allied to Hrinaceus, but is distinguished by the 
wider and completely ossified palate. In the Upper Eocene of 
Central France there are two genera, which appear to be most 
nearly allied to Gymnura, although connected by Palewoerinaceus with 
Erinaceus. Of these Necrogymnurus,! with which Cayluxotherium is 
apparently identical, has teeth like Gymnwra, but an imperfectly 
ossified palate like Erinaceus ; and the skull is remarkable for the 
peculiar rugose structure of the parietal and temporal regions. 
Comphotherium is distinguished by the presence of a cingulum to 
the lower molars, like that found in Gynmura. 


Family SorRiciDz&. 


Skull (Fig. 286) long and narrow, with no zygomatic arch or 
postorbital process, and the tympanic ring-like and not forming 
‘a bulla. Upper molars with the cusps arranged in a distinct W. 
No pubic symphysis. The tibia and fibula united. No cecum. 
Habits usually terrestrial, rarely aquatic. Distribution extensive. 

The Shrews are Rat-like or Mouse-like insectivores, with the 
body covered with hair, and the muzzle long and pointed. Their 
dentition (Fig. 286) is peculiar and characteristic. Thus the first 
upper incisor is large and hook-like, with a more or less developed 
basal cusp on the posterior border. Between this and the last pre- 
molar there are a variable number of small teeth, representing the 
other incisors, the canine, and the anterior premolars; although, 
owing to the early obliteration of the maxillo-premaxillary suture, 


) Originally given incorrectly as Vewrogymnurus. 


622 INSECTIVORA 


their homology is exceedingly difficult to determine. Three molars 
are invariably present, of which the third is much the smallest. In 
the mandible there are always six teeth, but in one species of 
Myosorex there may be a seventh. 
The first lower incisor is usually 
directed horizontally forwards; the 
second incisor (formerly reckoned as 
the canine) is the smallest tooth of 
the series, the fourth premolar being 
slightly larger. 

This family, which includes con- 
siderably more than half the re- 
presentatives of the order, has a 
distribution coextensive with the 
latter. Many classifications of this 
ait, Sante ey fe ical group have been attempted 
Te the cheatin, first incisor ; ¢, fost but according to the latest proposal 
incisor; p, canine; m, fourth premolar: of Dr. Dobson,1 the genera may be 
in the mandible—i, first incisor; c, second divided into two subfamilies, dis- 
incisor; p, fourth premolar; m,firstmolar. ~! x Clase 
(From Alston, Proc. Zool. Soc. 1877.) tinguished by the apparently trivial 

character of the colour of the teeth. 

Subfamily Soricinze.—Summits of the teeth coloured red. 

Sorex.2—Dentition : i 4,¢4, p 2, m %; total 32. Openings of 
male and female generative organs separated from the anal orifice ; 
penis cylindrical or tapering; ear well developed; tail long, 
covered with equal or subequal hairs. 

It has been shown by Brandt that the position of the pre- 
maxillo-maxillary sutures in the type of the genus is between the 
fourth and fifth tooth, so that it appears that we must regard this 
genus as differing from all other Eutherian mammals in having four 
upper incisors. Dr. Dobson, in his paper quoted, classes the tooth 
here reckoned as the upper canine with the premolar series in all 
the Shrews. Habits terrestrial. Species numerous, inhabiting the 
Palearctic and Nearctic regions. 

Of the two species found in the British Isles the Common 
Shrew (S. vulgaris, Fig. 287) is by far the most common in England, 
and is about the size of the House Mouse, to which it approximates 
in general form. The body is clothed with close long fur, very 
soft and dense, and varying in colour from light reddish to dark 
brown above, rarely speckled or banded with white. The under 
surface of both the body and the tail is grayish. The basal four- 
fifths of all the hairs above and beneath are dark bluish-gray ; the . 
hairs of the tail are less densely set and coarser. On each side of 
the body, at a point about one-third of the distance between the 
elbow and the knee, may be found, especially in the rutting season, 


1 Proc. Zool. Soc. 1890, p. 49. * Linn. Syst. Nat. 12th ed. vol. i. p. 73 (1766). 


SORICIDE 623 


a gland covered by two rows of coarse hairs. This secretes a 
peculiar fluid, on which the odour of the animal depends; this 
odour being evidently protective, and rendering the creature secure 
against the attacks of many predaceous animals. 

The geographical range of the Common Shrew is exceedingly 
-wide, extending eastwards through Europe and Asia (north of the 
Himalayas) to North America. 

The Lesser Shrew (8. pygmeus') is far less common in England 
and Scotland, although more abundant in Ireland, where S. vulgaris 
is unknown. It is distinguished from the latter not only by its 
inferior dimensions, but also by the circumstance that the third 
upper incisor is not longer than the fourth, and by the considerably 
shorter length of the forearm and manus. ‘This species extends 
through Europe and Asia as far as the island of Saghalin. Both 


Fic. 287.—The Common Shrew (Sorex vulgaris). 


this and the preceding species generally live in wooded districts, 
making their nests under the roots of trees, or in slight hollows. 
The great mortality noticeable among the Shrews in the early part 
of the autumn is probably due to insufficiency of food. The breed- 
ing season extends from the latter part of April to the beginning 
of August. The young, which are blind, naked, and toothless at 
birth, are very quickly developed. The number in a litter is 
usually from five to seven, but may be as many as ten. 

The Alpine Shrew (S. alpinus), which is restricted to the Alpine 
region of Central Europe, is slightly larger than the common 
species, from which it is distinguished by the longer tail, the length 
of which exceeds that of the head and body, by the fur being dark 
on both surfaces of the body, and also by the larger size of the 
upper canine. 

In North America S. bendirei is by far the largest species of the 
genus; and, as in many other species of the same country, the 
fourth upper incisor is relatively small. In S. hoyi (separated by 


1 Syn. S. minutus. 


624 INSECTIVORA 


some writers as Ificrosorer), of the same country, this tooth is 
rudimentary. 

Other North American Shrews, which are regarded by some 
zoologists as generically distinct under the name of Neosorez, are 
aquatic, and thus take the place of the Old World genus Crossopus. 
These are 8. palustris of the Rocky Mountains and S. hydrodromus of 
Unalaska Island, both of which resemble Crossopus in having the 
feet provided with swimming fringes, but agree with the other 
species of Sorex in their dentition and the character of the tail. 
The former species is about the size of Crossopus fodiens, while the 
latter is scarcely larger than S. pygmeus. 

Soriculus..Dentition : i #, ¢ 3, p , m %; total 30, or rarely 
32. Opening of male or female generative organs forming with the 
anal orifice a shallow cloaca. Ear and tail asin Sorex. First upper 
incisor with an internal cusp. Habits terrestrial. 

This genus is the only representative in the Oriental region of 
the Soricine, which are otherwise confined to the Palearctic and 
Nearctic regions. The Indian and Burmese species comprise 
S. nigrescens, S. caudatus, and S. macrurus. 

Notiosorea.?—Dentition: i 3, ¢ 3, p 3, m 3; total 28. Tail 
moderate ; first upper incisor without an inner cusp; other 
characters as in Sortculus. Habits terrestrial. 

This American genus is represented by S. crawfordi and S. evotis, 
which are found in Central America and Mexico, and are thus some 
of the most southerly representatives of the Shrews in that con- 
tinent. Their external appearance is very similar to that of the 
Old World genus Crocidura. 

Blarina.2—Dentition : 2 SB; C4, Py ms; total 32 or 30. Ear 
truncated above; tail short; otherwise as in Soriculus. This group of 
so-called Earless or Short-tailed Shrews is mainly North American, the 
common forms being B. dekayi and B. brevicauda. The species vary 
considerably in size; and B. mevicana and micrura extend the 
range of the genus into Mexico and Guatemala. The following 
account of the habits of B. brevicaudu is taken from Dr. Merriam’s 
Mammals of the Adirondack Region: “The rigours of our northern 
winters seem to have no effect in diminishing its activity, for 
it scampers about on the snow during the severest weather, 
and I have known it to be out when the thermometer indi- 
cated a temperature of - 20° Fahr. It makes long journeys 
over the snow, burrowing down whenever it comes to an 
elevation that denotes the presence of a log or stump, and I am 
inclined to believe that at this season it must feed largely upon 


1 Blyth, Journ. As. Soc. Bengal, vol. xxiv. p. 36 (1855). 2 Coues, Bull. 
U.S. Geol. Surv, Terrs. vol. iii. p. 646 (1877). ® Gray, Proc. Zool. Soc. 
1837, p. 124, 


SORICIDAE 625 


the chrysalides and larve of insects that are always to be found in 
such places.” Dr. Merriam has made the interesting discovery 
that the common short-tailed North American Shrew supplements 
its insectivorous fare by feeding on beech-nuts, which will account 
for the generally very worn state of the teeth in this species. 

Crossopus.—Dentition : i 3, ¢ 4, p 2, m 3; total 30. Opening 
of male or female generative organs enclosed within the same ring 
as the anal orifice ; penis broad, with lateral processes. ars small, 
not truncated. Tail long, with an inferior fringe of elongated 
hair ; feet also fringed. Habits aquatic. The Palearctic Water- 
Shrew (C. fodiens) is considerably larger than the Common Shrew, 
from which it is readily distinguished externally by its shorter and 
much broader muzzle, comparatively smaller eyes, and larger feet 
adapted for swimming,—the sides of the feet and toes being pro- 
vided with comb-like fringes of stiff hairs. The tail is longer than 
the body, and possesses a well-developed swimming fringe of 
moderately long, regularly arranged hairs, which extend along the 
middle of the flat under surface from the end of its basal third to 
its extremity. The fur of the body is long and very dense, varying 
much in colour in different individuals, and this has given rise to 
descriptions of many nominal species; the prevailing shades are 
dark brown, almost black, above, and more or less bright ashy 
tinged with yellowish beneath ; sometimes in the same litter there 
are individuals with the under surface more or less dark coloured. 
In the. number as well as in the shape of the teeth the Water- 
Shrew differs from the Common Shrew: there is a premolar 
less on each side above; the bases of the teeth are much more 
prolonged posteriorly ; and their cusps are much less stained brown, 
so that in old individuals with worn teeth they often appear alto- 
gether white. This species resembles the otter in its aquatic 
habits, swimming and diving with great agility. It frequents 
rivers and lakes, making its burrows in the overhanging banks, 
from which when disturbed it escapes into the water. Its food 
consists of insects and their larve, small crustaceans, and probably 
the fry of small fishes. It is generally distributed throughout 
England, is less common in Scotland, and as yet it has not been 
recorded in Ireland ; specimens have been obtained from many parts 
of Europe, and also from Asia as far eastward as the Altai Mountains. 

Subfamily Crocidurinze.—Teeth completely white. 

Myosorex.2—Dentition : i 3, ¢ 3, p <5, m %; total 30 or 32. 
Penis cylindroid and tapering; male or female generative organs 
opening close to anal orifice, but not forming a cloaca. Ears well 
developed ; tail long, clothed with equal or subequal hairs. Habits 
terrestrial. 

This genus is typically represented by M. varius, a very small 

1 Wagler, Zsis, 1832, p. 275. 2 Gray, Proc. Zool. Soc. 1837, p. 124. 
40 


626 INSECTITORA 


Shrew from the Cape, which is quite unique among the whole 
family in having a rudimental seventh pair of lower teeth. 

Crocidura..—Dentition : 7 3, ¢ 4, p SS m %; total 28 or 30. 
Male or female generative organs forming a short cloaca with the 
anal orifice. Tail long, with a mixture of long and short hairs. 
Other characters as in J/yossrev. Habits terrestrial. 

This Old World genus includes over seventy nominal species, 
which have been divided into four subgenera, (. aranea and C. 
suaveolens of Continental Europe, and ¢’. cerulea of India, being well- 
known forms. The species are very variable and difficult to dis- 
criminate. C. aranea has a very wide distribution, ranging from 
Central and Southern Europe to North Africa and Central Asia. 
The name Musk-Rat is popularly applied in India to C. ce@rulea, 
which frequents houses at night, hunting round rooms for cockroaches 
and other insects, and occasionally uttering a sharp shrill cry. The 
strong musky odour of this animal arises from large glands situated 
beneath the skin of the side of the body, a chars distanes behind 
the fore limbs. This odour is so powerful and penetrating that it 
is popularly believed in India that if the animal runs over a corked 
bottle of wine or beer it will infect the fluid within. Jerdon says 
that certainly many bottles are met with quite undrinkable from 
the peculiar musky odour of their contents, but, rejecting the 
possibility of its passing through the glass, he attributes it to 
the corks having been infected previously to bottling, stating in 
corroboration of this view that he has never found the odour in 
liquors bottled in England. 

Liiplomesodon.°—Dentition: i 3, ¢ 3, p 3, m 2: total 26. Tail 
moderate ; soles of the feet hairy. Other characters as in Crocidura. 
Habits terrestrial. 

This genus is represented only by D. pulchellus of the Kirghiz 
steppes, which is allied to the following form, although retaining 
the normal Shrew-like external contour. 

<fnurosorer.3—Dentition: i 2, ¢ 3, p 3, m 3: total 26. Ear 
very short ; tail rudimental or short; soles of feet naked. Other 
characters as in Diplomesodon. 

The two species of this genus are Mole-like terrestrial forms, of 
which the typical 1. sqguamipes occurs in Tibet, while f. assamensis 
is found in Assam. The latter species has the longer tail. The 
habits of both are probably fossorial. 

Chimarrogale.»—Dentition : i 3, ¢ 4, p 3, m 8; total 28. Penis 
broad, with lateral processes; male or female ‘generative organs 
opening within the same integumentary ring as the anal orifice. 


1 Wagler, Jsis, 1832, p. 278, ? Brandt, in Lehmann’s Reise.-Zool. Anh. 
p. 299 (1852). 3 Milne-Edwards, Compt:s Rendus, vol. xx. p. 341 (1870). 
* Anderson, Journ. As, Soc, Bengal, vol. xlvi. p. 282 877). 


SORICID.E. 627 


Tail long, with an inferior fringe of clongated hairs; cars small ; 
plantar callosities simple; toes free. Habits aquatic. 

This genus includes (. imulayica of the Himalaya and CL platy- 
cephalus of Japan, Both have the feet fringed, and, together with 
the next genus, may be regarded as the eastern analogues of (ruxso- 
pus among the ved-toothed series ; their structural resemblances to 
the latter, if Dr. Dobson’s classification is a natural one, being 
probably due to adaptation for a similar mode of life. 

Nectogale\—Dentition : 48,64, p4, m3; total 28. External 


Fia, vss.—Nectogale cleyans. (From Milne-Kdwards, Mammif. Tibet.) 


curs not forming a conch, valvular, Plantar callosities forming 
adhesive pads; toes webbed. Other characters as in Chimarroqule, 
Habits aquatic. 

The sole representative of this genus is the Tibetan Water- 
Shrew GY. elegans, Fig. 288), which differs from all other members 
of the family by the webbed toes and the presence of the disc-like 
adhesive pads on the under surface of the feet, which are believed 
to enable the creature to hold on to smooth rocks or stones in the 
heds of the streams it inhabits. This species is probably more 
completely aquatic in its habits than the allied Chimarroqule. 

Fossil Sorividie,—Remains of existing species of Soree or Crosso- 
pus ocent in the Norfolk Forest bed, while an extinet species has 
heen found in the Pleistocene of Sardinia. (recidura occurs in the 
eavern-deposits of Madras. Shrews from the Miocene and Upper 


! Milne-Edwards, Comptes Rendus, vol. xx. p. 341 (1870). 


628 INSECTIVORA 


Eocene of Europe have been referred to Surex and the genus Amphi- 
sorer, which is a synonym of C'rassopus. 


Fumily TALPIDE. 


Allied to the Svricide, but distinguished by the presence of a 
zygomatic arch and auditory bulla in the skull, and by the form of 
the teeth. The eyes are very small, and in some species covered 
with skin; the ears are short and concealed by the fur; the fore 
limbs are generally more or less modified for digging; there is no 
symphysis pubis; the intestine has no czecum; the tibia and fibula 
are united; and the unicuspidate first upper and lower incisors 
are not extended horizontally forwards. 

This family is connected with the Suricide by Urotrichus and 
Uropsilus. All the members are limited to the temperate regions 
of Europe, Asia, and North America; and the majority of them 
are of fossorial habits, although a few are aquatic or cursorial. The 
family has been divided into two subfamilies by Professor Mivart, 
and since this arrangement has been very generally adopted it will 
be followed here. From the presence of intermediate forms like 
Scaptonyz Dr. Dobson, in the second part of his JJonogruph of the 
Insectivora, has proposed a different arrangement, which, with the 
omission of some forms which are of not more than subgeneric 
value, is as follows :— 

MyocaLe—Myogale, 

CoyprLuRzZ— Condylura. 
Scapanus. 
Scalops. 
Tatpa—Talpa. 
Scaptonyx. 
Crotrichus. 
Unroprsit1— Uropsilus. 


SCALOPES { 


UROTRICHI { 


Subfamily Myogalinge.—Clavicles and humerus moderately 
elongated ; manus without falciform bone. : 

ALyogale.|— Dentition: i 3, ¢ 1, p 4,.m 3; total 44. Feet 
webbed. Habits aquatic. This genus is represented by the two 
species Jf. moschata (Fig. 289) and J. pyrenaica, of which the former 
is by far the largest member of the family, its total length being 
about 16 inches. Its long proboscis-like snout projects far beyond 
the margin of the upper lip; the toes are webbed as far as the bases 
of the claws ; and the long scaly tail is laterally flattened, so as to 
form a powerful instrument of propulsion when swimming. This 
species inhabits the banks of streams and lakes in South-East Russia, 
where its food consists of various aquatic insects. JL pyrenaica, 


* Cuvier, “Tabl. de Classif.” in Lecgons @’ Anat. Compar. vol. i. (1800). 


TALPID.E 629 


living in a similar manner in the region of the Pyrenees, is very 
much smaller, has a round tail, and a proportionally longer snout. 
Fossil remains of Jf. moschata occur in the Norfolk Forest bed, and 
were originally described under the name of Palwospalar. The 
genus is also represented in the Middle and Lower Miocene of the 
Continent. 

Urotrichus.1—Dentition : i 2, c+, p 4 or 3,m3; total 36. Feet 
not webbed; manus broad. Habits fossorial. The Mole-Shrews, 


Fic. 289.—The Desman (Myogale moschata). 4 natural size. 


as these animals are called, are represented by C. ¢ulpvides of the 
mountains of Japan and C7 gibbsi of North America. These two 
species are small and closely allied animals; the American form 
(which it has been proposed to separate subgenerically as Neuro- 
trichus) having p 3. 

Uropsilus.*—Dentition : 2 #, ¢ 4, p 3, m 3; total 34. Manus 
narrow; tail naked and scaly. Habits cursorial. The single 
species, U. svricipes, from the borders of Tibet, is a slate-coloured 
animal with the external form of a Shrew but the skull of a Mole. 


1 Temminck, Fw Japonica, vol. i, p. 22 (1842). ? Milne-Edwards, 
rch. du Museum, vol. vii. Bull. p. 92 (1872). 


630 INSECTIVORA 


Subfamily Talpinge.—Clavicle and humerus very short and 
broad ; manus with a large falciform bone. 

A. First upper incisor much larger than the second (New 

_ World Moles). 

Scalops..—Dentition: i #,¢34, p 3, m2; total 36. Extremity 
of muzzle simple ; hind feet webbed ; tail short and nearly naked. 
Represented by three species in the United States. 

Scapanus.-—Dentition : i 3,¢4, p4,m 4; total 44. Extremity 
of muzzle simple. The two North American species of this genus 
resemble Scalops in general characters, but have a dentition like 
Condylura. The habits are like those of the latter, and the right 
to generic distinction is doubtful. 

Condylura.2— Dentition: i 8, ¢ 4, p 4, m 2; total 44. Ex 
tremity of muzzle surrounded by filiform appendages. The Star- 
nosed Mole (C. cristata) derives its name from the star-like ring of 
appendages at the extremity of the muzzle, with the nostrils in the 
centre. The general contour is Mole-like, but the tail is nearly as 
long as the body, and the manus is somewhat less powerful, with 
its terminal phalanges not cleft. The length of the head and body 
is about 5 inches. This species is common in parts of North 
America, and forms tunnels in the ground like the Common Mole. 

B. First upper incisor scarcely larger than the second (Old 
World Moles). 

Scaptonyx.*—Dentition: i #, ¢4, p 4, m2; total 42. Manus 
moderately broad, as in Urotrichus. Represented only by 8S. fusi- 
caudatus of Eastern Tibet, which may be regarded as connecting 
Talpa with Urotrichus, having the head of the former and the limbs 
of the latter. 

Talpa.’,—Dentition (usually): 7 3, ¢ 3, p 4, m 3; total 44. 
Manus extremely broad. 

This genus includes the true Moles, of which the common 
English Mole ® (Tf. europea) is the type. This animal is about 6 
inches in total length, of which rather more than one inch is occu- 
pied by the tail. The body is elongated and cylindrical, and, owing 
to the very anterior position of the fore limbs, the head appears to 
rest between the shoulders; the muzzle is long and obtusely 
pointed, terminated by the nostrils, which are close together; the 
minute eye is almost hidden by the fur; the ear is without a conch, 
and opens on a level with the surrounding integument. The fore 
limbs are rather short and very muscular, terminating in broad, 
naked, shovel-shaped feet, with the palms normally directed out- 


* Cuvier, ‘‘Tabl. de Classif.” in Leon @ Anat. Comp. vol.i. (1800). 2 Pomel, 
Arch. Sct. Phys. Nat. vol. ix. p. 247 (1848). 3 Iliger, Prodromus Syst. Mamm. 
et Avium, p. 125 (1811). 4 Milne-Edwards, V. Arch. du Muséum, vol. vii. 
Bull. .p. 92 (1872). > Linn, Syst. Not. 12th ed. p. 73 (1766). The following 
account is taken almost entirely from Dr. Dobson. 


TALPIDA 631 


wards, and each with five subequal digits armed with strong flattened 
claws. The hind feet are long and narrow, and the toes are pro- 
vided with slender claws. The body is densely covered with soft, 
erect, velvety fur, the hairs being uniform in length and thickness, 
except on the muzzle and short tail. The colour of the fur is 
generally black, with a more or less grayish tinge, or brownish-black, 
but various paler shades up to pure white have been observed. 

The food of the Mole consists chiefly of the earth-worm, in 
pursuit of which it forms its well-known underground excavations. 
Its habits were many years ago studied and described by M. Henri 
le Court. Like many other mammals, the Mole has a lair to which 
it may retire for security. This consists of a central nest formed 
under a hillock, placed in some protected situation, as under a bank, 
or between the roots of trees. The nest, which is lined with dried 
grass or leaves, communicates with the main run by four passages, 
of which only one joins it directly, leading downwards for a short 
distance and then ascending again. The other three are directed 
upwards and communicate at regular intervals with a circular 
gallery constructed in the upper part of the hillock, which in turn 
communicates by five passages leading downwards and outwards 
with another much larger gallery placed lower down on a level 
with the central nest, from which passages proceed outwards in 
different directions, one only communicating directly with the main 
run, while the others, curving round, either soon join or end blindly. 
The main run is somewhat wider than the animal’s body ; its walls 
are smooth, and formed of closely compressed earth, the depth 
varying according to the nature of the soil, but ordinarily from 4 
to 6 inches. Along this tunnel the animal passes backwards and 
forwards several times daily, and here traps are laid by mole-catchers 
for its capture. From the main run numerous passages are formed 
on each side, along which the animal hunts its prey, throwing 
out the soil in the form of mole-hills. The Mole is one of the 
most voracious of mammals, and, if deprived of food, is said to die 
in from ten to twelve hours. Almost any kind of flesh is eagerly 
devoured by captive Moles, which have been seen by various 
observers, as if maddened by hunger, to attack animals nearly as 
large as themselves, such as birds, lizards, frogs, and even snakes ; 
toads, however, they will not touch, and no form of vegetable food 
attracts their notice. If two Moles be confined together without 
food, the weaker is invariably devoured by the stronger. Moles 
take readily to the water, in which respect they resemble their 
representatives on the North American continent. Bruce, writing 
in 1793, remarks that he saw a Mole paddling towards a small 
island in the Loch of Clunie, 180 yards from land, on which he 
noticed mole-hills. 

The sexes come together about the second week in March, and 


632 


INSECTIVORA 


the young—generally from four to six in number—which are 


sy 
any 


\ 


Fic. 290.—Skeleton of Mole x 2 (ower jaw 
removed to show base of skull). c, Caleaneum; 
¢.h., clavicular articulation of the humerus; cl., 
clavicle; e.c, external condyle of humerus; f, 
femur; fb, fibula; fc, falciform bone (radial sesa- 
moid); h, humerus; i.c, internal condyle of 
humerus ; i, left ilium; ip, ramus of the ilium 
and pubis; is., ischium; l.d, ridge of insertion of 
Jatissimus dorsi muscle ; J.t, lesser trochanter ; m, 
manubrium sterni; 0, fourth intercentral ossicle ; 
ol, olecranon; p., pubis widely separated from that 
of the opposite side; pa., patella; p.m., ridge for 
insertion of pectoralis major muscle; pt., pectineal 
eminence; r, radius; rb, first rib; s, plantar sesa- 
moid ossicle corresponding to the radial sesamoid 
(os faleiforme) in the manus; se., scapula; Sh, 
scapular articulation of the humerus; t, tibia; u, 
ulna. 


brought forth in about six 
weeks, quickly attain their 
full size. 

The Mole exhibits in the 
whole of its organisation a 
perfect adaptation to its 
peculiar mode of life. In 
the structure of the skeleton 
(Fig. 290) very striking de- 
partures from the typical 
mammalian form are notice- 
able. Thus the presternum 
is so much produced anteriorly 
as to extend forward as far as 
a vertical line from the second 
cervical vertebra, carrying 
with it the very short and 
almost quadrate clavicle, which 
is articulated with its anterior 
extremity and distally with 
the humerus ; being also con- 
nected ligamentously with the 
scapula. The fore limbs are 
thus brought opposite the 
sides of the neck, and from 
this position a threefold ad- 
vantage is derived: in the 
first place, as this is the 
narrowest part of the body, 
they add but little to the 
general width, which, if in- 
creased, would lessen the 
power of movement in a 
confined space ; secondly, this 
position allows of a longer 
fore limb than would other- 
wise be possible, and so in- 
creases its power ; and, thirdly, 
although the entire limb is 
relatively very short, its an- 
terior position enables the 
animal, when burrowing, to 
thrust the claws so far for- 
ward as to be in a line with the 
end of the muzzle, the import- 


TALPIDE 633 


ance of which is evident. Posteriorly, the hind limbs are similarly 
removed out of the way by approximation of the hip-joints to the 
centre line of the body. This is effected by inward curvature of 
the innominate bones at the acetabula to such an extent that they 
almost meet in the centre, while the pubic bones are widely separated 
behind. The shortness of the fore limb is caused by the great 
reduction in the length of the humerus, which has lost all resemblance 
to its normal shape. In addition to the usual articulation with the 
glenoid cavity of the scapula, the humerus also has a separate 
articulation with the extremity of the clavicle. The bones of the 
manus are enormously expanded laterally; this expansion being 
increased by the large sickle-like bone on the radial side of the 
carpus, which is considered by some anatomists to represent the 
prepollex. The skull is long and tapering, with very slender 
zygomatic arches and elongated nasals, which are ankylosed 
together, and in advance of which the mesethmoid is more or 
less ossified. The vertebre are usually C 7, D 13, L 6, 8S 6, 
C 10-12; all having very strong surfaces for mutual articulation. 
The upper incisors are chisel-like, and the canine has two roots ; 
the first three upper premolars are simple and conical, but the 
fourth is much larger, and canine-like. In the mandible the 
incisors are small and somewhat proclivous, while the canine can 
only be distinguished from them by its position; the first lower 
premolar is larger than the others. 

The Common Mole has an exceedingly wide distribution, 
ranging over the greater part of the Palearctic region, where it is 
met with in places so widely sundered as England and Japan. It 
occurs in both the Himalaya and Altai mountains. In Ireland it 
is unknown, and in Scotland it extends as far north as Caithness. 
Eight species of the genus are recognised, which may be grouped, 
from the characters of their dentition, as follows, viz.: 7 3, ¢ 4, p 4, 
m 3, T. wogura; 1 3, ¢ 4, p 4, m 4, T. ewropea, ceca, longirostris, 
mierura ; 13,64, p 3, m2, T. leucura, leptura ; i 3, ¢ 4, p 3, m 3, 
7. moschata. 

Except in I. europea, the eyes are covered by a membrane. In 
T. micrura the short tail is concealed by the fur. 7. ceca is found 
south of the Alps; the remaining species are Asiatic, and two only 
—T. micrura and T. leucura—occur south of the Himalaya. 7. 
moschata, of Tibet, is regarded by some zoologists as generically 
distinct under the name of Scaptochirus. 

Remains of 7. europea occur in the Norfolk Forest bed, while 
extinct species are found in the European Tertiaries as far down as 
the Lower Miocene, although it has been proposed to separate 
some of these forms generically. Protalpa, of the Upper Eocene 
Phosphorites of Central France, is very closely allied, but the 
structure of the humerus is somewhat less specialised. 


634 INSECTIVORA 


Extinct Genera. —A number of extinct Insectivora from the 
European Tertiaries more or less closely allied to the Moles have 
been described, but since our knowledge of most of them is 
extremely imperfect their precise affinities are in many instances 
problematical. Of these the Lower Miocene Teéracus is said to have 
affinity both with J/yogale and FErinaceus; while the forms 
described as Mysarachne and Echinogale, are considered to connect 
the present with the two preceding families. Plesiosorex is another 
Lower Miocene type known only by the mandible, in which there 
are ten teeth ; it is generally referred to the Alyogalinw. The minute 
Amphidozotherium, of the French Phosphorites, is considered to be 
allied to Urotrichus. 


Family ADAPISORICIDE 


This extinct family is represented by the genera Aduapisorex and 
.tdapisoriculus, of the lowest Eocene of Rheims, which are regarded 
as allied to the Soricide, but somewhat more specialised. In the 
type genus the formula of the lower teeth is 7 2,¢1, p 4, m3; 
the incisors and canine being proclivous, and the molars (of which 
the last is small and without a third lobe) quadritubercular. 
Adapisoriculus is a smaller form with differently shaped molars. 

Here also may be mentioned the genera Orthaspidotherium and 

b Pleuraspidotherium, from the above- 
‘ 7“ mentioned deposits, which are prob- 
me ably members of the present order. 
a @r They appear to have been animals 
p a ne ae oa ‘i heex, SOMeWwhat smaller than a Hedgehog, 
teeth Gf Pcinotninen Gennes with quadritubercular up Pp er molars 
from the Lowest Eocene of Rheims. pr, (Fig. 291), and the hinder premolars 
protocone ; me, metacone; pa, paracone; more complex than those of the 
ie ied ig a aa Erinaceide. In the first-named genus 
the dental formula is i $, ¢ 4, p 4, m 3; the third and fourth upper 
premolars having one outer column. Pleuraspidotherium has ap- 
parently only three premolars, of which the third and fourth 
(Fig. 291) have two outer columns. The humerus in both has 
no entepicondylar foramen, the femur has a third trochanter, and 
the astragalus is vertically perforated. 


Family POTAMOGALIDA, 


Skull with a small brain-case, no zygomatic arch or postorbital 
process, and the tympanic annulate and not forming a bulla. 
Upper molars with the cusps arranged in a broad V, and some- 
what intermediate in structure between those of the preceding and 
succeeding families. No clavicle; pubic symphysis ligamentous ; 


SOLENODONTIDA 635, 


tibia and fibula typically united distally. No cecum. Confined to 
the Ethiopian region. 

Potamogale.A—Dentition: 1 2, ¢ 1, p 3, m2; total 40. Repre- 
sented only by P. velox of Western Equatorial Africa. This animal 
(Fig. 292) inhabits the banks of streams, and is thoroughly adapted 
for an aquatic life; it is nearly 2 feet in length, the tail measuring 
about half. The long cylindrical body is continued uninterruptedly 
into the thick laterally compressed tail, the legs are very short, and 
the toes are not webbed, progression through the water evidently 
depending wholly on the action of the powerful tail, while the 
limbs are folded inwards and backwards. The muzzle is broad and 


Fic. 292.—Potamogale veloz. x 4. (From Allman, Trans. Zool. Soc. vol. vi. pl. i.) 


flat, and the nostrils are protected by valves. The fur is dark 
brown above, the extremities of the hairs on the back being of a 
metallic violet hue by reflected light, beneath whitish. This curious 
animal was discovered by M. du Chaillu. 

Geogale.-—Dentition : i ,¢ 4, p 3,m 3; total 34. This genus 
is known solely by G@. aurita, a small Mouse-like species from Mada- 
gascar, agreeing closely with Potamogale in the general form of the 
skull and teeth. The tibia and fibula are distinct, but it 1s not 
known whether a clavicle exists ; and the material at present avail- 
able is insufficient to definitely fix the natural position of the genus. 


Fumily SOLENODONTIDA. 


Skull with a small brain-case constricted between the orbits, no 


1 Du Chaillu, Proc. Boston Soc. Hist. Nut. vol. vii. p. 363 (1860). 
2 Milne-Edwards, Ann. Sci. Nat. vol. xv. p. 5 (1872). 


636 INSECTIVORA 


zygomatic arch or postorbital process, and the tympanic annulated 
and not forming a bulla. Upper molars tritubercular, the cusps 
being arranged in a V. Pubic symphysis short; tibia and fibula 
distinct. Vertebre: C 7,D 15,L4,85,C 23. Nocecum. The 
penis is carried forwards and suspended from the abdomen; the 
testes are received into perineal pouches ; the mammary glands are 
post-inguinal ; the uterine cornua end in cecal sacs. 
Solenodon.i—Dentition : 1 3,¢ 4, p 4,m 2; total 40. This genus, 
with S. paradorus and 8. cubanus (Fig. 293), from Hayti and Cuba 


Fic, 293.-—Solenodon cubanus. x }. (From Peters, Abh. Akad. Berlin.) 


respectively, alone represents the family. These species, which 
differ chiefly in the colour and quality of the fur, have a remark- 
ably long cylindrical snout, a long naked tail, feet formed for 
running, and the body clothed with long, coarse fur. 

The position of the mammz quite behind on the buttocks is 
unique among Insectivora. The first upper incisor is much enlarged, 
and this and the other incisors, canines, and premolars, closely 
resemble those of Myogale; the second lower incisor is, as in 
Potamogale, much larger than the anterior one, and is deeply 
hollowed out internally. While thus apparently showing relation- 
ship with the Talpide, the form of the crowns of the molar teeth 
connects them with the next family. 


1 Brandt, Mem. te. Linp. St. Pétersbourg, 1833, vol. ii. p. 459. 


CENTETIDA 637 


Family CENTETIDA, 


Skull (Fig. 294) with a small cylindrical brain-case not con- 
stricted between the orbits, no zygomatic arch or postorbital pro- 
cess, and the 
tympanic annu- 
late and not 
forming a bulla. 
Upper molars 
tritubercular. 
Pubic sym- 
physis short; 
and the tibia 
and fibula either 
united or free. 
No cecum. The 
penis is pendent and retractile within the fold of the integument 
surrounding the anus; the testes are abdominal; the mamme are 
thoracic and ventral; and the uterine cornua are terminated by 
the Fallopian tubes. All the species are limited to Madagascar. 

Subfamily Centetinze.—Tibia and fibula distinct; testes near 
kidneys ; fur with spines. 

Centetes.1—Dentition: i $,¢ 4, p 3, m 4; total 38. Vertebre: 
C7,D19,L 5,8 3,C 8. The single species is the well-known 
Tenrec (C. ecaudatus), characterised by the absence of a tail; it 
reaches a total length of from 12 to 16 inches, and is the largest 
known Insectivore. The adult males have long canines, the 
extremities of the lower pair being received into pits in front of 
the upper ones (Fig. 294). It is probably the most prolific of all 
mammals, since as many as twenty-one young are said to have been 
brought forth at a birth. The young have strong white spines 
arranged in longitudinal lines along the back, but these are lost in 
the adult animal, which is provided only with a nuchal crest of 
long rigid hairs. In rare instances a fourth upper molar may be 
developed. 

Hemicentetes.2—Dentition : 1 3, ¢ 4, p 3, m 3; total 40. This 
genus is represented by the two species HH. semispinosus (of which 
the skull is shown in Fig. 295) and H. nigriceps. It differs from 
Centetes by the presence of the third upper incisor, the much smaller 
canines, and by the form of the skull. Both species are very much 
smaller than C’. ecuudatus, and the dorsal spines are retained in the 
adult state. Vertebre: C 7, D16,L5,8 3,09. 


Fic. 294.—Left lateral view of the skull of the Tenrec (Centetes 
ecaudatus). Reduced. 


1 Illiger, Prodromus Syst. Mami. et Avium, p. 124 (1811). 2 Mivart, 
Proc. Zool. Soc. 1871, p. 72. 


638 INSECTIVORA 


Ericulus.i—Dentition: «2, ¢4, p 3,m; total 36. Vertebre: 
C7,D 17, L 6,8 4, C 9. The single species, £. setosus, is a 
Hedgehog-like animal, having the whole upper surface and the 
short tail densely covered with close-set spines. The facial bones 
are much shorter than in any of the preceding genera, and the 
first upper incisor is elongated, as in Erinaceus. Judging from 
the slight development of the cutaneous muscles compared with 
those of the true Hedgehogs, it is probable that complete involution 
of the body does not take place. 

Subfamily Oryzorietinze.—Tibia and fibula united ; testes near 
urethra ; fur without spines. 

Microgale.2—Dentition : 13, ¢4, p 3,m%; total 40. This genus 


BE 


Fic. 295.—Skull of Hemicentetes semispinosus. X 2. (From Mivart, Proc. Zool. Soc. 1871.) 


includes AL. longicaudata and AL. cowani, both of which are small 
Mouse-like species, the former with a tail double the length of the 
head and body, and having 43 caudal vertebre ; teeth like those of 
Centetes ecwudatus, but, owing to the comparatively much shorter 
muzzle, not separated by wide spaces, and the last premolar and 
molar with internal basal processes. 

Oryzorictes.-—Represented by two species, 0. hova and 0. tetra- 
dactylus, the latter distinguished by the presence of only four digits 
in the manus, the three inner having long laterally compressed 
fossorial claws. The general form of the head and body of the 
two species known is like that of a Mole. These animals burrow 
in the rice-fields and do much damage to the crops. 


Family CHRYSOCHLORID&, 


Skull conical, not constricted between the orbits, without post- 
orbital process, but with well-developed zygomatic arch and tympanic 


1 I. Geoffroy, Ann. Sei. Nat. ser. 2, vol. viii. p. 60 (1837). > Thomas, 
Journ. Linn. Soc.—Zool. vol. xvi. ». 319 (1882). ’ Grandidier, Rev. and 


Mag. Zool. 1870, p. 50. 


CHRYSOCHLORIDA 639 


bulla. Upper molars tritubercular, with the crowns very tall. 
No pubic symphysis; the tibia and fibula united. The eyes are 
covered by the hairy integument ; the ears short and concealed by 
the fur; the internal generative organs are as in Centetine ; the 
mamme are thoracic and inguinal and placed in cup-shaped depres- 
sions. Habits fossorial. Confined to the southern part of the 
Ethiopian region, not extending to Madagascar. 

This family is closely allied to the Centetide, occupying the 
same relative position with respect to that family that the Talpide 
does to the Soricide. Compared with the Talpide, we find the 
following differences in the structural adaptation to a fossorial life ; 
the manubrium sterni is not anteriorly elongated, neither are the 


Fic. 296.—The Golden Mole (Chrysochloris obtusirostris). 


clavicles shortened ; but this is compensated for by a deep hollowing 
out of the antero-lateral walls of the thorax, the ribs in these parts 
and the sternum being convex inwards. The long clavicles have 
their distal extremities pushed forward, and the concavities on the 
sides and inferior surface of the thorax lodge the thick muscular 
arms. 

Chrysochloris.A—Dentition : 1 3, ¢ 4, p 3, m SS ; total 40 or 36 
Vertebre: C 7,D 19, L 3,8 5, C 8. This genus includes some 
seven or eight South African species, commonly known as Golden 
Moles (Fig. 296). Those species, in which the molars are reduced 
to 2, with a basal talon to the lower ones, and without a prominence 
in the temporal fossa, have been placed in a separate genus, 
Chalcochloris, by Professor Mivart. Nearly all the species have the 
fur of the upper surface of a brilliant metallic lustre, varying from 
golden bronze to green and violet of different shades. The manus 


1 Lacépede, Mém. de 0 Institut, vol, iii. p. 493 (1801—read 1799). 


640 INSECTIVORA 


has four digits, of which the two outer are small, while the middle 
ones are large, with immensely powerful claws. 

Extinct Types.—The only fossil forms which can be referred to 
the section of the Insectivora with tritubercular molars are the 
Leptictide, of the Eocene and Miocene of North America. This 
family includes the genera Leptictis, MJesodectes, and Ictops, all of 
which are regarded by Dr. Schlosser as true Insectivora, although 
they were placed by Professor Cope with the Creodont Carnivora. 


Bibliography of Insectivora.—Peters, Reise nach Mossambique—Sdugeth. 1852; 
Id. ‘“‘Ueber die Classification der Insectivora,” JJonatsh. Akad. Wissensch. 
Berlin, 1865, and other papers; Mivart, ‘‘On the Osteology of Insectivora,” 
Journ. Anat. and Phys. 1867, 1868, and Proc. Zool. Soc. 1871; Gill, ‘Synopsis of 
Insectivorous Mammals,” Bull. Geal. and Geog. Survey, U.S.A. Washington, 
1875 (includes a general bibliography of the order) ; Dobson, Monograph of the 
Insectivora, Systematic and Anatomical, London, 1882-90. 


CHAPTER XIII 


THE ORDER CHIROPTERA 


MAmMALs, having their fore limbs specially modified for flight. 
The forearm consists of a rudimentary ulna, and a long curved 
radius. The carpus has six bones supporting a small pollex and 
four greatly elongated fingers, between which and the sides of 
the body and the hinder extremities a thin expansion of the 
integument (the wing-membrane or patagium) is extended. The 
knee is directed backwards, owing to the rotation of the hind limb 
outwards by the wing-membrane ; a peculiar elongated cartilaginous 
process (the calcar), rarely rudimentary or absent, arising from the 
inner side of the ankle-joint, is directed inwards, and supports part 
of the posterior margin of an accessory membrane of flight, extending 
from the tail or posterior extremity of the body to the hinder limbs 
(the inter-femoral membrane). The penis is pendent; the testes are 
abdominal or inguinal ; the mammary glands thoracic and generally 
postaxillary ; the uterus is simple or with more or less long cornua ; 
the placenta discoidal and deciduate; and the smooth cerebral 
hemispheres do not extend backwards over the cerebellum. The 
dental series includes incisors, canines, premolars, and molars and 
never exceeds 7 2, ¢ 4, p 3, m 4; total 38. 

The animals comprised in this order are at once distinguished 
by the presence of true wings, and this peculiarity is accompanied 
by other modifications of bodily structure having special relation to 
flight. Thus, in contrast to most other mammals, in which the hind 
limbs greatly preponderate in size over the fore, in the present 
order the fore limbs immensely exceed the short and weak hinder 
extremities. The thorax, as giving origin to the great muscles 
which sustain flight, and containing the proportionately large lungs 
and heart, is remarkably capacious, and the ribs are flattened and 
close together; the shoulder-girdle is also greatly developed in 
comparison with the weak pelvic bones. 

Linneus included the Bats among the Primates, mainly on 

41 


642 CHIROPTERA 


account of the number of their upper incisors, supposed to be 
always four, the thoracic position of the mamme, and the pendent 
condition of the penis. Many other zoologists, taking into con- 
sideration the placental characters and the form of the uterus, have 
followed him; but it is evident that the situation of the mammze 
is related to the necessarily central position of the young during 
flight, the shortness of the uterine cornua, observable in so many 
species, to the generally uniparous gestation requiring less room, 
while the discoidal deciduate placenta is equally present in and 
characteristic of the Insectivora, many species of which also have 
the penis pendent. Thus, the reasons for maintaining the Bats in 
this high position being disposed of, we find in the low organisation 
of their brain a proof of their inferior status; while furthermore, 
although they differ widely from all other mammals in external 
form, it is evident that this is only the result of special adaptation 
to aerial locomotion ; and, taking into account their whole bodily 
structure, we may accept the view of Professor Huxley that they 
should merely be regarded as exceedingly modified Insectivora. 

So thoroughly, however, has this adaptation for flight been 
carried out that of all animals the Bats are the least terrestrial, not 
one of them being equally well fitted for progression on the earth. 
This is due to the hind as well as the fore limbs being pressed into 
the service of aerial locomotion. Thus the hind limb is so rotated 
outwards by the wing-membrane that, contrary to what obtains in all 
other vertebrates, the knee is directed backwards, and corresponds 
in position to its serial homologue the elbow. It necessarily follows 
from this arrangement that when a Bat is on the ground it rests on 
all fours, having the knees directed upwards; while, in order: to 
bring it into a position for forward progression, the foot rotates 
forwards and inwards on the ankle. Walking under these circum- 
stances is at best only a kind of shuffle, and that this is fully 
recognised by the animal is evidenced by its great anxiety to take 
wing, or, if this be impracticable, to ascend to some point where it 
can hitch itself up by the claws of the hind legs in its usual position 
when at rest. 

The bones of the skeleton are characterised by their slender- 
ness and the great size of the medullary canals in those of the 
extremities. The vertebral column is short, and the vertebra differ 
very slightly in number and form throughout the species. The 
general number of the dorso-lumbar vertebre is 17, of which 12 
are dorsal; the cervicals are very broad, but short from before 
backwards, their breadth being due to the great transverse 
diameter of the spinal canal rendered necessary by the compara- 
tively large size of the spinal cord, which, after giving off the nerves 
to the fore limbs and thorax, rapidly diminishes in size, and in the 
lumbo-sacral region is reduced to a fine thread. Except in the 


CHIROPTERA 643 


frugivorous Pteropudide, the vertebre, from the third cervical back- 
wards, are devoid of neural spines. From the first dorsal to the 
last lumbar vertebra the spinal column forms a single curve back- 
wards, which is most pronounced in the lumbar region. The centra 
of the vertebre are but slightly movable upon each other, and in 
old individuals appear to become partially ankylosed together. 
The caudal vertebre are simple cylindrical bones without processes ; 
their number and length being extremely variable even in closely 
allied species ; and the anterior caudals are generally united to the 


Fic. 297,—Skeleton and flying-membranes of the Noctule Bat (Vesperugo noctula). x }. 
c, Clavicle; h, humerus; 7, radius; vu, ulna (rudimentary); d@}, pollex; ¢@°, d3, d4, @5, other 
digits of the manus supporting wm, the wing-membrane ; a, m, metacarpal bones ; phi, first 
phalanx ; ph?, second phalanx; pk, third phalanx; am, antebrachial membrane; jf, femur; 
t, tibia; fb, fibula (rudimentary) ; c, calear supporting im, the interfemoral membrane; pel, post- 
calcaneal lobe. 


ischial tuberosities. The relative development of the caudal 
vertebre is, indeed, intimately correlated to the habits of the 
animals ; the long tail in the insectivorous forms supporting and 
controlling the position of the large interfemoral membrane, which 
appears not only to aid their rapid motions when in pursuit of their 
prey by acting as a rudder, but also to assist in the capture and 
retention of the larger insects. In the frugivorous types, on the 
other hand, this is not required, and the tail is accordingly rudi- 
mentary or absent. In all Bats the presternum has a prominent 
keel for the attachment of the great pectoral muscles. In most 


644 CHIROPTERA 


species the ribs are much flattened, and in some they are partially 
ankylosed by their contiguous margins. 

The skull is subject to considerable structural variations, 
even within the limits of a single family. Postorbital processes 
to the frontals are found only in the Péeropodide and some 
Nycteride and Emballonuride. Pteropus leucopterus and Pteralopex 
are peculiar in having the orbit completely surrounded by 
bone. <A slender zygomatic arch is present, except in some of the 
Phyllostomatide. 

The milk-teeth are peculiar in that they are utterly unlike those 
of the permanent series. They are slender, with sharp recurved 
cusps ; and as a rule are shed at an early period (in the Rhino- 
lophide before birth), but may coexist with some of the fully 
developed permanent teeth. The permanent teeth are subject to 
great variation of form, although they always have distinct roots. 
In the Insectivorous types they are acutely cusped, the cusps in 
those of the upper jaw being arranged in a more or less distinct W ; 
but in the frugivorous forms, like the Pteropodide and some of the 
Phyllostomatide, the molars are longitudinally grooved or hollowed 
out. 

The pectoral girdle maintains a very constant type. Thus the 
clavicle is very long, strong, and curved; and the scapula large, 
oval, triangular, with a long curved coracoid process. The humerus, 
though long, is scarcely two-thirds the length of the radius. The 
ulna is rudimentary, its proximal extremity, which articulates with 
but a small part of the humerus, being ankylosed to the radius ; 
and immediately beyond the joint it is reduced to a slender splint- 
like bone, extending about as far as the middle of the radius. In 
all species a detached sesamoid bone exists in the tendon of the 
triceps muscle. The radius is very long, in some species actually 
equal to the length of the head and body. The proximal row of 
the carpus consists of a single bone formed by the united scaphoid, 
lunar, and cuneiform; which, with the extremity of the radius, 
forms the radio-carpal joint. In the distal row the trapezium, 
trapezoid, and magnum vary in size in the different families, the 
unciform appearing to be the most constant, and the pisiform being 
generally very small. 

The manus is always furnished with five digits. The first, 
fourth, and fifth digits consist of a metacarpal and two phalanges ; 
but in the second and third digits the number of phalanges is 
different in certain families. The pollex always terminates in a 
claw, which—like the proximal phalanx—is best developed in the 
frugivorous species. In most of the frugivorous Pteropodide the 
second digit is provided with a claw; but in all other Bats this and 
the remaining digits are unarmed. In the genus 7rianops alone a 
very peculiar short bony process projects from the outer side of 


CHIROPTERA 645 


the proximal extremity of the terminal phalanx of the fourth digit. 
The relative development of the digits and their phalanges will be 
noticed under each family. 

As might be expected from the small size of the posterior 
limbs, the pelvic girdle is relatively weak. The ilia are long and 
narrow. In the males of most species the pubic bones of opposite 
sides are very loosely united in front, while in females they are 
widely separated; and in the family [hinolophide alone do these 
bones form a symphysis. The ileo-pectineal eminence develops a 
long pectineal process, which in the subfamily Hipposiderine is con- 
tinued forwards to the anterior extremity of the ilium enclosing a 
preacetabular foramen unique among mammals. The acetabulum 
is small and directed outwards and slightly upwards; and with 
this is related the peculiar position of the hind limb already noticed 
as one of the chief characteristics of the order. The femur is 
slender and cylindrical, with a small head and very short neck, and 
scarcely differs in form throughout the order. The bones of the 
leg and foot are variable ; in the subfamily Molossine alone is there 
a well-developed fibula, while in all other species this bone is either 
very slender, or cartilaginous and ligamentous in its upper third, or 
reduced to a small bony process above the heel, as in Megaderma, 
or altogether absent, as in Nycteris. 

The foot consists of a very short tarsus, and of slender, later- 
ally compressed toes, with much curved claws. The hallux is 
composed of a metacarpal, a proximal and an ungual phalanx, and 
is slightly shorter than the other four toes, each of which has an 
additional phalanx, except in the subfamily Hipposiderine and in 
the anomalous genera Thyroptera and Jfyxopoda, where all the toes 
have the same number of phalanges as the first digit, and are equal 
to it in length. In the genus Chiromeles the first digit is thumb- 
like and separated from the others, and in the typical Molossinw 
the first and fifth digits are much thicker than the intermediate 
toes. 

The most noticeable peculiarities in the myology of the order 
consist in the separated bands or slips into which the platysma is 
divided, and in the presence of the remarkable muscle termed 
occipito-pollicalis, which extends from the occipital bone to the base 
of the terminal phalanx of the pollex. 

Although, as already mentioned, the brain presents a low type 
of organisation, yet probably no animals possess so delicate a sense of 
touch as the Chiroptera. It is undoubtedly this perceptive power 
which enabled the individuals deprived of sight, hearing, and smell, 
in Spallanzani’s well-known experiments, to avoid the numerous 
threads hung across the rooms in which they were permitted to fly 
about. In the common Bats the tactile organs evidently exist, not 
only in the delicate vibrissee which spring from the sides of the 


646 CHIROPTERA 


muzzle, but also in the highly sensitive and widely extended integu- 
mentary structures entering into the formation of the wing-mem- 
branes and ear-conchs; while in many other species, notably in the 
tropical Rhinolophine and Phyllostomatine Bats, peculiar foliaceous 
cutaneous expansions surrounding the nasal apertures or extending 
backwards behind them are added. These structures, collectively 
known as the “nose-leaf” (whence the term “leaf-nosed Bats”), 
have been shown by Dr. Dobson to be made up partly of the 
extended and thickened marginal integument of the nostrils, and 
partly of the highly differentiated glandular eminences occupying 
the sides of the muzzle, in which, in all the common Bats, the 
vibrissz are implanted. 

In all species of leaf-nosed Bats, and especially in the Phino- 
lopade, where the nasal appendages reach their highest development, 
the superior maxillary division of the fifth nerve is of remarkably 
large calibre. The nasal branch of this nerve, which is given off 
immediately beyond the infraorbital foramen, is by far the largest 
portion ; the palpebral and labial branches consisting of a few 
slender nerve-fibres only. This branch passes forwards and upwards 
on the side of the maxilla, but soon spreads out into numerous - 
filaments extending into the muscles and integument above, and 
into the base of the nose-leaf. The nerve supply of the nose-leaf is 
further augmented by the large nasal branch of the ophthalmic 
division of the fifth nerve. While the many foliations, elevations, 
and depressions which vary the form of the nose-leaf greatly increase 
the sensory surface supplied by the fifth nerve, and during rapid 
flight intensify the vibrations conveyed to it, the great number of 
sweat and oil glands which enter into its structure perform a func- 
tion analogous to that of the glands of the auditory canal in relation 
to the membrana tympani in maintaining its surface in a highly 
sensitive condition. The nasal appendages of the Chiroptera may 
thus be regarded as performing the office of an organ of a very 
exalted sense of touch standing in the same relation to the nasal 
branches of the fifth nerve as the aural apparatus to the auditory 
nerve ; for, as the latter organ collects and transmits the waves of 
sound, so the former receives impressions arising from vibrations 
communicated to the air by approaching objects. 

In no order of mammals is the ear-conch so greatly developed or 
so variable in form. Thus in most of the insectivorous species the 
ears are longer than the head, while in some, as in the common 
Long-eared Bat (Plecotus auritus), their length nearly equals that of 
the head and body. The form of the conch is very characteristic of 
the various families; in most the tragus is remarkably large, in 
some extending nearly to the outer margin of the conch; and its 
function appears to be to cause undulations in the waves of sound, 
and so intensify and prolong them. It is worthy of notice that in 


CAHIROPTERA 647 


the Rhinclophide, the only family of insectivorous Bats wanting the 
tragus, the auditory bull reach their greatest size, and the highly 
sensitive nasal appendages their highest development; and that in the 
typical group of the Aolossine the ear- 
conch is divided by a prominent keel ; 
and the antitragus is unusually large 
in those species in which the tragus is 
minute (see Fig. 298, a). In the frugi- 
vorous Bats the form of the ear-conch 
is very simple, and but slightly variable 
throughout the various types. 

In all Bats the ears are extremely 
mobile, each moving independently at 
the will of the animal. This has been 
observed even in the frugivorous Pferv- 
podide, in which the peculiar vibratory 


3 Z : Fra. 298. —Head of Molossus glaucinus. 
movements first noticed in 7fileus (prom Dobson, Proc, Zool. Soe. 1876.) a, 


perspicillatus may also be seen when Antitragus; , keel of the ear-coneh ; ¢, 


th e animals are alarm e d notch behind antitragus. 
a a ala . 


The opening of the mouth is anterior in most species, but in 
many it is inferior, the extremity of the nose being more or less 
produced beyond the lower lip,—so much so indeed in the small 
South-American species Lhynchonycteris naso as to resemble that of 
the Shrews. The lips exhibit the greatest variety in form, which 
will be referred to under each family. The absence of a fringe 
of hairs is characteristic of all fruit-eating Bats, and probably 
always distinguishes them from the insectivorous species, which they 
may resemble in the form of their teeth and other respects. 

The cesophagus is narrow in all species, and especially so in the 
sanguvorous Desmodont Phyllostematidie. The stomach presents two 
principal types of structure, which correspond respectively to the 
two great divisions of the order, the Megachiroptera and the Micro- 
chiroptera ; in the former (with the exception of Harpyia) the 
pyloric extremity is more or less elongated and folded upon itself, 
in the latter it is simple, as in the Insectivora Vera; a third 
exceptional type is met with in the Desmodont Phiyllostomutide, 
where the left or cardiac extremity is greatly elongated, forming a 
long narrow excum-like appendage. The intestine is comparatively 
short, varying from one and a half to four times the length of the 
head and body, being longest in the frugivorous and shortest in the 
insectivorous species. Only in Lhinopoma microphyllum and Aleqa- 
derma spasma has a very small cecum been found. 

The liver is characterised by the great size of the left lateral 
lobe, which oceasionally equals half the size of the whole organ; the 
right and left lateral fissures are usually very deep ; in the Mega- 
chiroptera (Harpyia excepted) the Spigelian lobe is ill-defined or 


648 CHIROPTERA 


absent, and the caudate is generally very large ; but in the Micro- 
chiroptera, on the other hand, the Spigelian lobe is very large, while 
the caudate is small, in most species forming a ridge only. The 
gall-bladder is generally well developed and attached to the right 
central lobe, except in the Rhinolophide, where it is connected with 
the left central. 

In most species the hyoids are simple, consisting of a chain of 
slender, elongated, cylindrical bones connecting the small basi-hyoid 
with the cranium, while the pharynx is short, the larynx shallow 
~ with feebly de- 

X veloped vocal 
cords, and 

ta guarded by a 

<< short, acutely- 

pointed epiglot- 
tis, which in 
some genera 
(Harpyia, Vam- 
pyrus) is almost 
obsolete. In 
Epomophorus, 
however, we 
find a remark- 
able departure 
fromthe general 
type. Thus 
the pharynx is 
long and very 
capacious ; the 
Fic. 299.—Head and neck of Epomophorus franqueti (adult male, aperture of the 
natural size). The anterior (a.ph.s) and posterior (p.ph.s) pharyngeal larynx is far re- 
sacs are opened from without, the dotted lines indicating the points 
where they communicate with the pharynx ; s, thin membranous septum moved from 
in middle line between the anterior pharyngeal sacs of opposite sides ; the fauces, and, 


s.m., sterno-mastoid muscle separating the anterior from the posterior . 7 
sac. (Dobson, Proc. Zool. Soc. 1881.) opposite fo it, 


atl ae = © 
SE | SS y 


(i 


opens a canal, 
leading from the narial chambers, and extending along the back 
of the pharynx ; the laryngeal cavity is spacious and its walls are 
ossified ; the hyoid bone is quite unconnected, except by muscle, 
with the cranium; the ceratohyals and epihyals are cartilaginous 
and greatly expanded, entering into the formation of the walls of 
the pharynx, and in the males of three species at least, supporting 
the orifices of a large pair of air-sacs communicating with the 
pharynx (Fig. 299). 

In extent, peculiar modifications, and sensitiveness the cutaneous 
system reaches its highest development in this order. As a sensory 
organ its chief modifications in connection with the external ear 


CHIROPTERA 649 


and with the nasal and labial appendages have been described when 
referring to the nervous system. It remains therefore to consider 
its relative development as part of the organs of flight. 

The extent and shape of the flyingmembranes depend mainly 
on the form of the bones of the anterior extremities, and on the 
presence or absence of the tail. Certain modifications of these 
membranes, however, are met with which do not depend on the 
skeleton, but are related to the habits of the animals, and to the 
manner in which the wing is folded in repose. 

These membranes consist of the ‘“antebrachial membrane,” 
extending from the point of the shoulder along the humerus and more 
or less of the forearm to the base of the pollex, the metacarpal bone 
of which is partially or wholly included in it; the “ wing-membrane,” 
which is spread out between the greatly elongated fingers, and 
extends along the sides of the body to the posterior extremities, 
generally reaching to the feet ; and the “interfemoral membrane,” 
the most variable of all, which is supported between the extremity 
of the body, the legs, and the calcar (Fig. 297). 

The antebrachial and wing-membranes are most developed in 
those species fitted only for aerial locomotion, which when at 
rest hang with the body enveloped in the wings; but in the family 
Emballonuride, and especially in the subfamily Molossine (the species 
of which are the best fitted of all Bats for terrestrial progression), 
the antebrachial membrane is reduced to the smallest size, and 
is not developed along the forearm, leaving also the pollex quite 
free, and the wing-membrane is very narrow and folded in repose 
completely under the forearm. 
The relative development of the 
interfemoral membrane has been 
referred to above in describing 
the caudal vertebre. Its small 
size in the frugivorous and sangui- « 
vorous species, in which its presence 
would be injurious as impeding 
their motions when searching for Fic. 300.—Frontal sac and nose-leaf in male 

5 and female of Hipposiderus larvatus. (Dobson, 
food as they hang suspended by Proc, Zoot. Soc. 1873.) 
their feet, is easily understood. 
Odoriferous glands and pouches opening on the surface of the outer 
skin are developed in many species, but in most cases more so in 
males than in females, and thus constitute secondary sexual char- 
acters, which will be referred to when treating of the peculiarities 
of certain species. 

All the fossil Chiroptera at present known are true Bats in every 
sense of the word, and therefore throw no light on the origin of the 
order. The earliest representatives of the order occur in beds 
of Upper Eocene (Lower Oligocene) age. 


650 CHIROPTERA 


The order is divided by Dobson into the suborders Mega- 
chiroptera and Microchiroptera. 


Suborder MEGACHIROPTERA. 


Frugivorous Bats, generally of large size. Crowns of molars 
smooth, marked with a longitudinal groove (cuspidate in Pteral- 
opex); bony palate continued behind the last molar, narrowing 
slowly backwards ; three phalanges in the index finger, the third 
phalanx generally terminated by a claw; sides of the ear-conch 
forming a complete ring at the base; tail, when present, inferior 
to (not contained in) the interfemoral membrane ; pyloric extremity 
of the stomach generally much elongated; the Spigelian lobe of 
the liver ill-defined or absent, and the caudate well developed. 

Limited to the tropical and subtropical parts of the eastern 
hemisphere. 

Mr. O. Thomas! considers that the ordinary type of molar 
dentition found in this suborder is a specialised adaptation from 
the cuspidate type of the Microchiroptera; the genus Pteralopex 
retaining the ancestral form of teeth. 


Family PTEROPODIDE. 


Since all the forms are included in this family its characters 
may be taken to be the same as those of the suborder. 

Subfamily Pteropodine.—Toneue moderate; molars well de- 
veloped. 


Epomophorus.-—Dentition ; iS c 4, p 3, mi; total 28 or 
26. Tail absent or very short, when present free from inter- 
femoral membrane; second digit of manus clawed; premaxille 
united. This genus includes some seven species inhabiting Africa 
south of the Sahara. The head is large and long, and the lips are 
expansible, and frequently with peculiar folds. The ears have a 
white tuft of hair on the margin ; and in the males of most species 
there are large glandular pouches in the skin of the side of the 
neck near the shoulder, from the mouth of which project long and 
coarse yellowish hairs, forming tufts on the shoulders, from which 
the genus takes its name. Another male secondary sexual 
character consists in the présence of a pair of large air-sacs 
extending outwards on each side from the pharynx beneath the 
integument of the neck, in the position shown in Fig. 299. These 
sacs are evidently capable of being greatly distended at the will of 
the animal, and their inflation probably occurs under the same 

1 Proc, Zool. Soc. 1888, p. 473. 
* Bennett, Trans. Zool. Soc. vol. ii. p. 38 (1835). 


PTEROPODIDA 651 


circumstances that the wattles of male gallinaceous birds swell up, 
namely, when engaged in courting the females. Other remarkable 
conditions in which these Bats appear to differ from all other species 
occur in the form of the hyoid bones and larynx. These Bats 
appear to live principally on figs, the juicy contents of which 
their large lips and capacious mouths enable them to swallow 
without loss. 

Pteropus.-_Dentition: i 3, ¢ 4, p 3, m 2; total 34. This 
genus has more than forty species, and thus includes more than 
half the members of the family. All are of large size, and the 
absence of a tail, the long pointed 
muzzle (Fig.301),and the woolly 
fur covering the neck render 
their recognition easy. They 
are commonly known as “Flying 
Foxes,” or Fox-Bats; and one 
of the species (P. edulis) in- 
habiting Java measures 5 feet 
across the fully extended wings, 
and is thus the largest known 
species of the order. All the 
species closely resemble one ne 
another in dentition, and are Fic. 301.—Head Ob BOX Bat (Pleropus personatus). 
mainly distinguished by the From Gray, Proc. Zool. Soc. 1866. 
form of the ears and the quality of the fur. P. scapulatus, from 
North-East Australia, approaches the species of the second sub- 
family in the remarkable narrowness of its molars and premolars. 

The range of this genus extends from Madagascar and the 
neighbouring islands through the Seychelles to India, Ceylon, 
Burma, the Malay Archipelago, Southern Japan, New Guinea, 
Australia, and Polynesia (except the Sandwich Islands, Ellice’s 
Group, Gilbert’s Group, Tokelau, and the Low Archipelago). Of 
the islands inhabited by it some are very small and remote from 
any continent, such as Savage Island in the South Pacific and 
Rodriguez in the Indian Ocean. Although two species inhabit the 
Comoro Islands, which are scarcely 200 miles from the African 
coast, not a single species is found in Africa; but in India, 
separated by thousands of miles of almost unbroken ocean, a 
species exceedingly closely allied to the common Madagascar 
Fox-Bat is abundant. The Malay Archipelago and Australia are 
their headquarters; and in some places they occur in countless 
multitudes. Mr. Macgillivray remarks of P. conspicillatus: ‘On 
the wooded slope of a hill on Fitzroy Island I one day fell in with 
this Bat in prodigious numbers, looking while flying in the bright 
sunshine (so unusual for a nocturnal animal) like a large flock of 

1 Geoflroy, Ann. du Muséum, vol. xv. p. 90 (1810).—£zx. Brisson. 


ORs CHIROPTERA 


rooks. On close approach a strong musky odour became apparent, 
and a loud incessant chattering was heard. Many of the branches 
were bending under their load of Bats. some in a state of inactivity, 


Fic. 302.—Female and young of Xantharpyia collaris. (From $ 
Proce. Zool. $2:..1870, p. 127.) 


suspended by their hind claws, others scrambling along among the 
boughs, and taking to wing when disturbed.” 

Meathar puta. 1_Dentition as in Pt: ys, but a short tail present, 
and the fur on the back of the neck similar to that of the body. 


7 


Gray, List, Spee. Mamm. Brit. Mus. pp. 37, 38 1843°: Sun. Cyno- 
nycteris. 


PTEROPODIDA 653 


This genus is represented by some nine species, which have a 
distribution very similar to that of Pteropus, except that they 
extend into Africa, and are not found in Australia and Poly- 
nesia. X. egyptiaca inhabits the chambers of the Great Pyramid 
and other deserted buildings in Egypt, and is probably the species 
so generally figured in Egyptian frescoes. Fig. 302 exhibits an 
African species of this genus in the attitude assumed by the Fox- 
Bats when at rest. 

Boneia.1.—This genus, as represented by B. bidens of Borneo, 
differs from Xantharpyia in having only a single pair of upper 
incisors. 

Cynopterus.2—Dentition : 4 aaa c+, p 3,m 2; total 32 or 30. 
Muzzle short, grooved like Pteropus in front; tail and fur generally 
as in Xantharpyia, but the former sometimes wholly absent. This 
genus, with seven species, is almost limited to the Oriental region. 
C. marginatus is very common in India, and extremely destructive 
to ripe fruit of every description. Dr. Dobson states that “he 
gave to a specimen of this Bat obtained at Calcutta a ripe banana, 
which, with the skin removed, weighed exactly 2 ounces; the 
animal immediately, as if famished with hunger, fell upon the 
fruit, seizing it between the thumbs and the index fingers, and took 
large mouthfuls out of it, opening the mouth to the fullest extent 
with extreme voracity. In the space of three hours the whole 
fruit was consumed. Next morning the Bat was killed, and found 
to weigh one ounce, or half the weight of the food eaten in three 
hours. Indeed the animal when eating seemed to be a kind of 
living mill, the food passing from it almost as fast as devoured, 
and apparently unaltered, eating being, as it were, performed only 
for the pleasure of eating.” 

Harpyia.2—Dentition : 14, ¢ 4, p #,m2; total 24. Premaxille 
well developed and united in 
front ; facial bones much ele- 
vated above the margin of 
the jaw, nostrils tubular (Fig. 
303); body and limbs as in 
Cynopterus. Includes two 
species from the Austro- 
Malayan subregion, readily 
recognised by the peculiar 
tubular and projecting ne ae 
nesirils, as shows im iho 7°" ee ee 
accompanying woodcut. 

Cephatotes.-—Dentition: i +, ¢ 4, p 3, m 3; total 28. Pre- 

1 Jentink, Notes Leyd. Mus. vol. i. p. 117 (1879).—Amended. 2 F. Cuvier, 


Dents des Mammiferes, p. 39 (1825). 3 Tlliger, Prodromus Syst. Mamm. et 
Avium, p. 118 (1811). 4 Geoffroy, Ann. du Muséum, vol. xvi. p. 99 (1810). 


654 CHIROPTERA 


maxille separate in front; nostrils simple; muzzle short; index 
finger without a claw; tail short. Includes one species, having 
the same distribution as Harpyia. The wing-membrane arises from 
the middle line of the back, to which it is attached by a longitudinal 
very thin process of the integument; the wings are quite naked, 
but the back covered by them is clothed with hair. 

Pteralopex.'—External characters as in Pteropus ; ears short and 
hairy; wings arising from the middle line of the back. Muzzle 
very short; plane of orbit directed more upwards than in Pteropus ; 
orbit surrounded by bone ; sagittal crest strongly developed. Teeth 
cuspidate; upper incisors with broad posterior ledges; upper 
canine short and thick, with a stout secondary cusp in the middle 
of the posterior border, and two smaller postero-internal basal 
cusps; cheek-teeth short and broad, with their anterior and 
posterior basal ledges so developed and the main cusps so nearly 
conical as to obliterate the longitudinal grooving of Pteropus. 
Lower incisors very disproportionate, the outer pair being nearly 
twenty times the bulk of the inner ; lower canine stout, with a simple 
posterior basal ledge. Represented by P. atrata of the Solomon 
Islands. As already mentioned, Mr. Thomas regards the dentition 
of this genus as the most generalised type found in the suborder. 

Subfamily Carponyeteriinz.—Facial part of skull much pro- 
duced ; molars narrow, and scarcely raised above the gum; and 
the tongue exceedingly long, attenuated in the anterior third, and 
armed with long recurved papille near the tip. 

Notopteris.s—Dentition : 7 2, ¢ 4, p 2, m 2; total 28. Index 
finger without a claw; wings arising from the middle line of the 
back ; tail long; first upper premolar long, with two roots. The 
single representative of the genus, NV. macdonaldi, inhabits the Fiji 
Islands, Aneiteum Island, and New Guinea. It is at once distin- 
guished from all other Bats of this family by the length of its tail, 
which is nearly as long as the forearm. 

Eonycteris.2—Dentition: 1 2,¢ 4, p $,m 3; total 34. First upper 
premolar small, with a single root. This genus is likewise repre- 
sented by a single species (Z. spelwa), from the Farm Caves, Moul- 
mein, Burma, which has somewhat the appearance of Xantharpyia ; 
but the absence of a claw to the index finger and the characteristic 
tongue and teeth at once distinguish it. 

Carponycteris* and Melonycteris,® each with a single species, are 
closely allied ; the index finger in both has a claw, and the number 
of the teeth is the same as in Konycteris. Carponycteris minima is 


1 0. Thomas, Ann. Jfag. Nat. Hist. ser. 6, vol. i. p. 155 (1888). 2 Gray, 
Proc. Zool. Soc. 1859, p. 36. 3 Dobson, Journ. As. Soe. Bengal, vol. xlii. 
p. 204 (1873). + New name: Syn. Iacroglossus, F. Cuvier, Dents des 


Mammiferes, p. 40 (1825). Preoccupied by Macroglossum, Scopoli, 1777. 
5 Dobson, Proc. Zool. Soc. 1877, p. 119. 


JMICROCHIROPTERA 655 


the smallest known species of the suborder, being much smaller than 
the common Noctule Bat of Europe, and its forearm scarcely longer 
than that of the Long-eared Bat. It is nearly as common in certain 
parts of India as Cynopterus marginatus (compared with which it is 
proportionally equally destructive to fruit), and extends eastward 
through the Malay Archipelago as far as New Ireland, where it is 
associated with Melonycteris melunops, distinguished from it by its 
larger size and the total absence of the tail. , 

Nesonycteris..—Dentition : 2 2,¢ 4, p 3, m2; total 32. Allied to 
Melonycteris, but distinguished by the absence of the inner pair of 
lower incisors, and of a claw to the index finger. Tail wanting. 
Represented by VV. woodfordi, of the Solomon Islands. 

Callinycteris.°—Dentition: 7 3, ¢4, p 3, m 3; total 32. Allied 
to the preceding, but with a short tail; no claw to index. One 
species from Celebes. 

Trygenycteris2=—Dentition: i2,¢ 4,p 3%, m 3; total 34. No 
external tail; a claw on index. One species from West Africa. 


HH 


Suborder MICROCHIROPTERA. 


Insectivorous (rarely frugivorous or sanguivorous) Bats, of com- 
paratively small size. Crowns of molars acutely cusped, marked 
by transverse grooves ; bony palate narrowing abruptly, not con- 
tinued backwards laterally behind the last molar ; one rudimentary 
phalanx (rarely two phalanges or none) in the index finger, which 
is never terminated by a claw; outer and inner sides of ear-conch 
commencing inferiorly from separate points of origin; tail, when 
present, contained in the interfemoral membrane, or appearing upon 
its upper surface ; stomach simple (except in the Desmodont Phyl- 
lostomutidw); Spigelian lobe of the liver very large, and the caudate 
generally small.. Inhabit the tropical and temperate regions of 
both hemispheres. The members of this suborder may be divided 
into two sections. 


Section VESPERTILIONINA. 


Tail contained within the interfemoral membrane; the middle 
pair of upper incisors never large, and separated from each other 
by a more or less wide space. Middle finger with two osseous 
phalanges only (except in Myxopoda aurita, Thyroptera tricolor, and 
Mystacops tuberculatus). First phalanx of the middle finger extended 
(in repose) in a line with the metacarpal bone. 


1 Q. Thomas, -lnn. Mag. Not. Hist. ser. 5, vol. xix. p. 417 (1887). 

2 Jentink, Notes Leyd. Mus. vol. xi. p. 209 (1889). 

3 New name: Syn. IWegaloglossus ; Pagenstecher, J. B. Mus. Hamburg, vol. 
ii, p. 125 (1885). Preoceupied by Ieyaglossa, Rond., 1865. 


656 CHIROPTERA 


Fumily RHINOLOPHID.E. 


In all the species of this family the nasal appendages are highly 
developed, and surround the sides of the nasal apertures, which are 
situated in a depression on the upper surface of the muzzle; the 
ears are large and generally separate, without trace of a tragus ; the 
premaxille are rudimentary, suspended from the nasal cartilages, 
and supporting a pair of very small incisors ; the molars have acute 
W-shaped cusps; the skull is large, and the nasal bones which support 
the large nasal cutaneous appendages are much expanded vertically 
and laterally ; in the females a pair of teat-like appendages are 
found in front of the pubis; and the tail is long and produced to 
the posterior margin of the interfemoral membrane. This family is 
found in the temperate and tropical parts of the eastern hemisphere. 

From whatever point of view the Lhinolophide may be con- 
sidered, they are evidently the most highly organised of insect- 
ivorous Bats. In them the osseous and cutaneous systems reach the 
most elaborate development. Compared with those of the present 
family the bones of the extremities and the flying-membranes of 
other Bats appear coarsely formed, and even their teeth seem less 
perfectly fitted to crush the hard bodies of insects. The very com- 
plicated nasal appendages, which evidently act as delicate organs of 
special perception, here reach their highest development, and the 
differences in their form afford valuable characters in the discrimi- 
nation of the species, which resemble one another very closely in 
dentition and in the colour of the fur. 

Subfamily Rhinolophine.—First toe with two, other toes with 
three, phalanges each ; ilio-pectineal spine 
not connected by bone with the antero- 
inferior surface of the ilium. 

Lhinolophus.\—Dentition: i 3, ¢ 1, p 3, 
m 3; total 32. Noseleaf (Fig. 304) with a 
central process behind and between the nasal 
orifices, posterior extremity lanceolate, anti- 
tragus large. Includes more than twenty 
species. J?. /uctus, in which the forearm has a 
length of 3 inches, isthe largest species, inhabit- 
ing elevated hill tracts in India and Malayana ; 

Fra. 804.—Head of Indian £2. Aapposiderus of Europe, extending into 
Horse-shoe Bat (hinolophus South England and Ireland, forearm 1:5 
mitratus). (From Dobson, . : 

Monogr. -Asiat. Chiropt.) inches, is one of the smallest ; and 2. Serrum- 

equinuin, with the forearm 2°3 inches in 
length, represents the average size of the species, which are mainly 
distinguished from one another by the form of the nose-leaf. The 


' Geoffroy, Nour, Dict. d' Hist. Nat. vol. xix, p. 383 (1803). 


RHINOLOPHIDA 657 


last-named species extends from England to Japan, and southward to 
the Cape of Good Hope. The genus is represented in the Himalaya 
by the closely allied /. fragutus, distinguished by having three 
vertical grooves on the lower lip, in place of the single groove found 
in R. ferrum-cquinwn.  Rhinolophus is vepresented in the Upper 
Kocene Phosphorites of Central France by 2. antiguas and J. 
dubivs ; the former appears to have the same dental formula as in 
the existing species, but differs slightly in the structure of some of 
the lower molars, so that it is separated generically by some writers 
under the name of Pseudorhinolophus. The face is also longer than 
in existing forms, and there are certain differences in the structure 
of the skull. -f/estor, from the same deposits, differs from Lhino- 
lophus by the extreme shortness of the nasal region. Pulwenyeteris, 
from the Lower Miocene of France, is said to be allied to Ihino- 
lophus, but the premolars are 8, and the limb bones are stated to 
resemble those of Jo/ossus. 

Subfamily Hipposiderinzs.—Toes equal, of two phalanges each ; 
ilio-pectineal spine united by a bony isthmus with a process derived 
from the antero-inferior surface of the ium. 

Hipposiderus\—Dentition : i3,¢4, p ~y*, m 3; total 30 or 2s. 
Tail well developed. This genus, of which more than twenty 
species have been described, differs 
from Phinolophus in the form of the 
nose-leaf, which is not lanceolate 
behind and is unprovided with a cen- 
tral process covering the nostrils. The 
largest species, //. armiger, appears 
to be the most northerly, having 
been taken at Amoy in China, and 
in the Himalaya at an elevation of 
5,500 feet. Many of the species are — 
provided with a peculiar frontal sac 1° ce Tia 
behind the nose-leaf, rudimentary is77. Viernes 
in’ females (Fig. 305), which the 
animal can evert at pleasure; the sides of this sac secrete a 
waxy substance, and its extremity supports a pencil of straight 
hairs. 

-lnthops°—Like Hipposiderus, but with the tail rudimentary, 
consisting merely of three or four vertebra: hidden in the base of 
the interfemoral membrane. Nose-leaf very complicated, its upright 
transverse portion emarginate above, and the projections rounded 


1 Gray, Proe, Zool. Soc. 1834, p. 53, The Bats of this genus are usually 
desevibed as Phyllorkina, but this uso has been shown to be incorrect ; see Blan- 
ford, Proc, Zool, Soc. USS7. p. 637. 

2-0, Thomas, dan. Mog, Vat. Hist. sev. 6, vol. i. p. 156 (1888). 

12 


658 CHIROPTERA 


and hollowed behind, and their substance quite thin. Premolars 2. 
Represented by -/. ornatus of the Solomon Islands. 

Mr. O. Thomas, the describer of this Bat, remarks that it is 
evidently more nearly allied to the preceding than to the succeeding 
genera, although it agrees with Cwlops in the rudimentary tail. 

Ehinonycteris? and Tricnops.2—These are two allied genera with 
well-developed tails; the former 
being represented by FR. auruntia 
from Australia, and the latter by 
T. persicus from Persia and Eastern 
Africa. Tvrienops (Fig. 306) is 
characterised by the remarkable 
form of its nasal appendages and 
ears, and the presence of a peculiar 
osseous projection from the 
proximal extremity of the second 
phalanx of the fourth finger. 

Cclops.-—This genus is known 
only by a single species, C. frithi, 
from the Bengal Sunderbans, 


Fic. 306.—Head of Tricenops persicus. x 2. 4 7 
(From Dobson, Monogr. Asiat. Chiropt.) Java, and Siam (in the roof of 


the great pagoda at Laos); and 
is distinguished, not only by the form of its nose-leaf, but also by 
the great length of the metacarpal of the index finger, as well as 
by the shortness of the calcar and interfemoral membrane and the 
rudimental tail. 


Family NYCTERIDA. 


This small family, including only two genera of Bats of peculiar 
aspect, limited to the tropical and subtropical parts of the eastern 
hemisphere, is distinguished from the Rhinolophide by the presence 
of a distinct tragus to the ear, and by the premaxille being cartila- 
ginous or small and separated from one another in front by a dis- 
tinct space. 

Megaderma.A—Dentition: i$, ¢ 4, p +, m 3; total 28 or 26. 
This genus, which is represented by five species, is readily recognised 
by the absence of upper incisors, the cylindrical narrow muzzle 
surmounted by an erect naked cutaneous nose-leaf, the base of 
which conceals the nasal orifices, by the immense connate ears with 
large bifid tragi, and by the great extent of the interfemoral 
membrane, in the base of which the very short tail is concealed. 
M. gigas (Fig. 307), from Central Queensland (length of forearm 4°2 


1 Gray, Proc. Zool. Soc. 1847, p. 16, ? Dobson, Journ. As. Soe. Bengal, 
vol. xl. p. 455 (1871). 3 Blyth, Journ. As. Soe. Bengal, vol. xvii. p- 251 (1848). 
* Geoffroy, Ann. du Muséum, vol. xv. p. 197 (1810). 


NYCTERIDAE 659 


inches), is not only the largest species of the genus but also of the 
suborder. Jf. lyra, common in India (forearm 2-7 inches), has been 
caught in the act of sucking the blood, while flying, from a small 
species of Vesperugo, which it afterwards devoured, so that it is 
probable that the Bats of this genus do not confine themselves to 


_Fia. 307.—Megaderma gigas. x 3. (From Dobson, Proc. Zool. Soc. 1880.) 


insect prey alone, but also feed, when they can, upon the smaller 
species of Bats and other small mammals. 

The Oriental J/. spasma and M. lyra differ from the Ethiopian 
M. cor and M. frons in having two upper premolars instead of one, 
and also in the shape of the frontals and nasals. 

Nycteris.1—Dentition : 1 2, ¢4,p 4, m3; total 32. This genus, 
of which there are seven species, differs so much from Megaderma 
that it may be considered the type of a separate subfamily. As in 
that genus, the frontal bones are deeply hollowed out and expanded 
laterally, the muzzle presents a similar cylindrical form, and the 
lower jaw also projects, but the single elevated nose-leaf is absent, 
and instead of it the face is marked by a deep, longitudinal, sharp- 
edged groove extending from the nostrils (which are on the upper 


1 Geoffroy, Vowv. Dict. d Hist, Nat. vol. xv. p. 501 (1803). 


660 CHIROPTERA 


surface of the muzzle, near its extremity) to the low band connect- 
ing the bases of the large ears; the sides of this depression being 
margined as far back as the eyes by small horizontal cutaneous 
appendages. All the species resemble one another closely, and are 
mainly distinguished by the form of the tragus and the size and 
relative position of the second lower premolar. With the exception 
of .V. javanica, they are all limited to the Ethiopian region. 


Family VESPERTILIONIDE. 


Nostrils opening by simple crescentic or circular apertures at 
the extremity of the muzzle, not surrounded by distinct foliaceous 
cutaneous appendages ; premaxillz small, lateral, and separated by 
a wide space in front; tragus distinct. In addition to these char- 
acters, it may be observed that the skull is of moderate size, the 
nasal and frontal bones not being much extended laterally or vertic- 
ally, nor furrowed by deep depressions. The number of incisors 
varies from 3 to 4, rarely (in Antrozous only) 4, premolars 3, or 2, 
or 3, rarely (in Vesperugo noctivaguns of North America) 3; the 
upper incisors are small, separated by a wide space in the middle 
line, and placed in pairs or singly near the canine; the molars are 
well-developed, with acute W-shaped cusps. 

This family, which may be regarded as occupying a central 
position in the suborder, includes the common simple-faced Bats of 
all countries, of which the well-known Pipistrelle and the Whiskered 
Bat (Vespertilio mystacinus) may be taken as familiar types, and its 
species number more than 150, or considerably more than one-third 
the total number of the known species of the entire order. The 
various genera may be conveniently grouped into the Plecotine, 
Fespertilionine, Miniopterine, and Thyropterine divisions. 

In the Pleeotine division, of which the common Long-eared Bat 
(Plecotus auritus) is the type, the crown of the head is but slightly 
raised above the face-line, the outermost upper incisor is close to 
the canine, and the nostrils are margined behind by grooves on the 
upper surface of the muzzle, or by rudimentary nose-leayes ; the 
ears also are generally very large and united. 

Plecotus.1—Dentition : i 3, ¢ 4, p 3, m 3; total 36. Outer 
margin of ear-conch ending abruptly near the angle of the mouth, 
the inner margin with a more or less prominent rounded projection 
directed inwardly above the base ; tragus very large, tapering up- 
wards, with a lobe at the base of its outer margin, rounded, and 
placed half horizontally. This genus is represented by the Euro- 
pean Long-eared Bat (P. auritus), and P. macrotis, restricted to 
North America. The latter is distinguished by the great size of 


? Geoffroy, Deseript. de l Egupte, vol. ii. p. 112 (1812). 


VESPERTILIONID A: 661 


the glandular prominences of the sides of the muzzle, which meet 
in the centre above and behind the nostrils. P. awritus extends 
over the greater part of the Palearctic region, occurring in Ireland 
in the west and the Himalaya in the east. 

Synotus.|—Dentition : 1 2, ¢ 4, p 2, m3; total 34. This genus 
is distinguished from the preceding by the loss of one lower pre- 
molar and by the outer margin of the ear being carried forwards 
above the mouth and in front of the eye; it includes the European 
Barhastelle Bat (S. barbastellus) and S. darjilingensis from the Hima- 
laya. 

Otonycteris.2— Dentition: 1 4, ¢ 4, p 4, m 4; total 30. The 
reduction in the number of upper incisors readily characterises this 
genus, which appears to connect the typical representatives of the 
section, through Scotophilus, with the Vespertilionine division. It is 
represented by a single species, 0. hemprichi, from North Africa and 
the Himalaya. 

Nyctophilus.2—Dentition: i 4, ¢4, p 4, m3; total 30. This 
and the following genera are distinguished from all the preceding 
by the presence of a rudimentary nose-leaf. The present genus 
contains IV. timoriensis of the Australian region, and N. amécrotis of 
New Guinea. 

Antrozvus.A—Dentition: 11, ¢ 4, p $, m4; total 28. Readily 
distinguished from the other members of the whole family by 
having but two lower incisors, and from the other species of the 
section by the separate ears. The single species, 4. pallidus, in- 
habits California. 

The /espertilionine division includes some nine-tenths of all the 
representatives of the family. They are distinguished from the 
preceding section by the simple nostrils, opening by crescentic or 
circular apertures at the extremity of the muzzle, the generally 
smill size of the ears, and the absence of grooves on the forehead. 

Vesperugo.,—Dentition : 7 mo ci,p ee m #; total 34, 30, 
or 36. This large genus comprises about one-third of the section, 
and is divided into groups or subgenera, according to the number 
of premolars and incisors; the latter varying from % to 4 in the 
subgenera Stotuzous and Ihogeé’sss, and the premolars from 3 to } (in 
the subgenus Lusionycteris 2). The Bats of this genus are generally 
easily distinguished by their comparatively thickly formed bodies, 
flat broad heads and obtuse muzzles, short, broad, and triangular 
obtusely-pointed ears, obtuse and usually slightly incurved tragus, 
short legs, and by the presence in most species of a well-developed 
post-calcaral lobule. This lobule (which is supported by a. cartil- 

1 Keyserling and Blasius, I irbelthiere Europ. p. 55 (1840). ? Peters, 
Monatsber, Ak, Berlin, 1859, p, 222. 3 Leach, Trans. Linn. Soc, vol. xiii. 
p. 78 (1822), + Allen, Proc. Ac. Nat. Sci. Philacl. 1862, p. 247. 

5 Keyserling and Blasius, Wieginann’s Archiv, 1839, p. 312. 


662 CHIROPTERA 


aginous process derived from the calcar) may act as a kind of 
adhesive disc in securing the animal’s grasp when climbing over 
smooth surfaces. J ¢sperugo probably contains the greatest number 
of individuals among the genera of Chiroptera, and, “with the excep- 
tion of J ¢spertilic, its species have also the widest geographical 
range, being almost cosmopolitan; and one of the species, the well- 
known Serotine (J” [Vesperus] serotinus) is remarkable as the only 
species of Bat known to inhabit both the Old and the New World ; 
one (J~ barealis) has been found close to the limits of the Arctic 
circle, and another (J~ imawellanicus) inhabits the cold and desolate 
shores of the Straits of Magellan, being doubtless the Bat referred 
to by Mr. Darwin in the Vaturalist's Foyase. The Common Pipis- 
trelle (J. pipistrellus), ranging over the greater part of the Palearctic 
region, is the best known species. 

Chalinolobus’—This genus agrees with Tesperugo in the dental 
formula, but is readily distinguished by the presence of a well- 
defined lobe projecting near the angle of the mouth from the lower 
lip, and by the unicuspidate first upper incisor. The species fall 
into two subgenera—Chalinolobus proper, with p 3, represented by 
C. morio from New Zealand, Tasmania, and Australia, and three 
other species from Australia; and Glauconycteriz, with p 4, limited 
to Southern and Equatorial Africa, and known by C. argentatus and 
two other species, the Bats of this subgenus being especially remark- 
able for their peculiarly thin membranes, traversed by very distinct 
reticulations and parallel lines. 

Scotophilus.*—Dentition: i 4, ¢ 4, p $, m 3; total 30. This 
genus comprises a comparatively small number of species nearly 
allied to 7% esperugo, and some of which 
approach so closely to the aberrant types of 
the latter ranged under the subgenus Svotozeus, 
as to render the definition of the present genus 
almost impossible. The species are restricted 
to the tropical and subtropical regions of the 
eastern hemisphere, though widely distributed 
within these limits. The more typical species 
nn A Head of “efe- are distinguished especially by the single pair 
philus¢marginatus. (Dobson, f > So o < 
Monogr. Asiat. Chirapt.) of unicuspidate upper incisors separated by a 

wide space and placed close to the canines, by 
the small transverse first lower premolar squeezed in between the 
canine and second premolar, and, generally, by their conical nearly 
naked muzzles and remarkably thick leathery membranes. S. kuhli 
is probably the commonest species of Bat in India, and appears 
often on the wing even before the sun has touched the horizon, 


Peters, Monatsber. Ak. Berlin, 1866, p. 672. 
Leach, Trans. Linn. Soe. vol. xiii. p. 71 (1822). 


1 
* See O. Thomas, Ann. Mus. Genova (2). vol. ix. pp.84-83 (1890). 


VESPERTILIONIDAS 663 


especially when the white-ants are swarming, feeding eagerly upon 
them as they rise in the air. S. gigas, from Equatorial Africa, 
with the forearm measuring 3°4 inches, is by far the largest 
species. 5. albofuscus, from the Gambia, which is readily distin- 
guished from the other species by its white wings, is an aberrant 
form, in which the lower premolars are long and not crowded 
together, and thereby so closely resembles Vesperugo (Scotozous) 
dormeri as to render it almost impossible to distinguish Scotophilus 
and Vesperugo. The figured species is from India. 
Nycticejus1—This genus, with the same dental formula as 
Scotophilus, is distinguished by the first lower premolar not being 
squeezed in between the adjoining teeth, and by the comparatively 
much greater size of the last upper molar. It includes only the 
common North American N. humeralis (crepuscularis), a small Bat 
scarcely larger than the Pipistrelle. It seems, however, as pointed 
out by Mr. O. Thomas, that the discovery of Scotophilus albofuscus 
renders the generic distinctness of Nycticejus no longer tenable, and 
that the species should be known as de! humeralis. 
Atalapha.2—Dentition: + 4, ¢ 1, p >, m4; total 32 or 30. 
The five species of this genus, which are confined to the New 
World, are generally characterised by the interfemoral membrane 
being more or less covered with hair (in the two commonest species, 
A. noveboracensis and A. cinerea, wholly covered), and by the peculiar 
form of the tragus, which is expanded above and abruptly curved 
inwards. These species have two upper premolars, of which the 
first is extremely small and quite internal to the tooth-row. 
Harpyiocephalus.’—Dentition : 23, ¢,4,p2,m 3; total 34. This 
genus includes some eight species of small Bats distinguished by 
their prominent tube-like nostrils and hairy interfemoral membrane. 
H. swillus, from Java and neighbouring islands, is the best-known 
species, and another closely allied (7. hilgendorfi) has been described 
by Professor Peters from Japan. The remaining six species are 
known only from the Himalaya and Tibet. All appear to be 
restricted to the hill tracts of the es in wich they are found. 
Vespertilio.A—Dentition: 1 2, ¢ 4, p 3, m4; total 38. Next 
to Vesperugo, this genus includes by far the largest number of species, 
amounting to over forty; it has, however, rather a wider geo- 
graphical distribution in both hemispheres, one species at least 
being recorded from the Navigators’ Islands. The species are 
easily recognised by the peculiar character of the upper incisors, 
the crowns of which diverge from each other ; by the large number 
of premolars, of which the second upper one is always very small ; 


1 Rafinesque, Journ. de Physique, vol. lxxxviii. p. 417 (1819). ®? Rafinesque, 
Précis des Decowvértes et Trav. Somiol. p. 12 (1814). 3 Gray, Ann. Mag. Nat. 
Hist, vol. x. p. 259 (1842) 4 Linn, Syst. Nat. 12th ed. vol. i. p. 46 (1766). 


664 CHIROPTERA 


and by the oval elongated ear and narrow attenuated tragus. In 
the British Isles this genus is represented by four species, viz. 
Bechstein’s Bat (V. bechsteini) ; the Reddish-Gray Bat (V. nattereri), 
of very local occurrence ; Daubenton’s Bat (V. daubentoni) ; and the 
Whiskered Bat (V. mystacinus). 

Cerivoula—This genus, which has the same dental formula 
as Vespertilio, is distinguished by the parallel upper incisors, 
and the comparatively large size of the second 
upper premolar. Some ten species have been 
described from the Ethiopian and Oriental 
regions, of which C. picta, from India and the 
Indo-Malayan subregion, is the best-known, 
being well characterised by its brilliantly 
coloured orange fur and conspicuously marked 
membranes, which are variegated with orange 
and black. This genus includes the most deli- 
cately formed and most truly insectivorous, 
tropical, forest-haunting Bats, which appear to 
SiS _~—s stand as regards the species of Vespertilio in a 

einai position similar to that occupied by Chalinolobus 
Fic. 309.—Side and : 
front views of the heaq With respect to Vesperugo. 
of Cerivoula hardwickei. The Miniopterine division includes only two 
eae Monogr. Asia. genera, and is characterised by the great eleva- 

“a tion of the crown of the head above the facial 
line, and by the upper incisors being separated from the canine 
and also in the middle line. 

Natalus.-—This genus, while having the divisional characters 
mentioned above, agrees in the dental formula and its general 
external form with Cerivoula, from 
which it is distinguished by the 
short triangular tragus. It in- 
cludes three species, restricted to 
South and Central America and 
the West Indies; the head of N. 
mucropus being shown in Fig. 310, 

Miniopterus.’—Dentition : i 3, 
¢4,p 2, m 3; total 36. In 
addition to the difference in the 
number of the teeth, this genus is 
distinguished by the shortness of 
the first phalanx of the middle finger and the great length of the 
tail, which is wholly contained within the interfemoral membrane ; 
it includes four species, restricted to the eastern hemisphere. Of 


1 Gray, Ann. Mag. Nat. Hist, vol. x. p. 258 (1842), Kerivoula. 
° Gray, Mag. Zool. Bot. vol. ii. p. 496 (1838). 
3 Bonaparte, Fauna Italica, fase. xxi. (1837). 


Fig. 310.—Head of Natalus mieropus. x8. 
(Dobson, Proc. Zool. Soc. 1880.) 


VESPERTILIONIDA 665 


these the best-known, J. schreibersi, is very widely distributed, being 
found almost everywhere throughout the tropical and warmer 
temperate regions of the eastern hemisphere; specimens from 
Germany, Madagascar, Japan, and Australia differing in no 
appreciable respect from one another. 

The last or Thyropterine division, which likewise comprises only 
two genera, is characterised by the presence of an additional osseous 
phalanx in the middle finger and an equal number of phalanges in 
the toes, and also by peculiar accessory clinging organs attached 
to the extremities. 

Thyropteru..Dentition : i 2, ¢ 1, p 8, m3; total 38. In the 
single species YT. tricolor of Brazil the clinging organs have the 
appearance of small, circular, pedunculated, hollow discs (Fig. 311), 
resembling in miniature the sucking cups of cuttle-fishes, and are 


Fic. 311.—Suctorial discs in Thyroptera tricolor. a, Side and }, concave surface, of thumb- 
dise ; c, foot with disc, and calear with projections (all much enlarged). Dobson, Proc. Zool. 
Soc. 1876. 


attached to the inferior surfaces of the thumbs and soles of the 
feet. With these the animal is enabled to maintain its hold when 
creeping over smooth vertical surfaces. 

Muyxopoda.2—The second genus is likewise represented only by 
a single species—AJf. aurita of Madagascar—and is distinguished 
from the preceding by the characters of the teeth and the form of 
the ears. The whole inferior surface of the pollex supports a 
large sessile horse-shoe-shaped adhesive pad, with the circular 
margin directed forwards and notched along its edge, and a 
smaller pad occupies part of the sole of the foot. 

Fossil Vespertilionidee.—It is not improbable that lesperugo is 
represented in the Upper Eocene of the Paris basin by J” pavi- 
siensis, which appears to be allied to 7”. serotinu, although it has 
been regarded by some writers as generically distinct, under the 
name of Nyctitherium.  Tesperugo (Nyctitherium) also occurs in the 
Bridger Eocene of the United States; Nyctilesies from the same 


1 Spix, Sim. and Vesp. Bresil, p. 61 (1828). 
2 A. Milne-Edwards, Bull. Soc. Philom. sér. 7, vol. ii. p. 1 (1878). 


666 CHIROPTERA 


deposits being an allied extinct genus. A number of European 
Miocene species have heen referred to Vespertilio, but the term in 
these cases must be used in a somewhat wide sense. Vespertiliavus, 
of the Phosphorites of Central France, differs oe Vespertilio in the 
proportions of its premolars. 


Section EMBALLONURINA. 


Tail perforating the interfemoral membrane and appearing on 
its upper surface, or produced considerably beyond the truncated 
membrane ; the middle pair of upper incisors generally large and 
close together. 


Family EMBALLONURID. 


First phalanx of the middle finger folded (in repose) on the 
dorsal surface of the metacarpal bone (except in Noctilio and 
Mystacops). Nostrils opening by simple circular or valvular aper- 
tures at the extremity of the muzzle, not surrounded or margined 
by foliaceous cutaneous appendages ; tragus distinct. 

The Emballonwride are generally easily distinguished by the 
peculiar form of the muzzle, which is obliquely truncated, the 
nostrils projecting more or less in front beyond the lower lip; by 
the first phalanx of the middle finger being folded in repose 
forwards on the upper surface of the metacarpal bone ; by the tail, 
which either perforates the interfemoral membrane or is produced 
far beyond it; and by the upper incisors, which are generally a 
single pair separated from the canine and also in the middle line. 
The family is cosmopolitan like the Vespertilionide, but rarely 
extends north or south of the thirtieth parallel of latitude. 

Subfamily Emballonurine.—Tail slender, perforating the inter- 
femoral membrane, and appearing upon its upper surface, or 
terminating in it; legs long, fibula very slender ; upper incisors 
weak. 

In the Furipterine division the tail terminates in the interfemoral 
membrane ; the crown of the head is greatly elevated above the 
face-line ; the thumb and first phalanx of the middle finger are very 
short ; and the dentition is 4 2, ¢ 4, p 2, m2; total 38. 

Represented by two genera, Furipterus } and Amorphochilus,? each 
including one species of peculiar aspect; the latter distinguished 
from the former by the widely separated nostrils and the great 
extension backwards of the bony palate. Habitat South America. 

In the typical or Emballonurine division part of the tail is 
included in the basal half of the interfemoral membrane, the remain- 

1 Bonaparte, Fawn. Ital. vol. i, (1832-41): Syn. Furia, F. Cuvier, Mém. du 
Muséum, vol. xvi. p. 150 (1828). Preoccupied by Linn. 1766. 
? Peters, Monatsber. Ak. Berlin, 1877, p. 185. 


EMBALLONURIDE 667 


ing part passing through and appearing upon its upper surface ; 
the crown of the head is slightly elevated; the pollex and first 
phalanx of the middle finger are moderately 
long; and the number of the premolars is 
always 2. 

Emballonura..—Incisors 3. Extremity of 
the muzzle more or less produced beyond the |’ 
lower lip, forehead flat. Contains some five ~ 
species, inhabiting islands from Madagascar 
through the Malay Archipelago to the Navi- 
gators’ Islands. Fic. 312.—EHar of Emballo- 

Coleiira.2—Incisors 4. Extremity of the eee ns) 
muzzle broad, forehead concave. Has two 
species from Kast Africa and the Seychelles Islands. 

Phynchonycteris.2A—This genus is distinguished from Coleiira by 
the much-produced extremity of the muzzle. The single species, P. 
naso, from Central and South America, is very common in the 
vicinity of streams throughout the tropical parts of these countries ; 
it is usually found during the day resting on the vertical faces of 
rocks, or on the trunks of trees growing over the water, and, owing 
to the peculiar grayish colour of the fur covering the body and 
growing in small tufts from the antebrachial membrane, so as to 
counterfeit the weathered surfaces of rocks and the bark of trees, 
easily escapes notice. As the shades of evening approach it appears 
early on the wing, flying close to the surface of the water, and 
seizing the minute insects that hover over it. 

Saccopteryx.*—Incisors 4. Antebrachial membrane with a pouch 
opening on its upper surface. This genus contains six species from 
Central and South America. In the adult males a valvular longi- 
tudinal opening is found on the upper surface of the membrane, 
varying in position in different species. This opening leads into a 
small pouch (in some species large enough to hold a pea), the 
interior of which is lined with a glandular membrane secreting an 
unctuous substance of a reddish colour with a strong ammoniacal 
odour. The presence of this sac only in males indicates that it 
is a secondary sexual character analogous to the shoulder-pouches 
of Epomophorus and the frontal sacs of Hipposiderus. It is quite 
rudimentary in the females. 

Taphozous.°—Incisors 4; upper pair deciduous. This genus, 
represented by some ten species, inhabiting the tropical and sub- 
tropical parts of all the eastern hemisphere except Polynesia, forms 
the second group of this division, distinguished by the cartilaginous 


1 Temminck (Van der Heeven), Tijdsch. Nat. Ges. 1839, p. 22. 

2 Peters, Monatsber. Ak. Berlin, 1867, p. 479. > Peters, loc, cit. p. 477. 
4 Tlliger, Prodromus Syst. Mami. et Aviwm, p. 121 (1811). 

5 Geoffroy, Deseript. de ? Egypte, vol. ii. p. 126 (1812). 


668 CHIROPTERA 


premaxillaries, deciduous upper incisors, and the presence of only 
two lower incisors. Most of the species have a peculiar glandular 
sac (Fig. 313) placed between the angles of the lower jaw. This 
is.a sexual character, for, while always more developed in males 
than in females, in some species, although distinct in the male, 
it is quite absent in the female. An open gular sac is wanting 
in both sexes in ZT. melanopogon, but about its usual position the 
openings of small pores may be seen, the secretion exuding from 


Fic. 313.—Heads of Taphozous longimanus, showing relative development of gular sacs in 
male and female. (Dobson, Proc. Zool. Soc. 1873.) 


which probably causes the hairs to grow very long, forming the 
black beard found in many male specimens of this species. 

In the Diclidwrine division there is but a single genus, repre- 
sented by two species. 

Diclidwrus..—Dentition: 1 4, ¢ 4, p 2, m 3; total 32. Both 
species are from the Neotropical region, the typical D. albus ranging 
as far north as Central America. This Bat resembles the species 
of Taphogous in the form of the head and ears, but, besides other 
characters, differs from all other Bats in possessing a peculiar pouch, 
opening on the centre of the inferior surface of the interfemoral 
membrane ; the extremity of the tail enters this, and perforates its 
fundus. 

The Noctilionine division is likewise represented only by a single 
genus, with two species. This genus connects the present with the 
following family, possessing characters common to both, but also so 
many remarkable special peculiarities as almost to warrant the 
formation of a separate family for its reception. 

Noctilio.2—Dentition: i 2, c4, pi, m3; total 28. The two 
species NV. leporinus and N. albiventer inhabit Central and South 
America. The typical N. leporinus is a Bat of very curious aspect, 
with strangely folded lips, erect cutaneous processes on the chin, 
and enormous feet and claws. The first upper incisors are close 
together, and so large as to conceal the small outer ones, while in 
the lower jaw there is one pair of small incisors. This apparent 
resemblance toa Rodent actually led Linneus to remove this species 
from the Bats and place it in the Rodents. This Bat is remark- 


1 Wied, Jsis, 1819, p. 1629. * Linn. Syst. Nat. 12th ed. vol. i. p. 88 (1766), 


EVMBALLONURIDE 669 


able for feeding on fish—a circumstance which has only recently 
been fully authenticated. 

The remaining genus of this subfamily is regarded as repre- 
senting another division, which may be known as the Lhinopomu- 
tine division. : 

Lhinepoma.|—This genus, represented by the single species 
R. micrephyllum, might also be elevated to the rank of a family, for it 
is difficult to determine its exact 
affinities, a kind of cross relationship 
attaching it to the \ycteride on the 
one hand and to this family, in which 
it is here placed provisionally, on the 
other. This species, distinguished 
from all other Microchiroptera as well 
by the presence of two phalanges in Fic. 314.—Skull of Rhinczoma micro- 
the index finger as by its remarkably Phylum. x2. (Dobson, Monogr. <isiat. 
long and slender tail projecting far sai 
beyond the narrow interfemoral membrane, inhabits the subterranean 
tombs in Egypt and deserted buildings generally from North-East 
Africa to Burma. 

Subfamily Molossine.—Tail thick, produced far beyond the 
posterior margin of the interfemoral membrane (except in J/ysta- 
cops); legs short and strong, with well-developed fibula; upper 
incisors strong. This subfamily includes all the species of Emba/- 
lonuride with short and strong legs and broad feet (whereof the 
first toe, and in most species the fifth also, is much thicker than 
the others, and furnished with long curved hairs), well-developed 
callosities at the base of the thumbs, and a single pair of large 
upper incisors occupying the centre of the space between the 
canines. In all the species the feet are free from the wing- 
membrane, which folds up perfectly under the forearm and legs; the 
interfemoral membrane is retractile, being movable backwards and 
forwards along the tail, and this power of varving its superficial 
extent must confer upon these Bats great dexterity in quickly 
changing the direction of their flight, as when obliged to double in 
pursuing their swift insect prey, which their extremely expansible 
lips evidently enable them to secure with ease. Like the preceding 
subfamily, the genera may be arranged in divisions, of which there 
are two. 

The Jfolossing division is characterised by the production of the 
tail beyond the posterior margin of the interfemoral membrane ; it 
includes three senera. 

Chiremeles.-—Dentition: i 4, ¢4, p 4, m 3; total 26. Hallux 
much larger than the other toes and separable from’ them, ears 
1 Geoffroy, Descript. de T Egupte, vol. ii. p. 123 (1212). 

2 Horstield, Zool. Ressarch Java 1324). 


670 CHIROPTERA 


separate. This genus is represented by a single species, C. torquatus, 
of large size (forearm 3:1 inches) and peculiar aspect, inhabiting 
the Indo-Malayan subregion. This Bat is nearly naked, a collar 
only of thinly spread hairs half surrounding the neck; and is 
further remarkable for its enormous throat-sac and curious nursing- 
pouches. The former consists of a great semicircular fold of skin 
forming a deep pouch round the neck beneath, and concealing the 
orifices of large subcutaneous pectoral glands, which discharge an 
oily fluid of insufferably offensive smell. The nursing-pouch is 
formed on each side by an extension of a fold of skin from the side 
of the body to the inferior surfaces of the humerus and femur. In 
the anterior part of this pouch the mamme are placed. 

Molossus.:—Dentition : 7 a ci,p , m2; total 24 or 28. 
Upper incisors close together in the middle line. There are some 
ten species, restricted to the tropical 
and subtropical regions of the New 
World. The woodcut of the head of 
M. glaucinus (Fig. 315) exhibits the 
general physiognomy of the Bats of 
this genus. IM. obscurus, a small species, 
is very common in tropical America. It 
inhabits the hollow trunks of palms and 
other trees, and also the roofs of houses. 
The males and females live apart (as, 
indeed, appears to be the case in most, 
Fic, 315.—Head of Molossusglaucinus. if not in all, RPECIES of Bats). In the 

(Dobson, Proc. Zool, Soc. 1876.) hollow trunk of a palm two colonies 

were discovered, one consisting of from 

150 to 200 individuals, exclusively males, while the other was 
composed almost entirely of females. 

Nyctinomus.2—Dentition : ts cap a, m %; total 32 or 
28. Upper incisors separated in the middle line. The genus con- 
tains about twenty-five species, inhabiting the tropical and sub- 
tropical parts of both hemispheres. The lips of the Bats of this 
genus are even more expansible than in Jfolossus, in many of the 
species (as in the woodcut of the head of NM. macrotis, Fig. 
316) showing vertical wrinkles. NV. twniotis, one of the largest 
species, alone extends into Europe, and has been taken as 
far north as Switzerland. NN. johorensis, from the Malay Penin- 
sula, is remarkable from the extraordinary form of its ears. 
NV. brasiliensis is nearly as common as Molossus obscwrus in tropical 
America, and extends farther north (California) and south than 
that species, 


1 Geoffroy, Ann. du Muséwm, vol. vi. p. 154 (1805). 
* Geoffroy, Descript. de U Egypte, vol. ii. p. 114 (1812). 


EMBALLONURIDA 671 


In the Mystacopine division the tail perforates the interfemoral 
membrane and appears upon the upper surface. 

Mystacopsi—This genus includes only M. tuberculatus of New 
Zealand, where, together with Chalinolobus tumorio, it represents 
the whole indigenous mammalian fauna of 
the islands. There are three distinct 
phalanges in the middle finger; the 
greater part of the wing-membrane is 
exceedingly thin, but a narrow portion 
along the forearm, the sides of the body, 
and the legs is remarkably thick and 
leathery ; beneath this thickened portion 
the wings are folded. With the wings 
thus encased, this species is the most sn 
quadrupedal of Bats. Other peculiarities Rie mone ee 
of structure are found in the remarkable js75) — git beak ele 
form of the claws of the thumbs and toes, 
which have each a small talon projecting from its concave surface 
near the base, also in the sole of the foot and inferior surface of 
the leg, as shown in Fig. 317. The plantar surface, including the 
toes, is covered with soft and very lax integument deeply wrinkled, 


Fic. 317.—Pollex and leg and foot of Mystacops tuberculatus, enlarged. (Dobson, 
Proc. Zool. Soc. 1876.) 


and each toe is marked by a central longitudinal groove with short 
grooves at right angles to it. The lax wrinkled integument is 
continued along the inferior flattened surface of the ankle and leg. 
These peculiarities appear to be related to climbing habits in the 
species. 

Fossil Emballonuride.—In the cavern-deposits of Madras remains 
of the existing Tuphozous saccolenus are not uncommon ; while in 
the corresponding beds of Brazil bones of a Molossus, probably refer- 
able to M. temmincki, now inhabiting the same region, are met with. 


1 New name: Syn. Mystacina ; Gray, Voyage of the ‘‘ Sulphur,” ‘‘Mamm.’ 
p. 28 (1843). Preoccupied by Mystacina, Boie, 1822. 


672 CHIROPTERA 


It has been suggested that remains from the Upper Eocene Phos- 
phorites of Central France may indicate the existence of the genus 
Taphozous at that early epoch. 


Family PHYLLOSTOMATIDA. 


Middle finger with three well-developed bony phalanges ; first 
phalanx of the middle finger short ; nostrils in the front part of the 
cutaneous nasal appendages, or opening by simple apertures at 
the extremity of the muzzle; chin with warts or erect cutaneous 
ridges ; premaxille well developed, united in front. 

The members of this family are readily distinguished by the 
third phalanx in the middle finger, associated either with distinct 
cutaneous nasal appendages, or with well-developed first upper 
incisors, or with both. Unlike the Rhinolophide, their eyes are 
generally large; and the tragus is well developed, maintaining 
almost the same form throughout the species, however much the 
other parts of the body may vary. The fur is of a dull colour, and 
the face and back“(in the Stenodermatine division especially) are often 
marked with white streaks, as in the Pteropodide, of which these 
Bats take the place in the western hemisphere. A few species, 
probably all those with the tail and interfemoral membrane well 
developed, feed principally on insects, while the greater number of 
the species of the ’ampirine and Glossophagine divisions appear to 
live on a mixed diet of insects and fruits; and the Desmodontine 
division, of which two species only are known, are true blood- 
suckers, and have their teeth and intestinal tract specially modified 
in accordance with their habits. The family is restricted to the 
tropical and subtropical parts of Central and South America. 

Subfamily Chilonyeteriinze.—Nostrils opening by simple aper- 
tures at the extremity of 
the muzzle in front, not 
margined by a distinct nose- 
leaf; chin with expanded 
leaf-like appendages. It 
includes two genera. 

Chilonycteris.1.—Dentition: 
i$, c4, p 2, m3; total 34. 
The crown of the head is 
moderately elevated above 


Via. 318.—Head of Mormops blainvillet. (Dobson % . i 
Cat. Chiropt. Brit. Mus.) > the facial line, and the basi- 


cranial axis is almost in the 
same plane as the facial. There are about half a dozen species. 
Mormops.2—The two species of this genus are distinguished 
1 Gray, Ann. Mag. Nat. Hist. vol. iv. p. 4 (1839). 
* Leach, Zrans. Linn. Soe, vol. xiii. p. 76 (1820-22).—Amended. 


PHVLLOSTOMA TIDE 673 


from Chilonycteris by the great elevation of the crown of the head 
above the line of the face, as well as by the basicranial plane being 
nearly at right angles to the facial. Both species are noticeable 
for their peculiar physiognomy, as is shown in the accompanying 
woodcut (Fig. 318). 

Subfamily Phyllostomatins.—Nostrils opening on the upper 
surface of the muzzle, the nasal apertures more or less surrounded 
or margined by well-developed cutaneous appendages, forming a 
distinct nose-leaf; chin with warts. The numerous genera, most 
of which can only be mentioned here by name, may be arranged 
under four divisions. 

In the first or Vampirine division the muzzle is long and narrow 
in front ; the distance between the eyes is generally less than, rarely 
equal to, that from the eye to the extremity of the muzzle; the 
nose-leaf is well developed, horse-shoe shaped in front, and lanceolate 
behind ; interfemoral membrane well developed; tail generally 
distinct, rarely absent ; inner margin of the lips not fringed. The 
dentition is: 7 =n c+, p sys m%; total 32. The cusps of the 
upper molars are usually well developed, and arranged in a W. 
Nearly all the species of this division appear to be insectivorous, so 
that the name applied to them must not be considered as having 
any relation to their habits. Vampyrus spectrum, a large Bat 
inhabiting Brazil, of forbidding aspect, which was long considered 
by naturalists to be sanguivorous in its habits, and named accord- 
ingly by Geoffroy, has been shown by the observations of modern 
travellers to be mainly frugivorous, and is considered by the 
inhabitants of the countries in which it is found to be perfectly 
harmless. It is the largest Bat in America, the length of the 
forearm being 4°2 inches. Otopterus waterhousei appears to prey 
occasionally on small species of Bats, like Megaderma lyra of the 
eastern hemisphere, which it resembles in many respects. 

Lonchorhina,’ Otopterus,? and Dolichophyllwm.?—These three genera 
are characterised by the tail continuing to the hinder margin of the 
interfemoral membrane. Lonchorhina is represented by the single 
species L. auwrifa, in which the nose-leaf is much elongated, and the 
ear-conch and tragus are unusually large. 

Vampyrus,* etc-—In all the remaining genera of this division the 
tail perforates the interfemoral membrane, so as to appear upon its’ 
upper surface. These genera are Vampyrus, Lophostoma, Micronycteris,° 


1 Tomes, Proc. Zool. Soc. 1863, p. 81. 2? New name: Syn. Macrotus ; 
Gray, Proc. Zool. Soc. 1843, p. 21. Preoccupied by Aacrotis, Dej. 1833. 
® New name: Syn. Macrophyllum ; Gray, Mag. Zool. Bot. vol. ii. p. 489 (1838). 
Preoccupied by Macrophylla, Hope, 1837. 4 Leach, Trans. Linn. Soc. vol. 
xiii. pp. 74, 75 (1822). For the references to the other genera see Dobson, Cat. 
Chiropt. Brit. Mus. ° Gray, Proc. Zool. Soc. 1866, p. 113. Syn. Schizostoma ; 
Gervais, 1855. Preoccupied by Broun, 1835. 

43 


674 CHIROPTERA 


Trachyops, Phylloderma, Phyllostoma, Anthorhina,) M mon, Hemiderma 3 
and Rhinophylla, all, with the exception of the last, being distinguished 
from one another chiefly by the form of the skull and the presence 
or absence of the second lower premolar. 
Trachyops, Phylloderma, and the three 
last-named genera are each represented 
by a single species. Phyllostoma has- 
tatum, in which the forearm has a 
length of 3°2 inches, and next in point 
of size to J“ampyrus spectrum, is a well- 
known species in South America; P. 
elongatum (Fig. 319) differs in its smaller 
size and much larger nose-leaf. Hemz- 
. derma brevicauda is a small species, 
Fig, 819.—Head of Phyllostoma elon- which forms a connecting link between 
gatum. (From Dobson, Proc. Zool. Soc. és eS ese 5% 
1866.) this and the next division. Ahinophyllu 
pumilio, the smallest known species 
of the family, is further distinguished by the narrowness of its 
molars, which do not form W-shaped cusps, and by the very small 
size of the last upper molar; characters connecting it with the 
Stenodermatine division. 

In the second or Glossophagine division of the subfamily the 
muzzle is long and narrow; the tongue remarkably long and exten- 
sible, much attenuated towards the tip, and beset with very long 
filiform recurved papille ; lower lip with a wide groove above, and 
in front margined by small warts; nose-leaf small; tail short or 
absent. Dentition: i 4, ¢ 3, p =i m <3 ; teeth very narrow ; 
molars with narrow W-shaped cusps, sometimes indistinct or absent ; 
lower incisors very small or deciduous. 

The ten species included in this division are arranged under 
seven genera,’ distinguished principally by differences in the form 
and number of the teeth and the presence’ or absence of the 
zygomatic arch. The form and position of the upper incisors are 
extremely variable. In Glossophaga and Phyllonycteris the upper 
incisors form, as in the Vampyrine division, a continuous row between 
the canines; in JMonophylla and Leptonycteris* they are separated 
into pairs by a narrow interval in front; while in Lonchoglossi, 
Glossonycteris, and Cheronycteris they ave widely separated and placed 
in pairs near the canines. In the first four genera the lower incisors 
are present (at least up to a certain age), while in the last three 


' New name: Syn. Zylostoma ; Gervais, 1855. Preoceupied by Sharpe, 1849. 
2 Gervais, Castlenau’s Exped,-Zool, p. 48 (1855): Syn. Carollia, Gray, 1888. 
Preoceupied by Carolia, Cantraine, 1837, 3 The references to the genera of 
this and the following division will be found in Dobson’s Cataloyue. + New 
name: Syn. Jschnoglossa, Saussure, 1860. Preoccupied by Kraatz, 1856, 


PHYLLOSTOMATIDA 675 


they are deciduous even in youth. The zygomatic arch is wanting 
in Phyllonyeteris, Glossonycteris, and Cheronycteris. 

The typical species is Glossophaga soricina, which so closely 
resembles Hemiderma brevicauda, both in external form and dentition, 
that it has frequently been confounded with it. Its long fimbriated 
tongue, which it possesses in common with other species of the 
division, led Spix to 
describe it as a blood- 
sucker, believing that 
this organ was used to 
increase the flow of 
blood. This view is, 
however, without found- 
ation, and from later 
observations it is evident 
that the peculiarly Fic. 320.—Head of Cheronycteris menica net, showing 

: fimbriated tongue. (Dobson, Cat. Chiropt. Brit. Mus.) 

shaped tongue is used 

by the animal to lick out the pulpy contents of fruits having hard 
rinds. The food of the species of this division appears to consist 
of both fruit and insects, and the long tongue may also be used for 
extracting the latter from the deep corolle of certain flowers. This 
type of tongue is shown in the woodcut of the head of Cheronycteris 
(Fig. 320); and it is paralleled among the Megachiroptera by the 
Carponycteriine Pteropodida. 

The Stenodermatine division is characterised by the muzzle being 
very short and generally broad in front, the distance between the 
eyes nearly always exceeding (rarely equal to) that from the eye to 
the extremity of the muzzle; nose-leaf short, horse-shoe shaped in 
front, lanceolate behind (except in Brachyphylla and Centurio) ; 
interfemoral membrane always concave behind; tail none; inner 
mar, on . the lips fringed with conical papille. Dentition : 


tr =v. pF ca 2 ; the number of the molars being either 3, 3, 


or 3 in ees elds premolars and molars very broad (except 
in Stur nira), the latter with concave or flat crowns margined exter- 
nally by raised cutting-edges. Although the members of this division 
are usually distinguished from those of the Vampirine division by 
the peculiar shortness and breadth of the muzzle and the form of 
the molars, yet certain species of the latter closely resemble those 
of the former in external appearance, agreeing almost absolutely in 
the form of the nose-leaf, of the ears and tragus, and of the warts 
on the chin. These resemblances indicate that, while the form of 
the teeth and jaws has become modified to suit the nature of the 
food, the external characters, being but slightly: affected by this 
cause, have remained much the same. The food of these Bats 
appears to be wholly or in great part fruit. The twenty species 
have been grouped into nine genera, distinguished by the form of 


676 CHIROPTERA 


the skull and teeth. Artibeus, with six species, includes the well- 
known frugivorous Bat, 4. perspicillatus. Waterton believed that 
A. planivostris, a common Bat in British Guiana, usually found in 
the roofs of houses, and now known to be frugivorous, was the true 
blood-sucking Vampire. Stenoderma achradophilum, found in Jamaica 
and Cuba, associated with Artibeus perspicillatus, from which it is 
scarcely distinguishable externally except by its much smaller size, 
differs altogether in the absence of the horizontal plate of the 
palatal bones. Sturnira lilium, while 
agreeing with the above in the form of 
the nose-leaf and ears, differs from all 
the species of the family in its longi- 
tudinally-grooved molars, which resemble 
those of the Pteropodide more closely than 
those of any other Bats; and the presence 
of tufts of long differently coloured hairs 
comets Eee over glands in the sides of the neck shows 
1G. 321.—Head of Centurio senex. : 
(Dobson, Cat, Chiropt. Brit. Mus.) another common character still more 
remarkable, which can scarcely be con- 
sidered the result of adaptive change. Centurio senex is the type 
of a genus distinguished from Stenoderma and other genera of this 
division by the absence of a distinct nose-leaf ; its facial aspect, as 
shown in Fig. 321, is altogether bizarre. 

In the last or Desmodont division the muzzle is conical and 
short; there is a distinct nose-leaf; the interfemoral membrane is 
very short; and the tail is wanting. Dentition: 7 3, ¢ 41, p 3, 
m 3 total 24 or 20. Upper incisors very large, trenchant, 
occupying the whole space between the canines; premolars very 
narrow, with sharp-edged longitudinal crowns; molars rudimentary 
or wanting; stomach greatly elongated, intestiniform. There are only 
two genera, the single species of each of which are the true blood- 
sucking Vampires. They appear to be confined chiefly to the 
forest-clad parts, and their attacks on men and other warm-blooded 
animals were noticed by some of the earliest writers. Thus Peter 
Martyr (Anghiera), who wrote soon after the conquest of South 
America, says that in the Isthmus of Darien there were Bats which 
sucked the blood of men and cattle when asleep to such a degree 
as to kill them. Condamine, a writer of the eighteenth century, 
remarks that at Borja (Ecuador) and in other places they had 
entirely destroyed the cattle introduced by the missionaries. Sir 
Robert Schomburgk relates that at Wicki, on the river Berbice, no 
fowls could be kept on account of the ravages of these creatures, 
which attacked their combs, causing them to appear white from loss 
of blood. Although these Bats were known thus early to Europeans, 
the species to which they belonged were not determined until about 
sixty years ago, several of the large frugivorous species having been 


PHYLLOSTOMATIDA 677 


wrongly set down as blood-suckers and named accordingly; and it 
fell to the lot of Darwin to determine at least one of the blood- 
sucking species, the following being his account of the circumstances 
under which the discovery of the sanguivorous habits of Desmodus 
rufus was made: “The Vampire Bat is often the cause of much 
trouble by biting the horses on their withers. The injury is gener- 
ally not so much owing to the loss of blood as to the inflammation 
which the pressure of the saddle afterwards produces. The whole 
circumstance has lately been doubted in England; I was therefore 
fortunate in being present when one was actually caught on a horse’s 
back. We were bivouacking late one evening near Coquimbo, in 
Chili, when my servant, noticing that one of the horses was very 
restive, went to see what was the matter, and, fancying he could 
detect something, suddenly put his hand on the beast’s ‘withers 
and secured the Vampire.” 

These Bats present, in the extraordinary differentiation of the 
manducatory and digestive apparatus, a departure from the type of 
other members of the family unparalleled in any of the other orders 
of Mammalia, standing apart from all other mammals as being fitted 
only for a diet of blood, and capable of sustaining life upon that 
alone. Travellers describe the wounds inflicted by the large sharp- 
edged incisors as similar to those caused by a razor when shaving: 
a portion of the skin being shaved off and a large number of 
severed capillary vessels thus exposed, from which a constant flow 
of blood is maintained. From this source the blood is drawn 
through the exceedingly narrow gullet—too narrow for anything 
solid to pass—into the intestine-like stomach, whence it is probably 
gradually drawn off during the slow process of digestion, while the 
animal, sated with food, is hanging in a state of torpidity from the 
roof of a cave or the inner side of a hollow tree. 

Desmodus.1—No true molar, and no calcar. The Common 
Vampire (D. rufus) is widely spread over the tropical and sub- 
tropical parts of Central and South 
America from Oaxaca to Southern Brazil 
and Chili. It is a comparatively small 
species, a little larger than the common 
Noctule, the head and body being about 
3 inches in length, the forearm 24, with 
a remarkably long and strong thumb; 
it is destitute of a tail, and has a 
peculiar physiognomy, well represented — Fie. 322.—Head of Vampire Bat 
in Fig. 322. The body is covered with sc imiabtsiats 
rather short fur of a reddish-brown colour, but varying in shade ; 
the extremities of the hairs being sometimes ashy. The teeth 
are peculiar and admirably adapted for the purposes for which they 

1 Wied, Beitr. Natgesch. Brasil, vol. ii. p. 231 (1826). 


678 CHIROPTERA 


are employed. The upper incisor is greatly enlarged, and of some- 
what triangular shape (Fig. 323); the canine, although smaller 
than the incisor, is large and sharp; but the cheek-teeth are very 
small, with laterally compressed crowns rising but slightly above 
the level of the gum, their longitudinally disposed. cutting-edges 
being continuous with the base of the canine and with each other. 
The lower incisors are small, bifid, and separated from the canine, 
with a space in front. The 
lower cheek-teeth are nar- 
row, like those in the upper 
jaw, but the anterior tooth 
is slightly larger than the 
others, and separated by a 
small space from the canine. 
Behind the lower incisors 
the jaw is deeply hollowed 
out to receive the ex- 
tremities of the large upper 
incisors. The exceedingly 
narrow cesophagus opens at 
right angles into the slender, intestinelike stomach, which almost 
immediately terminates on the right, without a distinct pylorus, 
in the duodenum, but on the left forms a greatly elongated fundus, 
bent and folded upon itself, appearing at first sight like part of the 
intestines. This cardiac extremity of the stomach is, for a short 
distance to the left of the entrance of the cesophagus, still very 
narrow, but soon increases in size, till near its termination it 
attains a diameter quite three times that of the short pyloric 
portion. The length of this cardiac diverticulum of the stomach 
appears to vary from 2 to 6 inches, the size in each specimen 
probably depending on the amount of food obtained by the animal 
before it was captured. 

Diphylia.imA small true molar in each jaw, and a rudimentary 
calcar. The single species D. ecaudata inhabits Brazil, and appears 
to be much less abundant than Desmodus rufus, from which, in 
addition to the characters already mentioned, it is distinguished by 
its slightly smaller size, the absence of a groove in the front of the 
lower lip, the non-development of the interfemoral membrane in the 
centre, and the peculiar form of the lower incisors, which are much 
expanded in the direction of the jaws and pectinated, forming a 
semicircular row touching each other, the outer pair being wider 
than the inner ones, and having six notches, the inner pair having 
only three notches. 

Fossil Phyllostomatide.—Remains of Tampyrus spectrum, as well 
as of several species of Phyllostoma or closely allied types, are found 

1 Spix, Sim. e¢ Vesp. Brasil, p. 68 (1823). 


view of upper teeth; b, left lateral view of upper and 
lower teeth. 


PHYVLLOSTOMATIDA 679 


in the cavern-deposits of Brazil. The mandible of a large Bat from 
the Upper Eocene Phosphorites of Central France, described as 
Necromantis, has been referred to this family—a determination 
which, if confirmed, will be of great interest from a distributional 
point of view. 


Bibliography of Chiroptera.—G. E. Dobson, Catalogue of the Chiroptera in the 
Collection of the British Museum, 1878, including descriptions of all the species 
of Bats then known ; subsequent papers by the same author in Rep. Brit. Assoc., 
Proc. Zool. Soc., Ann. Mag. Nat. Hist., and Bull. Soc. Zool. de France ; by 
Peters in Monatsb. Akad. Wiss. Berlin; by O. Thomas in Ann. Mag. Nat. Hist., 
Proc. Zool. Soc., and Ann. Mus. Genova ; and by J. Scully in Ann. Mag. Nat. Hist. 
and Journ. As. Soc. Bengal; H. A. Robin, Recherches Anatomiques sur les Mam- 
- miferes de V Ordre des Chiroptéres, Paris, 1881; W. T. Blanford, “Notes on Indian 
Chiroptera,” Journ. As. Soc. Bengal, vol. lviii. (1888). See also papers by Jentink, 
Bocage, and others. 


CHAPTER XIV 
THE ORDER PRIMATES 


TuIs order in the system of Linnzus includes Man, the Monkeys, 
the Lemurs, and the Bats. By common consent of all zoologists 
the last-named animals have been removed into a distinct order ; 
but with regard to the association of the others there has been, 
and still is, much difference of opinion. 

That all the Monkeys, from the highest Anthropoid Apes to 
the lowest Marmosets, form a natural and tolerably homogeneous 
group seems never to have been questioned; but whether the 
Lemurs on the one hand and Man on the other should be united 
with them in the same order are points of controversy. If, in 
accordance with the traditional views of zoologists, the former are 
still considered to be members of this order, they must form a sub- 
order apart from all the others, with which they have really very 
little in common except the opposable hallux of the hind foot, a 
character also met with in the Opossums, and which is therefore of 
very secondary importance.! 

Using the term Primates in this wider sense it is not easy to 
give any precise definition of the order. The dentition is diphy- 
odont and heterodont ; the number of incisors being very generally 
3, and that of the molars, with the exception of the Hapalide, 
being 3. The cheek-teeth are adapted for grinding, the molars 
being more complex than the premolars, and usually having four 
main tubercles, which may be either subconical or more or less 
compressed. The orbit is invariably surrounded by a ring of bone; 


1 For the arguments in favour of placing the Lemurs in a separate order 
see Milne-Edwards, ‘‘Observations sur quelques points de l’embryologie des 
Lemuriens et sur les affinités zoologiques de ces animaux,” in the Azz. des 
Sciences Nat. October 1871; and P. Gervais, ‘‘Encephale des Lemures,” in 
Journ, de Zoologie, tom. i. p. 7. For those for retaining them among the 
Primates, see Mivart, ‘‘On Lepilemur and Chirogaleus, and on the Zoological 
Rank of the Lemuroidea,” in Proc. Zool. Soc, 1878, p. 484. 


PRIMATES 681 


the clavicles are well developed ; and the radius and ulna are never 
united. The scaphoid and lunar of the carpus, and commonly also 
the centrale, remain distinct from one another. There are usually 
five digits furnished with well-developed nails in both the manus 
and the pes; but the pollex may be rudimentary or wanting. The 
hallux, except in Man, is opposable to the other digits, and has a 
flat nail (absent in Simia) ; and the pollex, 
when present, is usually also more or less 
opposable. The terminal phalanges of 
the digits are flattened (except in the 
second digit of the pes of the Lemu- 
roidea), and not cleft at their extremities. 
The fingers and toes generally do not 
taper towards their extremities, but (ex- 
cept in Chiromys) are dilated, flattened, 
and rounded at their tips. The humerus 
has no entepicondylar foramen, nor the 
femur a third trochanter. In the ali- 
mentary canal (Fig. 324) the stomach is 
generally simple, although sacculated in 
the subfamily Semnopithecine of the 
Cercopithecide ; and there is always a 
cecum, which is generally of large size. 
The placenta may be either non-deciduous, 
or discoidal and deciduous. There are 
always two mammez in the pectoral 
region, except in Chiromys; and the gg 394 alimentary canal of 
testes descend into a scrotum. Galago, the greater part of the small 
The Lemuroidea are decidedly low in itestine being omitted. d, duo- 
: a a denum; 7, ileum; em, cecum; 7, 
the scale of organisation, their placenta- yoctum, 
tion being of a lower type than that 
of the Insectivora; and all the Primates retain generalised features 
in their pentadactylate limbs and more or less bunodont cheek-teeth. 
In respect to cerebral characters and other features the higher 
representatives of the order have, however, acquired a specialisation 
clearly indicating their right to occupy the highest position in the 
animal kingdom. So far as the available material admits of forming 
an opinion, fossil forms appear to indicate an intimate connection 
between the Lemuroidea and Insectivora, so that in some cases it is 
almost impossible to determine whether an extinct type should be 
referred to the former or to the latter group. It is noteworthy 
that while in all existing Primates the upper molars are of a quadri- 
tuberculate type, in the extinct Lemuroid genus Anaptomorphus 
they are trituberculate. 


682 PRIMATES 


Suborder LEMUROIDEA. 


The Latin term Lemur was applied by Linnzus to the typical 
representatives of the present group of Primates, having been sug- 
gested by the nocturnal habits and strange ghost-like appearance 
of some of its members. As these animals had previously no 
vernacular appellation in English, this name has been generally 
adopted, and is now completely anglicised, making “Lemurs” in 
the plural. The French call them Makis, and the Germans Halbaffen, 
in allusion to their forming a transition from monkeys to ordinary 
quadrupeds. For the same reason they are called Prosimie by 
some systematic writers. When the name was bestowed by 
Linneus only five species were known, of which one, L. volans, 
Linn. (Galeopithecus volans of modern writers), is now removed by 
common consent from the group. Notwithstanding the discovery 
of many new and curious forms, the Lemurs remain a very natural 
and circumscribed division of the animal kingdom, though no longer 
considered a single genus, but divided up into many genera and 
even families. 

The existing species are not numerous, and do not diverge 
widely in their organisation or habits, being all of small or moderate 
size, all adapted to an arboreal life, climbing with ease, and, as they 
find their living, which consists of fruits, leaves, birds’ eggs, small 
birds, reptiles, and insects, among the branches of the trees, they 
rarely have occasion to descend to the ground. None are aquatic, 
and none burrow in the earth. Many of the species, although by no 
means all, are nocturnal in their habits, spending the day in sleep- 
ing in holes, or rolled up in a ball, perched on a horizontal branch, 
or in the fork of a tree, and seeking their food by night. Their 
geographical distribution is very peculiar; by far the larger pro- 
portion of species, including all those to which the term “ Lemur ” 
is now especially restricted, being exclusively inhabitants of Mada- 
gascar, where they are so abundant and widely distributed that it 
is said by M. Grandidier, who has contributed more than any other 
traveller to enrich our knowledge of the structure and manners of 
these animals, that there is not a little wood in the whole island 
in which some of them cannot be found. From Madagascar as a 
centre a few species less typical in character extend through the 
African continent westward as far as Senegambia, and others are 
found in the Oriental region as far east as the Philippine Islands 
and Celebes. 

The following are the essential characters by which the sub- 
order as a whole is distinguished from the Anthropoidea. Skull 
with the orbit opening freely into the temporal fossa beneath the 
postorbital bar (except in Tarsius); and the lachrymal foramen 


LEMURIDA 683 


situated externally to the margin of the orbit (Fig. 327). The 
pollex and hallux are always well developed, the latter being 
especially large ; the second or index digit of the manus may be 
rudimentary ; while in the pes the second digit invariably termin- 
ates in a long pointed claw. The cerebral hemispheres do not 
completely overlap the cerebellum, and are but slightly convoluted. 
The uterus is bicornuate. The placenta is non-deciduate, and either 
diffused or bell shaped—the whole of the chorion except the 
cephalic pole being covered with villi; and the allantois is of very 
great size. There may be abdominal mamme. Except in Chiromys, 
the first pair of upper incisors are separated in the middle line. 
In marked contrast to the Anthropoidea, the middle or transverse 
portion of the colon is almost always folded or convoluted on 
itself. (See Fig. 324.) 

In subdividing the group for the purpose of a more detailed 
description of the different animals of which it is composed it must 
first be noted that there are two very aberrant forms, each repre- 
sented by a single species—the little Tarsius of the Indian archi- 
pelago, and the singular Chiromys or Aye-aye, which, though an 
inhabitant of Madagascar, the headquarters of the suborder, and living 
in the same forests and under the same external conditions as the 
most typical Lemurs, exhibits a most remarkable specialisation in 
the structure of its limbs and teeth, the latter being modified so as 
to resemble, at least superficially, those of the Rodents, in which 
order it was once placed. The differences between these two forms 
and the remaining Lemurs are so great that the whole suborder 
naturally divides itself into three families, the first of which may 
be again divided into four subfamilies. 


Family LEMURIDE. 


Upper incisors two on each side, small and separated by an 
interval in the middle line. Upper canine large, conical, com- 
pressed, and pointed. Premolars two or three, molars three on 
each side above and below, with numerous more or less pointed 
cusps. In the front of the lower jaw are on each side two or three 
closely approximated, long, slender teeth lying almost horizontally 
and projecting forwards. These are generally considered to repre- 
sent the incisors and canine, but there is some doubt about their 
homologies, and they may be all considered as incisors, the canine 
being absent. The first lower premolar larger than those behind 
it, and shaped like a canine, of which it performs the function 
(Fig. 327). The orbit and temporal fossa widely continuous beneath 
the bar of bone (formed by the frontal and jugal) constituting the 
posterior boundary of the former cavity. The fibula well developed 
and distinct from the tibia. All the digits of both feet (except the 


684 PRIMATES 


second of the hind foot) with flat nails, and corresponding form of 
ungual phalanges. 

Subfamily Indrisinee.—The dentition of the adult consists of 
thirty teeth, usually expressed by the formula 7 3, ¢4, p 2, m3; 
but, as indicated above, they may be i 2,¢ 4,» 3, m3. In the 
milk-dentition there are twenty-two teeth, the true molars of course 
not being represented, but there are two additional teeth in the 
fore part of the lower jaw which have no successors in the permanent 
series. Hind limbs greatly developed, but the tarsus normal. 
Hallux of large size, and very opposable. The other toes united 
at their base by a fold of skin, which extends as far as the end of 
the first phalanx. Mamme two, pectoral. Czecum very large, and 
colon extremely long and spirally coiled. 

The animals of this group are, as their organisation indicates, 
essentially arboreal, and feed exclusively on fruit, leaves, buds, and 
flowers. They are restricted geographically to the island of 
Madagascar. Among them are the largest members of the sub- 
order. A detailed and beautifully illustrated account of their 
characters, external and internal, and distribution and_ habits, 
is given in the Histoire Naturelle de Madagascar, by A. Grandidier 
and Alphonse Milne-Edwards (1875). The species are not numerous 
and are distributed into three genera. 

Indvis.\— Upper incisors subequal in size. Upper canine larger 
than the first premolar. Muzzle moderately long. Ears exserted. 
Carpus without an os centrale. Tail rudimentary. Vertebre : 
C7,D12,L9,8 4,09. 

_ The only well-established species is the Indris (J. brevicaudata, 
Fig. 325), discovered by Sonnerat in 1780. It is the largest of 
the Lemurs, the length of the head and body being about 2 feet, 
and the tail 2 inches, It is very variable in colour, for although 
usually nearly black, marked with whitish spots principally in the 
lumbar region and forearm, individuals have been found quite 
white. It inhabits exclusively the forests of a part of the east 
coast of Madagascar, living in small troops of four or five in number, 
and resembling in most of its habits the animals of the next genus. 

Propithecus.-—Second upper incisor much smaller than the first. 
Upper canine larger than the first premolar. Muzzle rather short. 
Ears short, concealed by the fur. An os centrale in the carpus. 
Tail long. Vertebre: C 7, D 12, L 8, S 3, © 28. 

The species are all subject to great variations in colour, which 
has led to much difficulty in discriminating them, and to much 
confusion of synonymy. Grandidier and Milne-Edwards recognise 
three as certainly distinct —P. diadema, P. verreauaii, and P. | 
coronatus (Fig. 326). Some of these are to be found in almost 

1 Geoffroy, Mag. Encyclop. 2d ann. vol. i. p. 46 (1796), ‘Indri.” 
* Bennett, Prov. Zool. Soc. 1832, p. 20. 


LEMURID.E 685 


every part of the island of Madagascar, living in the woods in small 
bands of six or eight together, and feeding exclusively on buds, 
flowers, and berries. Their powerful hind limbs enable them to 
leap from tree to tree, often to a distance of 10 yards, without any 
apparent effort, and thus seeming to fly through the air. When 
obliged to descend to the ground to pass from one clump of trees 


Fic. 325.—Indris (Indris brevicaudata). From Milne-Edwards and Grandidier, 
Mammiferes de Madagascar, pl. 12. 


to another they do not run on all fours, but stand erect, and 
throwing their arms above their heads progress by a series of short 
jumps, producing an effect which is described by travellers who 
have seen them thus in their native haunts as exceedingly ludicrous. 
They are not nocturnal, but most active in the morning and even- 
ing, remaining seated or coiled up among the branches during the 
heat of the day. They are naturally of a quiet and gentle disposi- 
tion, and do not show much intelligence. All the species are also 
less vociferous than the true Lemurs, only when alarmed or angered 


686 PRIMATES 


making a noise which has been compared to the clucking of a fowl. 

Like the rest of the subfamily they never have more than a single 

young one at a time. - 
Avahis.1—Second upper incisor larger than the first. Upper 


Fic, 326.—Propithecus coronatus. (From Milne-Edwards and Grandidier, Mammiféres de 
Madagascar, pl. 7.) 


canine scarcely larger than the first premolar. Muzzle very short. 
Ears very small and hidden in the fur, which is very soft and 
woolly. Carpus without an os centrale. Taillong. Vertebre: C 7, 
D 11,1 9, $ 3, € 23. 

One species, -f. laniger, the Woolly Lemur, or Avahis, consider- 
ably smaller than any of the last genus. It differs from them in 


? Jourdan, Mem. de ’Institut, vol. ii. p. 231 (1834). 


LEMURID 687 


its habits, being quite nocturnal, and not associating in small troops, 
but being always met with either alone or in pairs. It is very 
slow in its movements, and rarely descends to the ground, but 
when it does it walks upright like the other Indrisinw. It is found 
throughout the forests which clothe the mountains on the east coast 
of Madagascar, and also in a limited district on the north-west 
coast, the specimens from the latter locality being of smaller size 
and rather different in colour. 

Subfamily Lemurinse.—The dentition in the adult consists of 
thirty-six teeth, which, as usually enumerated, are i 2, ¢ 4, p 4, m 3. 
In the fore part of the lower jaw are on each side three elongated, 
compressed, procumbent teeth, of which the outer, usually con- 
sidered the homologue of the canine, is larger than the others. All 
the forms have long tails. Hind limbs not of the same dispropor- 
tionate size as in the last group; and the cecum much less devel- 
oped. ‘Tarsus but slightly elongated, the caleaneum being always 
less than one-fourth the length of the tibia. Toes of the hind feet 
free to the base. Habitat, Madagascar, and some of the adjacent 
Comoro Islands. 

This group contains the typical Lemurs, or rather those to 
which the term is now chiefly restricted. Two somewhat aberrant 
members make it necessary to divide it into three genera. 

Lemur..—Upper incisors separated by an interval in the middle, 
and not in contact with each other or the canine, in front of which 
they are both placed. Muzzle elongated. Ears conspicuous and 
tufted. Mamme two, pectoral. Vertebre: C 7, D 12, L 7 (or D 
13, L 6), 8 3, C 27. 

Animals much about the size of a common Cat, with Fox-like 
faces, soft thick fur, and long tails well clothed with hair. Not 
having the same 
disproportionate 
size of the limbs 
as the last group, 
they are much 
more quadru- 
pedal in their 
actions, walking 
on the ground 
or running along 
the branches of 


bees on all Tous Fic. 327,—Skull of Ring-tailed L (L ta). + 
1G, 327,—Skull of Ring-tailed Lemur (Lemur cattu). x {. ue, 
feet, but also Upper canine ; lc, lower canine; pm, premolars; m, 1nolars. 


jumping with 
marvellous agility. They are gregarious, living in small troops, 
are diurnal in their habits, hut most active towards evening, when 


1 Linn. Syst. Nat. 12th ed. vol. i. p. 44 (1766). 


688 PRIMATES 


they make the woods resound with their loud cries. They feed 
not only on fruits and buds, but also on eggs, young birds, 
and insects. When at rest or sleeping they generally coil 
their long, bushy tails around their bodies, apparently for the 
sake of the warmth it affords. They have either one or two 
young ones at a birth, which are at first nearly naked, and are 


BY! 
reason Ae: 


Fic. 328.—The Ring-tailed Lemur (Lemur catta). 


carried about, hanging close to and almost concealed by the hair of 
the mother’s belly. After a while they change their position and 
mount upon the mother’s back, where they are carried about until 
they are able to climb and leap by themselves. Though no member 
of the Indrisine has as yet lived long enough in captivity to be 
brought alive to Europe, various species of Lemurine are commonly 
seen in menageries, and often breed in England. They present a 
great tendency to variation in their colouring, in consequence of 
which many nominal species have been made. The most distinct, and 
at the same time most beautiful, is the Ring-tailed Lemur (L. catta, 


LEMURIDE 689 


Fig. 328), of a delicate gray colour, and with a long tail marked 
with alternating rings of black and white. This is said by Mr. G. 
A. Shaw! to be an exception to all the other Lemurs in not being 
arboreal, but living chiefly among rocks and bushes. Pollen, how- 
ever, says that it inhabits the forests of the south-west parts of 
Madagascar, living, like its congeners, in considerable troops, and 
not differing from them in its habits. He adds that it is extremely 
gentle, and active and graceful in its movements, and utters at 
intervals a little plaintive cry like that of a domestic cat. All the 
others have the tail of uniform colour. The largest species is L. 
varius, the Ruffed Lemur, sometimes black and white, and some- 
times reddish-brown, the variation apparently not depending on 
sex or age, but on the individual. In ZL. macwco the male is black 
and the female red. LZ. mongoz, L. collarvis, and L. albifrons are 
other well-known species. 

Hapalemur.2—Upper incisors very small, subequal, separated 
widely in the middle line. Those of either side in contact with each 
other and with the canine, the posterior one being placed on the 
inside, and not in front of the latter. Muzzle very short and 
truncated. Mammez four. There is apparently but one species, 
H, griseus, smaller than any of the true Lemurs, of a dark gray 
colour, with round face and short ears. It is quite nocturnal, and 
lives chiefly among bamboos, subsisting on the young shoots. A 
second species has been named H. sinus, but it is doubtful if it is 
more than a variety. 

Lepidolemur.2—Upper incisors absent or rudimentary. Muzzle 
more elongated than in the last. No distinct os centrale in the 
carpus. JL. mustelinus is the best-known species. It has, at all 
events when adult, no upper incisors. It is rare, and like 
Hapalemur nocturnal in its habits. A second closely allied species, 
but with better developed premaxille, containing a pair of small 
styliform incisors, has been described by Peters* under the name 
of Myxoeebus caniceps. 

Subfamily Galagingze.—Dentition as in Lemuwrine, from which 
the members of this subfamily are distinguished by the elongation 
of the tarsus, caused by a peculiar modification of the caleaneum 
and the navicular, the distal portion of the former and the whole 
of the latter having the form of almost cylindrical rods placed side 
by side, while the other bones retain nearly their normal form and 
proportion. 

Chirogaleus.o—Last wpper premolar very much smaller than the 
first molar, with only one external cusp. The animals included 


1 Proc, Zool. Soc. 1879, p. 182. 2 Gray, Proc. Zool, Soc. 1870, p. 829. 
3 T. Geoffroy, Cat. Mus. Hist. Nat. Paris, p. 75 (1851). Amended from 
Lepilemur. + Monatsb. Ak. Berlin, 1874, p. 690. 


5 Geoffroy, dan. du Muséwin, vol. xix. p. 171 (1812). 
y> 
44 


690 PRIMATES 


under this name appear to form a transition between the true 
Lemurs and the Galagos. The genus was originally established by 
Geoffroy St. Hilaire in 15312 for the reception of three species 
only known at that time by drawings made in Madagascar by the 
traveller Commerson. Subsequent discoveries have brought to 
light several others that may be referred to it, including one or 
two which are sometimes considered as forming a genus apart under 
the name of Vicroretus. They are all small, some being less than 
a rat in ae long-tailed, and nocturnal in their habits. ‘One of the 
largest,  pureifer, is of a reddish-gray colour, and distinguished 
by ¢ a a median stripe on its back which divides on the “top of 
the head into two branches, one of which passes forwards above 
each eye. The most interesting peculiarity of these animals, a 
knowledge of which we owe to M. Grandidier, is that certain species 
(C. samati, C. glirsides, ('. milii, ete.) during the dry season coil them- 
selves up in holes of trees and pass Into a state of torpidity like 
that of the hibernating animals in the winter of northern climates. 
Before this takes place an immense deposit of fat accumulates 
upon certain parts of the body, especially upon the basal portion of 
the tail, which has then dimensions corresponding to that of the 
well-known tat-tailed Sheep of the Cape, but which by the time 
they emerze from their torpor has acquired its normal proportions. 
The smallest species, to which many names have been given 
(C2 pusillus, rufus, sinithi, ete.), lives among the small branches on 
the tops of the highest trees, feeding on fruit and insects, and 
making nests which resemble those of birds. 

Galago.\.—Last upper premolar with two large external cusps, 
and nearly equalling the first molar in size. Calecaneum about one- 
third the length of the tibia, and the navicular much longer than 
the cuboid. Vertebre: C7, D 13, L 6,8 3, C 22-26. Tail long, 
and generally bushy. Ears large, rounded, naked, and capable of 
being folded at the will of the animal. Mammze four, two pectoral 
and two inguinal. 

The Galagos differ from all the Lemuroids previously mentioned, 
inasmuch as they are inhabitants, not of Madagascar, but of the 
African continent, being widely distributed in the wooded districts 
from Senegambia in the west to Abyssinia in the east, and as far 
south as Natal. They pass the day in sleep, but are very active at 
night, feeding on fruit, insects, and small birds. When they 
descend to the ground they sit upright, and move about by jump- 
ing with their “hind legs, like jerboas and kangaroos. They are 
pretty little animals, varying in size from that of. a small cat to less 
than a rat, with large eves and ears, soft woolly fur, and long tails. 
There are several species, of which G. crass sicandutus, from Mozam- 
bique, is the largest. A similar species, or perhaps variety, from 

1 Geoffroy, v7. Encyclop, 2d ann. vol. i. p. 49 (1796). 


LEMURIDZ 691 


Angola is G. montieri. G. garnetti, alleni, maholi, demidofi, and 
senegalensis are other recognised species. The last-mentioned was 
the first known to science, having been brought from Senegal by 
Adanson, and described in 1796 by Geoffroy, who adopted the 
name Galago, by which it was said to be called by the natives. 

Subfamily Lorisinze.—Dental formula as in Lemurine. Index 
finger very short, sometimes rudimentary and nailless. Fore and 
hind limbs nearly equal in length. Tarsus not specially elongated. 
Pollex and hallux diverging widely from the other digits, the hallux 
especially being habitually directed backwards. Tail short or quite 
rudimentary. Mammez two, pectoral. 

A small group of very peculiar animals, of essentially nocturnal 
habits, and remarkable for the slowness of their movements. They 
are completely arboreal, their limbs being formed only for climbing 
and clinging to branches, not for jumping or running. They have 
rounded heads, very large eyes, short ears, and thick, short, soft 
fur. They feed not only on vegetable substances, but, like many 
of the Lemuride, on insects, eggs, and also birds, which they steal 
upon while roosting at night. None of the species are found in 
Madagascar. One of the greatest anatomical peculiarities of these 
animals is the breaking up of the large arterial trunks of the limbs 
into numerous small parallel branches, constituting a rete mirabile, 
which is found also in the Sloths, with which the Loris are some- 
times confounded on account of the slowness of their movements. 
The animals of this group are usually divided into four genera, 
though the characters by which they are separated are very trivial. 
There are more properly two natural divisions. 

A. Characterised by the index finger being small, but having 
the complete number of phalanges, and by their Asiatic habitat. 

These form the genus Loris of Geoffroy St. Hilaire (1796), 
Stenops of Illiger (1811), but they were in 1812 divided by Geoffroy 
into two genera, Mycticebus and Loris, a division which has been 
accepted by most modern zoologists. 

Nycticebus..First upper incisor larger than the second, which 
is often early deciduous. Inner margins of the orbits separated 
from each other by a narrow flat space. Nasal and premaxillary 
bones projecting but very slightly in front of the maxilla. Body 
and limbs stout. No external tail. Vertebre: C7,D17,L 6,58 3, 
C12. The species are WV. tardigradus, the common Slow Lemur or 
Loris, of the Malay Countries, Sumatra, and Borneo; NV. javanicus, 
of Java; and WN. cinereus (Fig. 329) of Siam and Cochin China. The 
habits of all are much alike. They lead a solitary life in the 
recesses of large forests, chiefly in mountainous districts, where they 
sleep during the day in holes or fissures of large trees, rolled up 
into a ball, with the head between the hind legs. On the approach 

1 Geoffroy, Ann. du Muséum, vol. xix. pp. 162, 163 (1812). 


692 PRIMATES 


of evening they awake; and during the night they ramble among 
the branches of trees, slowly and quietly, in search of their food, 
which consists of tender leaves and fruit, small birds, insects, and 
mice. When in quest of living prey they move noiselessly till quite 
close, and then suddenly seize it with one of their hands. The 
female produces but one young one ata time. L. tardigradus was 
placed by Linneus at the head of the list of species of his genus 
Lemur, and its habits doubtless suggested the generic name which 


Fic. 329.—The Gray Loris (Nycticebus cinereus). From A. Milne-Edwards, N. Archives 
du Muséum, vol. iii. pl. 3. 


was transferred by Geoffroy to the less nocturnal and spectre-like 
Madagascar members of the group.! 

Loris.2—Upper incisors very small and equal. Orbits very large, 
and only separated in the middle line above by a thin vertical plate 
of bone. Nasals and premaxille produced forwards considerably 
beyond the anterior limits of the maxille, and supporting a pointed 
nose. Body and limbs slender. No external tail. Vertebre: C 7, 
D 14,L 9,8 3,C6. This genus is represented only by the Slender 
Loris (Z. gracilis) of Southern India and Ceylon (Fig. 330). This 
species is common in some of the forest regions of Southern India, 
and may be purchased in the bazaars at Madras, its eyes being 
regarded as a remedy by the natives for ophthalmic diseases. It is 
a strange-looking creature, about the size of a squirrel, of a 
yellowish-brown colour, with large, prominent eyes, pointed nose, 


1 For the anatomy of this genus, see J. L. C. Shroeder van der Kolk and 
W. Vrolik, ‘‘ Recherches d’Anatomie comparée sur le genre Stenops d’Illiger,” in 
Bijdragen tot de Dierkunde, Part I, Amsterdam, 1848-54. 

? Geoffroy, Mag. Encyclop. 2d ann. vol. i. p. 48 (1796). 


LEMURID.1- 693 


long thin body, long, angularly bent, slender limbs, and no tail. 
Its habits, according to Mr. W. T. Blanford,! are “very similar 
to those of 
Nyeticebus tardi- 
gradus, except 
that the Slender 
Loris is rather 
quicker in its 
movements, 
though still slow 
in general. Like 
its ally, it is 
purely nocturnal 
and arboreal, 
living upon 
shoots and young | 
leaves, insects, 
birds’ eggs, birds, 
and lizards. It 
is said to be very 
fond of honey or 
syrup. It sleeps 
rolled up in a 
ball with its head between its legs, grasping its perch with its arms.” 

B. Index fingers reduced to a mere tubercle without nail. Both 
the known species are from West Africa. 

Perodicticus.°—A short tail, about a third of the length of the 
trunk. Two or three of the anterior dorsal vertebra have very 
long slender spinous processes which in the living animal project 
beyond the general level of the skin, forming distinct conical pro- 
minences, covered only by an exceedingly thin and naked integu- 
ment. The Potto, P. potto, is one of the oldest known members of 
the lemuroid group, having been described in 1705 by Bosman, 
who met with it in his voyage to Guinea. It was, however, lost 
sight of until 1825, when it was re-discovered in Sierra Leone, and 
fully described by Bennett in 1830 under the name of Perodicticus 
geofroyi. Bennett's generic name has been retained, but the specific 
name bestowed by Gmelin, adopted from Bosman, has been restored. 
It is also found in the Gaboon. It is strictly nocturnal, and slower 
in its movements even than Nycficebus tardigradus, which otherwise 
it much resembles in its habits. 

A second species, the Awantibo (P. calabarensis), rather smaller 
and more delicately made, with smaller hands and feet and rudi- 
mentary tail, constitutes the genus —frefocebus of Gray. It is found 

1 Mammalia of British India, p. 48 (1888). 
2 Bennett, Proc, Zool. Soc. 1839, p. 109. 


Fic. 330,—The Slender Loris (Loris grucilis). Fron. Blanford, 
Mammalia of British India, p. 47. 


694 PRIMATES 


at Old Calabar, and is very rare, only a few individuals having as 
yet been met with. Vertebrae: C 7,D 15,L. 7,8 3,C 9.1 


Family TARSIUD. 


Dentition: i 2, ¢ 4, p 3, m 3; total 34. The first upper 
incisor large, and in contact with its fellow of the opposite side. 
Canine of moderate size. Molars with numerous pointed cusps. 
Lower canine semi-erect, its apex diverging from that of the single 
incisor. First lower premolar smaller than those behind it. Orbit 
to a large extent separated from the temporal fossa by a bony 
partition. Fibula slender, with its lower half confluent with the 
tibia. Second and third digits of the hind foot with compressed 
claws ; all the other digits of both feet with flat nails. Calcaneum 
and navicular bone of the foot elongated as in the Chirogales and 
Galagos, but to a still greater extent. Colon short and not folded. 
Vertebre : C7, D 13, L 6,8 3, C 27. 

Tarsius.2—The family contains the single genus Tarsius, of 
which but one species is known, 7. spectrum, the Tarsier, a very 
singular little animal, rather smaller than an English squirrel, with 
very large eyes and ears, a long thin tail, tufted at the end, and 
immensely elongated tarsal portion of the foot, in allusion to which 
its generic name was given to it. It inhabits the forests of many 
of the islands of the Indo-Malayan archipelago, including Sumatra, 
Borneo, Celebes, and some of the Philippines, feeds chiefly on insects 
and lizards, sleeps during the day, but is tolerably active at night, 
moving chiefly by jumping from place to place, an action for which 
the structure of its hind legs, which present a curious resemblance 
to those of a frog, seems particularly well adapted. It is rare, not 
more than two being generally found together, and only brings 
forth one young ata time.? 


Family CHIROMYIDE. 


Dentition of adult: 7 4, ¢ 3, p $, m3; total 18. Incisors very 
large, compressed, curved, with persistent pulps and enamel only in 
front, as in Rodents. Teeth of cheek series with flat, very indis- 
tinctly tuberculated crowns. In the young the first set of teeth 
more resemble those of the normal lemurs, being i 2, ¢ 3, m 2, all 
very small. Orbit surrounded by a ring of bone posteriorly, beneath 
which it communicates freely with the temporal fossa. Fibula well 


1 For the anatomy of P. potto, see Van der Hoeven and Van Campen (Ontlced- 
kundige Onderzock van den Potto van Bosman, 1859) for P. calabarensis, Huxley, 
Proc. Zool. Soc. 1864, p, 314. * Storr, Prodromus Meth. Mamm. (1780). 

3H. Burmeister, Beitriige zur niéihreren Kenntiiss der gattung Tarsius, 1846, 


CHIROM VID. 695 


developed and distinct from the tibia. All the digits of both feet 
with pointed rather compressed claws, except the hallux, which has - 
a flattened nail. Middle digit of the hand excessively attenuated. 
Vertebrae: C 7, D 12, L 6,8 3, © 27. 

Chiromys..cThis family, like the last, is formed for the recep- 
tion of a single genus, Chiromys,? containing one species, C. mada- 
gascariensis, the Aye-aye, an animal about the size of a cat, with a 
broad rounded head, short face, and large and naked ears. It has 
very large hands and long thin fingers with pointed claws, one of 
which (the middle 
or third) is remark- 
able for its extreme 
slenderness.. The 
foot resembles that 
of the other lemurs 
in itslarge opposable 
hallux, with a flat 
nail, but all the 
other toes have 
pointed compressed 
claws, like that of 
the second toe in 
the Lemurine and 
the second and third 
in the Turstide. Tail . 
long and bushy. General colour dark brown, the outer fur being 
long and rather loose, with a woolly undercoat. Mammz two, 
inguinal in position. It is a native of Madagascar, where it was 
discovered by Sonnerat in 1780. The specimen brought to Paris by 
that traveller was the only one known until 1860. Since then many 
others have been obtained, and they may frequently be seen living in 
the gardens of the Zoological Society of London. Like so many of 
the Lemurs, the Aye-aye is completely nocturnal in its habits, living 
either alone or in pairs, chiefly in the bamboo forests. Observations 
upon captive specimens have led to the conclusion that it feeds princi- 
pally on succulent juices, especially of the sugar-cane, which it obtains 
by tearing open the hard woody circumference of the stalk with its 
strong incisor teeth. It is said also to devour certain species of 
wood-boring caterpillars, which it obtains by first cutting down 
with its teeth upon their burrows, and then picking them out 
of their retreat with the claw of its attenuated middle finger. It 


Fia, 331.—Skull of Aye-aye (Chiromys madagascariensis) xX £ 
Mus. Roy. Coll. Surgeons. 


1 Cuvier, ‘Table de Class.” in Légons @’ Anat. Comp. vol. i. (1800). 

2 It was first named Daubentonia by Geoffroy; but this name was withdrawn 
by its author in favour of Chiromys, as it had been previously given to a genus 
in the vegetable kingdom. This would not, however, constitute preoccupation 
according to the modern rules of nomenclature. 


696 PRIMATES 


constructs large ball-like nests of dried leaves, lodged in a fork 
of the branches of a tree with the opening on one side. The 
resemblance of its teeth to those so characteristic of the Rodentia 
caused it to be placed formerly in that order, and it was only when 
its anatomical characters were fully known that its true affinities 
with the Lemurs became apparent.! 


Extinct LEMUROIDS. 


The discoveries of the last few years have revealed the former 
existence, both in Europe and North America, of a number of 
extinct animals more or less closely allied to the living Lemurs, 
which are of especial interest as showing in some instances characters 
of a more generalised type than is the case with the living 
representatives of the suborder. It is, however, in some cases very 
difficult to determine whether these extinct forms should be referred 
to the Lemuroidea or Insectivora ; and if those naturalists are right 
who regard these groups as survivors of a very generalised ancestral 
type of mammalian organisation, it is to be expected that as we 
recede in time we should find that the two groups show more and 
more marked signs of a natural connection. The earliest reference 
of one of these extinct Upper Eocene types to the Primates was 
made in 1862 by Professor L. Riitimeyer, of Basle, who described 
part of an upper jaw with three teeth from the so-called Bohnerz 
of Egerkingen, near Soleure in Switzerland, under the name of 
Cenopithecus lemuroides, regarding the animal to which the specimen 
belonged as partaking of the characters both of the Lemurs and the 
American Monkeys. Most other paleontologists refused, however, 
to accept this determination, and it was not until many years 
later that the researches of Gaudry and Filhol showed not only 
that Ccnopithecus was indeed a true Lemuroid, but also that it was 
either identical with or closely allied to a form described by Cuvier 
in the early part of this century under the name of Adapis and 
regarded as referable to the Ungulata. Later researches have 
brought to light other Lemuroids in the Tertiaries of both the Old 
and the New World ; and it is very noteworthy that all these types 
seem to have disappeared from both regions with the close of the 
upper portion of the Eocene period. 

Among the more interesting of the forms which are generally 
regarded as true Lemuroids we may first mention a small species 
from the Quercy Phosphorites, of which the hinder cheek-teeth are 
shown in Fig. 332, 4, which was originally described as Necrolemur 
antiquus, but appears to be generically identical with JLicrocherus 

1 R. Owen, “On the Aye-aye,” in Zrans. Zool. Soc. 1862, vol. v. p. 38 ; 
W. Peters, ‘Ueber die Siiugethier-Gattung Chiromys,” in Abhand. Kénigl. 
Akad. der Wissenschaften, Berlin, 1865, p. 79. 


EXTINCT LEMUROIDS 697 


erinaceus, of the upper Eocene of Hampshire, of which the corre- 
sponding teeth are shown in B of the same figure. In this genus, 
according to Dr. Schlosser, the dental formula is i 2, ¢ 4, p 3, m 3, 
or the same as in the existing Twrsius ; but it is not improbable that 
in some instances the first lower premolar may have been developed. 
The upper molars of Af erinaceus differ from those of MZ. antiquus 
by the simpler structure of their columns and the smaller size of 
the external cingulum, which lacks the median cusp found in the 
latter. The angle of the mandible is produced into a large hook- 
like flange which at once 
distinguishes the genus 
from all existing Lemurs; 
and the anterior lower 
premolar is not canine- 
like. M. antiquus is of 
very small size, but the 
larger M. edwardsi of the 


same deposits comes : 

nearer in dimensions to Fic, 332.—The last five right upper cheek-teeth of Micro- 
. cherus antiquus (A) and Microcherus erinaceus (B). Twice 

M. erinaceus. The upper natural size, and natural size. 


molars decrease in size 

from the first to the third, the first and second having a median 
cusp in the external cingulum, by which they are readily dis- 
tinguished from the corresponding teeth of the under-mentioned 
genus Hyopsodus. The third upper molar differs from that of 
Hyopsodus by its small size and the abortion of its posterior columns. 
The skull approximates to that of the living genus Galago, exhibit- 
ing the same inflation of the auditory bulla. The upper molars 
are also not unlike one species of that genus, but the fourth upper 
premolar has but one outer cusp, as in Chirogaleus. 

The small ./naptomorphus, from the North American Eocene, 
has a skull of about the same size as that of the smallest species of 
Microcherus, but the dental formula is i 2, ¢ 4, p 2, m 3, and the 
upper molars are of the tritubercular type. 

The well-known Adapis (Aphelotheriwn or Paleolemur), of the 
Upper Eocene of France and England, differs from all existing 
Lemuroids in possessing four premolars!; the dental formula being 
i 2,¢4,p4,m 3. The fourth upper premolar has two outer cusps, 
and the upper molars (Fig. 333) resemble those of Lepidolemur and 
Hapalemur, while the lower canine is a well-developed tooth per- 
forming the usual function of biting against the canine of the upper 
jaw. The lower incisors have upright, spatulate crowns, as in the 
true Apes. The skull is said to approximate in contour to that of 
Propithecus. The typical A. parisiensis is of comparatively small 
size, but the species of which the upper cheek-teeth are shown in 


1 One specimen has been seen with only three lower premolars. 


698 PRIMATES 


the woodcut is of much larger dimensions. The skull of 4. magna, 
which measures upwards of 4 inches in length, resembles that of 
A. parisiensis in its general characters, but is modified much in the 
way that the skulls of larger animals differ from the smaller ones of 
the same natural group. Thus the brain-chamber and orbits are 
relatively smaller, the face larger, the muscular crests more 
developed, and the constriction be- 
tween the cerebral and the facial 
portion of the skull more marked. 
These modifications remove the skull 
in its general characters still farther 
from the existing Lemurs—so much 
so that M. Filhol refers it and the 
other species of Adapis to a distinct 
Fic. 33.—The left upper cheek-teeth_ roolocical type, intermediate between 
of Adapis magna, from the Upper Eocene 
of Hampshire. the lemurs and the pachyderms, to 
which he gives the name of Pachy- 
lemuriens, but later researches do not support this view. As 
mentioned above, it has been suggested that Cenopithecus lemuroides 
is inseparable from Adapis parisiensis, but the postero-internal 
column of the upper molars is said to be larger. The genera 
Tomitherium and Notharctus, of the Eocene of the United States, 
appear to be allied to Adapis, but the second has a larger lower 
canine. The same deposits have also yielded more or less imper- 
fect remains of other forms departing more widely from the existing 
Lemuroid type. Of these Hyopsodus, of the Wasatch and Bridger 
Eocene of the United States, has the dental formula i 2, ¢ 4, p 4, 
m 3. The quadrituberculate upper molars have well-developed 
accessory intermediate columns (protoconule and metaconule), and 
thus resemble those of Microcherus,; the external surfaces of the 
outer columns of their teeth being flattened, with vertical ridges 
and a distinct cingulum. The third upper molar has its postero- 
internal column (hypocone) partly aborted, but is otherwise as well 
developed as the preceding molars. Microsyops, of the North 
American Kocene, appears to have been 
an allied form in which there were prob- 
ably only three premolars. 

The genera Protoadapis and Plesiadapis, 
from the lowest Eocene of Rheims, may 
not improbably be regarded as primitive 
Lemuroids. The lower molars are quin- iq. 334.—'The right upper 
quetubercular, and not unlike those of cheek-teeth of Plesiadapis remen- 
Microsyops ; the dental formula of the 74)" me Ciera 
lower jaw is 7 2, ¢ 1, p 3-4, m 3 in the m, 1, 2,3, molars. (From Osborn.) 
first-named genus, but in the second the 
dentition is reduced to i 2, ¢ 3, p 2, m3. In Plesiadapis the lower 


ANTHROPOIDEA 699 


and the first upper incisor are enlarged, the upper molars 
(Fig. 334) tritubercular, and the lower quadritubercular. Indrodon, 
of the lowest Eocene of the United States, resembles Plesiadapis in 
its tritubercular upper molars, and appears to have a nearly similar 
dental formula. Miaodectes, of the same deposits, was probably a 
more or less closely allied type. Pelycodus of the Wasatch Eocene 
of North America, in which the hallux was not opposable, and 
Cryptopithecus of the German Eocene, may be regarded as very 
generalised Lemuroids. 


Bibliography.—Besides the works and memoirs on particular families and genera 
referred to above, see St. G. Mivart, ‘‘Notes on the Crania and Dentition of 
the Lemuride,” in Proc. Zool. Soc. 1864 (pp. 611-648) and 1867 (pp. 960-975) ; 
Mivart and Murie, ‘‘On the Anatomy of the Lemuroidea,” in Trans. Zool. Soc. 
1872, vol. vii. pp. 1-113; W. Turner, ‘‘On the Placentation of the Lemurs,” in 
Phil. Trans, vol. clxvi. pp. 569-587 ; F. Pollen and D. C. Van Dam, Recherches 
sur la Faune de Madagascar, 2”° parte, ‘‘Mammiferes,” 1868. For the fossil 
types see M. Schlosser, ‘‘Die Affen., Lemuren, etc,, des Europiiischen Tertiars,” 
in Beitr. Pal. Gstr-Ungar, 1888. 


Suborder ANTHROPOIDEA. 


This suborder includes the whole of the remaining members of 
the Primates, namely, those animals commonly known as Marmosets, 
Monkeys, Baboons, and Apes, together with Man himself. The 
characters by which the Anthropoidea are distinguished as a whole 
from the Lemuroidea may be summarised as follows. Skull with 
the orbit separated from the temporal fossa by a vertical plate of 
bone joining the postorbital bar, and the lachrymal foramen situated 
within the margin of the orbit. Pollex sometimes rudimentary or 
absent; second digit of manus always well developed, and that of 
the pes usually with a flattened nail (not so in Hupalide). The 
cerebral hemispheres of the brain either completely or almost 
completely cover the cerebellum, and are much convoluted. 
Uterus not bicornuate. The placenta is deciduate and discoidal ; 
and the allantois is small. There are never abdominal mammex. As 
additional points of distinction from the Lemuroidea, it may be 
mentioned that the anterior cornu of the hyoid is shorter than the 
posterior; the inner pair of upper incisors are in contact in the 
middle line ; and the transverse portion of the colon extends unin- 
terruptedly across the abdomen. 

The Anthropoidea may be divided into the five families—Hapa- 
lide, Cebide, Cercopithecide, Simiide, and Hominide, of which the 
first and second are confined to the New, and the third and fourth 
to the Old World. 

In noticing some of the salient features in the external and 
internal structure of the Anthropoidea it will be found convenient 


700 PRIMATES 


to allude to all the members of the first four families as Apes, in 
contradistinction to Man. In respect to relative size the extremes 
are found in the Gorilla on the one hand and Hapale on the other; 
the difference in this respect between these two forms being greater 
than that between Man and a Squirrel. The relative proportions 
between the limbs and the body, and also between the fore and 
hind limbs, are subject to great variation. Thus in Hylobates and 
Aieles both pairs of limbs are much elongated; in the former case 
the pectoral being much longer than the pelvic pair (Fig. 335). 
In other cases, as in the Orang (Fig. 354), while the arms are very 
long, the legs are short; but in the ‘subfamily Cercopithecine both 
pairs are short and subequal. Only in the Hapalide and some of 
the Cebide are the legs proportionately as long as in Man. 

The tail is as much as three times the length of the body in 
Aleles ; while in the Simiide it is totally absent. In the majority 
of genera it is long in all the species; but in some cases, as in 
Macacus, it may be either long, short, or absent in the different 
species of a single genus. 

Equally marked variations occur in the shape of the head. 
Thus in Ateles it is rounded; while in the Orang it is elevated 
vertically ; in Chrysothrix it is produced posteriorly ; and in the 
Baboons (Cynocephalus) it is characterised by the great production 
of the muzzle and the terminal position of the nostrils, whereby a 
characteristic Dog-like form is assumed. The eyes are always 
directed forwards, and are never more separated from one another 
than in Man, although, as in Chrysothriz, they may be closer 
together. They are of very large size in Nyctipithecus, while in the 
Baboons they are relatively small in proportion to the size of the 
head. The ears are invariably well developed, and are usually 
pointed at their postero-superior angle. Those of man are charac- 
terised by the soft depending portion known as the “lobule,” of 
which there is a rudiment in the Gorilla. In the majority of Apes 
the nose is but very slightly prominent; but it attains an extra- 
ordinary development in Nasalis larvatus, and is scarcely less 
remarkable in Semnopithecus rowellane (Fig. 349). Among the 
Gibbons the Hoolock has a distinctly aquiline nose. The nostrils 
are terminal in the true Baboons; and while in all the Old World 
Apes they are approximated, in those of the New World they are 
separated by a broad septum. With the exception of the Orang, 
the lips of the Apes are thin. 

The pollex makes a nearer approach in form to the human 
thumb in the Chimpanzee than in any other Ape. Man differs 
from all the Apes in having the hallux frequently longer instead of 
shorter than the other digits of the foot. The hallux of the Orang 
is peculiar in having no nail, but in other cases the nail is flat; the 
nails of the other digits of the Apes are never quite flat, and in 


ANTHROPOIDEA 701 


some of the Cebidw they are decidedly compressed laterally, while 
in the Hapalide they assume the form of sharp and curved claws. 

All the Apes have the greater part of the body well clothed 
with hair. In the Gibbons and the Cercopithecide the buttocks have 
naked ischiatic callosities, which attain their greatest development 
in Cynocephalus and its allies. The male of the Orang has a well- 
developed beard, and in Cercopithecus diane there is long hair on the 
cheeks and chin, while in Macacus silenus the face is surrounded by 
a fringe of long hair, separated by an interval on the forehead. 
Long hair is found on the head in Hapale wdipus and in some species 
of Semnopithecus; while in the Bonnet Monkey (Afucacus sinicus) it 
radiates in all directions from a central point on the vertex. Long 
hair clothes the shoulders in Cynocephalus humadryas and Hapale 
humeralifer; and the end of the tail has a tuft in two species of 
Cynocephalus and in Afacacus sinicus. Many of the African Colobdi 
and some species of the Howlers have very long hair on the flanks ; 
and in Pithecta this development of hair extends to the greater part 
of the body and the tail, P. safanas also having a long beard. In 
all the lower Apes the hairs on the arm and fore-arm are directed 
towards the hand quite down to the wrist ; and the same arrange- 
ment obtains in Hylobates. In the other Simiide, however (as in 
man), the points of the hairs of the arm and fore-arm converge 
at the elbow. Darwin’s explanation of this peculiarity is that these 
Apes are accustomed to sit with the arms bent, so that the rain is 
thus enabled to run off at the elbow. 

In one species of Hapale the hair is of a silky texture, and in 
the South American Lrivdes and Afacacus tibefanus (as in all the 
mammals inhabiting the arid and severe climate of Tibet) it becomes 
woolly. 

The development of very brilliant colours on the naked parts of 
the body, such as the face, sexual organs, and ischiatic callosities is 
a marked feature of many of the Cercopithecid and some other Apes. 

With the exception of the long tail found in most forms, the 
general structure of the skeleton of the Apes is very similar to 
that of man, but there are marked differences in the form of the 
jaws and of the innominate bones. The proportion of the facial to 
the cranial region of the skull varies with the shape of the head, 
of which brief mention has already been made; the greatest 
development of the facial portion being in the Baboons. Curiously 
enough, some of the lower American Monkeys, and more especially 
Chrysothrix, have the greatest relative development of the cranial 
part of the skull of all the Apes; this character being, however, 
one common to all the smaller representatives of particular groups, 
and obviously necessary to provide the requisite amount of brain- 
space. In the convexity of the frontal region of the skull the 
American: forms, and more especially Pithecia, make the nearest 


702 PRIMATES 


approximation to man, and the same is true with regard to the 
occipital production, which is most developed in Chrysothriz. - Most 
of the Simide exhibit, however, a distinct convexity of the occiput, 
and thereby differ markedly from the Cercopithecide, in which this 
region is flat. The rotundity of the cranium is obscured in the 
larger Apes, such as the Orang (Fig. 353) and Gorilla, by the 
development of prominent bony ridges for muscular attachment ; 
these attaining their maximum in the males of the species last 
named, where the sagittal crests and the supraorbital ridges are 
very prominent. The mastoid process is always smaller in the 
Apes than in Man, and as it diminishes in size the petrosal tends 
to assume an inflated or bullate condition. The orbits in shape 
are most like that of Man in the Gorilla; and, in accordance with 
the size of the eyes, they are of enormous dimensions in Nyctipithecus. 

The angle formed by the plane of the foramen magnum with 
that of the basicranial axis is subject to variation according to the 
degree of convexity of the occiput, but is generally smaller than in 
Man, although larger in Chrysothriz. There is an external bony 
meatus auditorius in Man, the Simide, and the Cercopithecide, but 
none in the Cebide and Hapalide. 

The premaxille of the Apes are always large; and, except in 
the Chimpanzee, the premaxillo-maxillary suture persists until after 
the permanent dentition has been developed. The nasals are 
smaller and flatter than in Man, but are largest in A/ycetes. The 
two rami of the mandible are invariably completely ankylosed at 
the symphysis in the adult. The Siamang (Hylobates syndactylus) is 
the only ape in which the mandibular symphysis shows a slight 
projection in front corresponding to the human chin. In Mycetes 
the angle of the mandible attains an enormous development (Fig. 
338) to protect the huge inflated basihyal. The frontal sinuses, 
though present in the Simiidw, are generally replaced in the 
Cercopithecide by a coarse diploé, but they are present in the 
Cebide and Hapalide, being especially large in Cebus. In fully 
adult individuals the cranial sutures become obliterated, the inter- 
nasal suture disappearing at an early age in the Simiidw and most 
of the Cercopithecide. As in many Carnivora, the tentorium, or 
membrane separating the cerebrum from the cerebellum, may 
become ossified in some of the American forms. 

The number of the teeth in the Old World Apes is invariably 
the same as in Man, namely 7 3, ¢ 4, p 3, m 3, total 32; but in the 
Cebide the cheek-teeth are p 3, m 3, and in the Hapalidw p 3, m 2. 
It is probable that the two pairs of incisors correspond to the first 
and third of the typical series of three. In all Apes the dental 
series is interrupted by a diastema, and the canines of the males 
are large. Man alone has an uninterrupted dental series of a 
horse-shoe-form, without prominent canines. 


ANTHROPOIDEA 703 


According to recent researches the Chimpanzee and some of the 
other Simidw exhibit a more or less close approximation to the 
sigmoid curvature of the vertebral column which is so characteristic 
of Man, and there is also some approach to it in the Baboons. 
The number of dorsal vertebree in the Apes may vary from eleven, 
as in some species of Cercopithecus and Macacus, to fourteen in 
certain forms of Hylobates, and to fifteen in Nyctipithecus. The 
Cebidew generally have thirteen; and the same number obtains in 
the Chimpanzee and Gorilla, while the Orang resembles Man in 
having but twelve. The lumbar vertebre show a range in number 
of from four to seven. In the Stmiide there are four or five of 


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lf 
LF 


Fic. 335,—Skeleton of the Black-handed Spider Monkey (Ateles geoffroyi). From De Blainville. 


these vertebrae, the length of the lumbar region being shorter in 
this family than in the other Apes, with the exception of <dAfeles. 
The shortness of the lumbar region in the last-named genus is 
compensated by the relative length of the dorsal region, as is 
shown in Fig. 335. 

The sacrum is longest in the Siwiide and Man, its greatest 
absolute length occurring in the Gorilla, and the relative greatest 
length being found in Hylobates, The Simiide never have less than 
five, and may have six sacral vertebre ; while in the lower forms 
there are generally only two or three, although occasionally four in 
some of the American forms. The Orang and some of the Baboons 
make the nearest approximation to Man in the marked angle 
formed at the junction of the sacrum with the lumbar vertebra. 
Except in the Simiide and JMacacus inuus, the number of caudal 
vertebree in the Apes always exceeds four, but they may be 


704 PRIMATES 


reduced to five in the Mandrill (Cynocephalus maimon). In Macacus 
and Uacaria the shortness of the tail is attained by the small 
size of the vertebre themselves, the number of which may be 
from fifteen to seventeen. Other forms usually have from twenty 
to thirty-three caudals, the latter number occurring in Afeles 
(Fig. 335), where the tail is relatively the longest. The tail is, 
however, absolutely longest in Semnopithecus, Colobus, and their 
allies, the length being partly due to the size of the component 
vertebra. Chevron bones are present in all forms having a distinct 
tail; and, together with other processes for muscular attachment, 
attain their greatest development in Afeles. 

The vertebral processes known as metapophyses and anapophyses, 
which are generally inconspicuous in Man, and are but small in the 
Simiide, attain a large development in the lower forms. The 
metapophyses generally commence in the eighth or ninth dorsal, 
and continue to the anterior caudals, where they gradually merge 
in the prezygapophyses. The anapophyses, which are most de- 
veloped in the Cebidw, project outwards and backwards from one 
vertebra to embrace the prezygapophyses of the succeeding one. 
They occur generally in the same region as the metapophyses, but 
usually cease at the penultimate lumbar, although in some cases 
they can be traced on to the posterior cervicals and anterior 
caudals, in the latter region passing into the transverse processes. 

In most Apes the sternum is narrow, and consists of a more or 
less enlarged manubrium, followed by a chain of subequal and 
antero-posteriorly elongated bones, from three to six in number. In 
the Simzide alone is there a broad sternum, or one consisting of a 
manubrium, followed by a single bone only, as in Hylobates. The 
Orang presents a peculiarity, in that the sternum long remains 
made up of ossifications arranged in pairs, side by side, successively. 
The true ribs are seven in number on each side in the highest 
forms, but in Hylobates there are sometimes eight. In Ateles there 
are sometimes nine pairs. In Hapale the number varies from six 
to eight, and it is seven or eight in the other genera. The angles 
of the ribs are never so marked as in Man; although most marked 
in Hylobates.  Pithecia is distinguished by the greater relative 
breadth of the ribs. In no Ape is the thorax half as broad again 
as it is deep from back to breast ; but in the Simiidw its transverse 
diameter exceeds its depth by from about one-fourth to a little 
under one-third of the latter. In Ateles, and sometimes in Mycetes, 
the thorax is wider than deep, but in all the rest it is deeper than 
wide. 

In regard to the appendicular skeleton it may be observed that 
the Gorilla and Orang make the nearest approach to Man in the 
form of the scapula; and that the supraspinous fossa of this bone is 
largest in Gorilla and Afycetes, being remarkably small in Simia. 


ANTHROPOIDEA 705 


The Cebide have a distinct suprascapular notch which is often 
converted by a bar of bone into a foramen; this bar in Mycetes 
giving rise to a peculiar flat process. The acromion and coracoid 
processes are most developed in the Simiide and Ateles. 

The relative length of the fore and hind limbs has been already 
briefly touched upon. The humerus closely resembles that of 
Man throughout the suborder ; the nearest approximation occurring 
in the Simiide. As in the Lemuroidea, this bone never has an 
entepicondylar foramen, but in many of the American forms it has 
a supracondylar perforation. The radius and ulna, like the tibia 
and fibula, are always perfectly distinct throughout their length ; 
and the hand can-be pronated and supinated upon the forearm. 
Man, the Gorilla, and the Chimpanzee differ from other forms in 
having no os centrale in the carpus. 

The brain of Apes is always much smaller in absolute dimensions 
than in Man. Thus, according to Professor Mivart,! “the cranial 
capacity is never less than 55 cubic inches in any normal human 
subject, while in the Orang and Chimpanzee it is but 26 and 27} 
cubic inches respectively. The relative size of the brain varies 
inversely with the size of the whole body, but this is the case in 
warm-blooded vertebrates generally. The extreme length of the 
cerebrum never exceeds, as it does in Man, two and a quarter times 
the length of the basicranial axis. The proportion borne by the 
brain to its nerves is less in the Apes than in Man, as also is that 
borne by the cerebrum to the cerebellum. In general structure 
and form the brain of Apes greatly resembles that of Man. Each 
half of the cerebrum contains a triradiate lateral ventricle, and 
though in some Cercopithecide the posterior. cornu is relatively 
shorter than in man, it again becomes elongated in the Cebide, and 
in many of the latter it is actually longer relatively than it is in 
man. The posterior lobes of the cerebrum are almost always so 
much developed as to cover over the cerebellum, the only exceptions 
being the strangely different forms Mycetes and Hylobates syndactylus. 
In the latter the cerebellum is slightly uncovered, but it is so con- 
siderably in the former. In Chrysothria the posterior lobes are much 
more largely developed relatively than they are in man. The 
cerebrum has almost always a convoluted external surface. In this 
group, however, as in mammals generally, a much-convoluted cere- 
brum is correlated with a considerable absolute bulk of body. Thus 
in Hapale (and there only) we find the cerebrum quite smooth, the 
only groove being that which represents the Sylvian fissure. In 
Simia and Gorilla and Anthropopithecus, on the contrary, it is very 
richly convoluted. A hippocampus minor is present in all Apes, 
and in some of the Cebide it is much larger relatively than it is in 
Man, and is absolutely larger than the hippocampus major. Of all 

1 Article APE, Encyclopedia Britannica, ninth edition. 
45 


706 PRIMATES 


Apes, the Orang has a brain which is most like that of Man; 
indeed, it may be said to be like Man’s in all respects, save that it 
is much inferior in size and weight, and that the cerebrum is more 
symmetrically convoluted and less complicated with secondary and 
tertiary convolutions. If the brain of Simia be compared with that 
of Gorilla and Anthropopithecus, we find the height of the cerebrum 
in front greater in proportion in the former than in the latter ; also 
the bridging convolutions, though small, are still distinguishable, 
while they are absent in the Chimpanzee. Nevertheless this 
character cannot be of much importance, since it reappears in Afeles, 
while two kinds of the genus Cebus (so closely allied as to have been 
sometimes treated as one species) differ strangely from each other 
in this respect. The corpus callosum, in Apes generally, does not 
extend so far back as in Man, and it is very short in Pithecia. In the 
Orang and Chimpanzee there are, as in Man, two corpora albicantia, 
while in the lower Monkeys there is but one. The vermis of the 
cerebellum is larger in the Cebide than in the Simiide and Cerco- 
pithecide. In all Apes below the Stiniide each lateral lobe of the 
cerebellum gives off a small lobule, which is received into a special 
fossa of the petrous bone. Certain prominences of the medulla 
oblongata, termed corpora trapezoidea, which are found in lower 
mammals, begin to make their appearance in the Cebid.” 

The organs connected with the functions of alimentation, circu- 
lation, and excretion, as well as the muscles, conform generally to 
the type obtaining in Man, of which full description will be found 
in works on human anatomy. The tongue is longer in Apes than 
in Man ; and a uvula is generally present, although rudimentary in 
the Cebide. The peculiar sacculation of the stomach in the sub- 
family Semnopithecine has been already mentioned ; this sacculation 
is most developed at the cardiac extremity, where it somewhat 
resembles a colon spirally coiled. In Hylobates the stomach is very 
like that of Man, differing only in the more elongated and distinct 
pylorus. Pithecia has a more globular stomach, while in Hapale 
the cardiac and pyloric apertures are approximated. The intestine 
of Apes is devoid of valvule conniventes, and it is only in Man and 
the Simiidw that the colon is furnished with a vermiform appendage. 
The colon varies from a fully sacculated form in Hylobates to a 
smooth one in Cebus. 

The liver of Apes is subject to a considerable amount of varia- 
tion. In the Simiide it comes more or less close to the human 
type; that of the Orangs being usually divided only into two 
principal lobes by the umbilical vein, and showing no trace of 
lateral fissures. In the Gorilla these fissures are present, so as to 
produce right and left lateral and central lobes. Hylobutes has a 
liver (Fig. 352) which perhaps is nearer to the human than that of 
any of the other Simiidw. In the Cercopithecide the liver differs 


ANTHROPOIDEA 707 


from that of the Simiide by the deeply cleft lateral fissures, and 
has a comparatively small and pointed caudate lobe. The enormous 
size of the stomach in Colobus causes the liver to be very narrow, 
and pushed to the left side. The liver of the Cebide (Fig. 336) 
and Hapalide, in ad- ; 

dition to the deeply 
cleft lateral fissures, is 
characterised by the / 
great size and quad- {ij 
rangular form of the || 
caudate lobe (c¢), which 
attains its maximum 
development in dieles. 
The gall-bladder is 
always present. 

The larynx is in many 
Apes furnished with sac- 
like appendages, which 
are variable in different 
species as regards 
number, size, and situ- 


(aa a 


. Fic. 336.—Under surface of the liver of the Black-handed 
ation. They may be Spider Monkey (Ateles melanochir). u, Umbilical fissure ; 
dilatations of the laryn- ve, vena cava; Ul, left lateral lobe ; Ic, left central lobe; re, 
geal ventricle, as in pee fe agi, ro lobe ; s, Spigelian lobe ; 
Simia, Gorilla, and 
Anthropopithecus, or they may open above the false vocal chords 
so as to be extensions of the thyro-hyoid membrane, as in Hylobates. 
There may be but a single median opening in the front part of 
that membrane at the base of the epiglottis, as in the Cercopithecide. 
There may be a single median opening at the back of the trachea, 
just below the cricoid cartilage, as in Ateles; there may be but a 
single sac, or there may be five, as sometimes in Mycetes. These 
may be enormous, meeting in the middle line in front and extend- 
ing: down to the axille, as in the Gorilla and Orang. A sac may 
occupy the cavity of the expanded body of the hyoid, as in Mycetes. 
The hyoid has its basilar part generally somewhat more convex 
and enlarged than in Man ; but in Mycetes it becomes greatly enlarged 
and deeply excavated, so as to form a great bony bladder-likestructure. 
The posterior cronua of the hyoid (thyro-hyals) are never entirely 
absent, but the anterior or lesser cornua may be so, as in Afycetes. 
The anterior cornua never exceed the posterior cornua in length ; 
but they may be (eg. in Cercopithecus) more largely developed 
relatively than in Man, and may even be jointed, as in Lagothria. 
The lungs have generally the form of those of man; but the 
right lung may have four lobes, as in Hylobates. The great arterial 
trunks in Simia, Gorilla, and Anthropopithecus are arranged as in 


708 PRIMATES 


Man. In Aylobates and the lower Apes, however, the left carotid 
artery may take its origin from the innominate artery. 

In regard to their distribution in time the earliest record that 
we as yet have of the occurrence of Apes is in the Middle Miocene 
of Europe, where forms are met with apparently so closely allied to 
some of the higher existing types that it is evident we must look 
much farther back before we can get any clue to the origin of the 
suborder. Since all the known fossil Old World Apes are referable 
to the Stmiudw or Cercopithecide, and no representatives of these 
families have been obtained from the Tertiaries of America, it would 
appear that the distinction of the Apes of the Old World from 
those of the New is of very old standing. 

At the present day Apes are mainly confined to tropical and 
subtropical regions. In the Old World Afacacus inuus is found as 
far north as Gibraltar, Jf tibetanus and Semnopithecus roxellane 
inhabit western Tibet, while in Japan we have JZ. speciosus. In the 
New World one species of 4éeles is known to occur as far north as 
latitude 19° in Southern Mexico, and may range a few degrees 
higher. To the southward species are found near the Cape, in 
Timor, and the Malay Archipelago; while in America they range 
in Brazil and Paraguay to about latitude 30°. The Tibetan species 
are found at a very high elevation; and in the outer Himalaya the 
Langurs (Semnoptthecus) may be seen in winter and spring leaping 
from bough to bough of snow-covered pines. 

Apes are very abundant in the Ethiopian and Oriental regions, 
as well as in that part of America which extends from Panama to 
Southern Brazil. Ceylon, Borneo, Sumatra, and Java may be 
mentioned as islands where Ape-life attains great development ; but 
they are unknown in Madagascar and the West Indian Islands, and 
of course in the Australasian region. 

We have already alluded to the circumstance that while the 
Simiide and Cercopithecide are exclusively confined to the Old World, 
the Cebide and Hapalide are equally restricted to the New, and we 
may accordingly proceed to notice a few points in relation to generic 
distribution. Of the larger Simidw the Gorilla and Chimpanzee 
are confined to Equatorial Africa, and the Orang to Malayana; but 
there is evidence of the former existence of a species of Chimpanzee 
(<{nthropopithecus) and not improbably of an Orang (Simic) in Northern 
India. The Gibbons (Hylobates) are now exclusively Oriental. 
Europe has only Aacacus inwus of Gibraltar, also found in Africa 
north of the Sahara, and therefore strictly Palearctic in distribu- 
tion. The Ethiopian region includes in the Cercopithecida the genus 
Colobus (the African analogue of Semnopithecus), Cercopithecus, and 
the Baboons (Cynocephalus, etc.) The Baboons range, however, into 
Arabia and Syria, and also existed during the Pliocene epoch in 
Northern India. Semnopithecus and Macacus are very characteristic 


HAPALIDE 709 


of the Oriental region ; but, as already mentioned, outlying species 
extend into various parts of the Palearctic region. Macacus has 
indeed a very wide distribution, extending from Gibraltar and 
North Africa to Japan. The allied Cynopithecus, represented only 
by C. niger of Celebes, approximates to the Baboons ; while the one 
species of Nasalis is peculiar to Borneo. Remains of Semnopithecus 
and Macacus occur in the Tertiaries of India and Europe, which also 
yield allied extinct types noticed in the sequel. 

In America, north of Panama, the genera known to be repre- 
sented are Chrysothria, Nyctipithecus, Cebus, Ateles, Mycetes and 
Hapale in Veragua; Nyctipithecus, Cebus, Ateles, and Mycetes in 
Costa Rica and Nicaragua; Ateles and Mycetes in Guatemala ; and 
Ateles in Southern Mexico. Brazil is the headquarters of the 
American Apes; but different portions of that vast region have a 
somewhat distinct Ape fauna. Thus the genus Eriodes appears in 
South-Eastern Brazil to represent the species of Ateles inhabiting the 
more northern and western parts of the empire. Southwards, the 
genera Cebus, Mycetes, Chrysothriz, and Callithria extend farthest; 
but they do not probably all extend to the farthest limit yet known, 
namely 30° 8. The species found farthest south are Mycetes caraya, 
Cebus fatuellus, and Callithrix personatus. 


Family HAPALIDA. 


2 


Dentition: i 2, c 4, p 3, m2; total 32. No bony external 
auditory meatus, a broad internarial septum, and no cheek-pouches. 
Tail non-prehensile ; no ischiatic callosities. Pollex not opposable ; 
a long, curved, and pointed claw to all the digits except the hallux. 

This family, which includes the smallest representatives of the 
suborder, commonly known as Marmosets, is confined to the New 
World. In addition to the diagnostic characters given above, it may 
be mentioned that the pollex is elongated and the hallux very 
small, while the pectoral limbs are not longer than the pelvic pair ; 
and the tail is long and more or less thickly covered with elongated 
hairs. 

The dentition of the Marmosets sufficiently distinguishes them 
from all other members of the suborder, although they are evidently 
nearly allied to the Cebide. The small size of the hallux, and the 
total incapacity of the pollex to oppose itself in the least degree to 
the other digits, as well as the presence of claws on all the digits of 
the manus, are, however, equally characteristic features. These 
animals (Fig. 337) are not larger than Squirrels, and are of active 
arboreal habits, living in small companies, and adding insects to the 
ordinary fruit diet. Frequently, as in the figured species, the head 
is furnished on either side with a long tuft of hair projecting out- 
wards and backwards. They give birth to as many as three young 


710 PRIMATES 


ones at a time, and thereby differ from all other members of the 
suborder, in which one is the normal number. They are divided 
into two genera, according to the proportionate size of the lower 
canine to the incisors; but some species present an intermediate 
condition, so as to render this distinction of somewhat doubtful 
value. 

Hapale.\—Lower canine not longer than the incisors. A number 
of species have been described, among which may be mentioned 


Fic, 337.—The Golden Marmoset (Midas chrysoleucas). From Proc. Zool. Soc. 1868. pl. 24 
3 » pl. 24. 


Al. jacchus, H. albicoilis, H. aurita, and H. humeralifer. Remains of 
species of this genus have been found in the cavern-deposits of 
Brazil. 

Midas.’—Lower canine considerably longer than the incisors, N. ft) 
less than twenty-four species of this genus have been named amon 
which the Silky Marmoset (J. rosalia) of Columbia, the Ting 
Monkey (M. edipus) of South-Eastern Brazil, and the Golden 
Marmoset (J. chrysoleucas, Fig. 337) are well-known types. 


* Hliger, Prodromus Syst. Mamm. et Avium, p. 71 (1811). 
* Geoffroy, dan. du Muséwm, vol. xix. p. 120 (1812). 


CEBID.£E Far 


Family CEBID-E. 


2 


Dentition: ¢ 2, ¢ 4. p 3, m &; total 36. Tail frequently 
prehensile : digits with nails: other characters as in the Huapalide. 

The members of this American family are at once distinguished 
by the dental formula, which is numerically higher than in any 
other Apes. The various species range over the whole of tropical 
America, but are most abundant in the dense forest regions 
of Brazil, where they live a completely arboreal life, to which the 
prehensile tails of many of them are so specially adapted. They 
are In most respects closely 
allied to the Hapalidr, but 
the pollex diverges some- 
what from the plane of the 
other digits : while the re- 
tention of the third molar 
is a very distinctive feature. 
None of the species attain 
the dimensions of the larger 
Ceorupithectde of the Old 
World. The genera are 
usually arranged in five 
subfamilies. 

Subfamily Myecetinse.— 
Lower incisors vertical : 
hyoid bones enormously 
inflated ; tail long and pre- 
hensile, naked beneath at 
the end; pollex well de 
veloped. 

Uyectes4—The sole re- 
presentatives of this sub- 
family are the well-known 
Howling Monkeys, all of 
which are included in the = 
genus JM yoctes, They are Fie. 388. Side view of skull and hyoid bone of the 
of more bulky build, and Red Howling Monkey (Mucetus seniculus), From De 
have more produced muzzles P"""* 
than the other members of the family. The truncated occipital 
region, and the extraordinary development of the rami of the 
mandible, especially of their angular and ascending portions, are 
the chief peculiarities by which the skulls (Fig. 338) of the 
members of this genus are characterised. The last named char- 
acter, which is more marked in the male than in the female sex, 


1 Wliger, Prodromus Sust. Mamin, ef Avi, p. 70 (1811). 


712 PRIMATES 


is related to the enormous size of the vocal organs, which the rami 
of the mandible enclose and protect. The inflated hyoid bone, 
which forms a deep cup, is shown in the figure. The Howlers are 
subject to great individual and sexual variation of colours, so that 
the discrimination of species from local races is difficult. In one 
species the male is black and the female straw-coloured ; and several 
of the species have bright red or golden hair on the flanks. In 
disposition these creatures are sluggish and stupid, but their chief 
characteristic is their prodigious power of voice. Mr. Bates, in his 
Naturalist on the Amazons, observes that “when Howlers are seen in 
the forest there are generally three or four of them mounted on the 
topmost branches of a tree. It does not appear that their harrow- 
ing roar is emitted from sudden alarm; at least it was not so in 
captive individuals. It is probable, however, that the noise serves 
to intimidate their enemies.” 

Several species have been described, the Red Howler (JZ. seniculus) 
and the Ursine Howler (Jf. wrsinus) being well-known forms. 
Remains of this genus probably referable to existing types are found 
fossilised in the cavern-deposits of Brazil. An allied fossil form 
from the South American Pleistocene has been described as 
Protopithecus. 

Subfamily Pitheeiinse.—Lower incisors inclined forward at their 
summits ; hyoid bone normal ; tail long or short, non-prehensile ; 
pollex well developed. Two genera are included in this subfamily, 
readily distinguished by the length of the tail. 

Pithecia.\—The Sakis, as the representatives of this genus are 
commonly termed, are readily characterised by the length of the 
tail; the angle of the mandible is expanded, although less so than 
in Mycetes. A number of species have been described, the Black 
Saki (P. satanas) of the Lower Amazons, being one of the best 
known. While some species, like P. hirsuta, have long hair covering 
the whole of the head, body, and tail, in others only the head, or 
the cheeks and chin, are so clothed. 

Uacaria.2—The Uakari Monkeys differ from all the other 
Cebide by their short Baboon-like tail. The Bald Uakari (U. calva) 
of the Rio Negro, and the closely allied U. rubicunda of the Upper 
Amazons, are remarkable for their scarlet face, which forms a striking 
contrast to the long, silky, whitish hair covering the body. Accord- 
ing to Mr. Bates, the Uakaris live in forests which are inundated 
during a great part of the year, and never descend to the ground ; 
they appear to be rare and of local distribution. The third species, 
U. melanocephala, differs considerably from both the others. The 
cecum of U. calvw, according to Mr. F. E. Beddard, measures 

’ Geoffroy, Ann. du Muséum, vol. xix. p. 115 (1812). 
* Gray, Proc. Zool. Soc. 1849, p. 9. Amended from Ouakaria : Syn. Brachy- 
urus ; Spix, Sim. et Vesp. Brasil,'p, 11 (1823). Preoceupied by Fischer, 1814. 


CEBID 713 


upwards of “10 inches along the greater curvature ; it is separated 
from the colon by a very marked constriction ; it is not sacculated, 
and when fully distended with air is curved on itself into a little 
less than a circle; it is furnished with a well-developed median 
frenum carrying blood-vessels.” A similar type of cecum is also 
found in Callithria and Pithecia. 

Subfamily Nyetipithecinz.—Lower incisors vertical; hyoid 
normal ; tail long, non-prehensile ; pollex well developed. 

Three genera are included in this subfamily, the species being 
partly insectivorous. 

Callithria..—Head small, depressed, and not elongated; nares 


Fia. 339.—The Moloch Teetee (Callithria moloch). From Archives du Muséwm, vol. iv. pl. 3. 


widely separated ; canines small; angle of mandible expanded as in 
Pithecia ; tail with long hair. 

This genus comprises several small species, mostly from Brazil 
and the Amazons, and commonly known as Teetees, one of the 
best-known species (C. moloch, Fig. 339) being represented in the 
accompanying woodcut. The smaller eyes and the more widely 
separated nostrils distinguish them from Nyctipithecus , while the 


1 Geoffroy, Ann. du Muséum, vol. xix. p. 112 (1812), 


714 PRIMATES 


small canines and the bushy tail readily mark their distinction from 
Chrysothvix. Remains of Callithria have been found in the Brazilian 
caves. 

Chrysothrix.'—Head greatly elongated ; orbits large and closely 
approximated; canines well developed; tail with comparatively 
short hair. 

The small Squirrel Monkeys, of which four species have been 


Fic. 340.—The Lemurine Douroucouli (Nyctipithecus lemurinus). From Archives du Muséum, 
vol. iv. pl. 2. 


described, are characterised by the great backward projection of the 
occipital region of the skull, and by orbits approximating in size to 
those of the next genus. 

Nyctipithecus.’—Head rounded ; orbits very large, separated by 
a narrow septum ; nares somewhat approximated. 

The Douroucoulis (Fig. 340), as the members of this genus are 
called, are of nocturnal habits, in association with which the eyes 
are of enormous dimensions, as in the Lemuroid genus Loris, The 
following account of two species of this genus is taken from Mr. 


? Kaup, Thierreich, vol. i. p. 51 (1835). 
° Spix, Sim. ct Vesp. Brasi, p. 25 (1823). 


CEBIDE 715 


Bates’s Naturalist on the Amazons: “They sleep all day long in 
hollow trees, and come forth to prey on insects and eat fruit only 
in the night. They are of small size, the body being about a foot 
long, and the tail 14 inches, and are thickly clothed with soft gray 
and brown fur, similar in substance to that of the Rabbit. Their 
physiognomy reminds one of the Owl or Tiger-Cat; the face is 
rounded and encircled by a ruff of whitish fur; the muzzle is not 
at all prominent ; the mouth and chin are small; the ears are very 
short, scarcely appearing above the hair of the head; and the eyes 
are large and yellowish in colour, imparting the staring expression 
of nocturnal birds of prey. The forehead is whitish, and decorated 
with black stripes, which in one of the species (NV. trivirgatus) 
continue to the crown, and in the other (NV. felinus) meet on the 
top of the forehead. WV. trivirgatus was first described by Humboldt, 
who discovered it on the banks of the Cassiquiare, near the head- 
quarters of the Rio Negro.” 

Subfamily Cebinze.—Lower incisors vertical; hyoid bone 
normal ; tail long and prehensile ; pollex present or absent. 

This subfamily includes the typical members of the family, 
which are arranged in four genera. 

Ateles1—Form slender; limbs very long; fur not woolly ; 
pollex absent ; tail naked beneath distally ; nails not much laterally 
compressed and pointed. 

This genus includes the well-known Spider Monkeys (Fig. 341), 
which by their long limbs and tail are admirably adapted to a 
purely arboreal life, although they lack the active and agile habits 
of the Old World Gibbons. The tail with the under surface of its 
extremity naked affords the most completely prehensile type of 
this organ, and can sustain the weight of the whole body. 
Objects are not unfrequently grasped by it and brought within 
reach of the hand or mouth. Owing to the absence of the pollex 
the power of grasping is very imperfect in the hand. At least 
fourteen species of this genus have been described, among the 
best-known being 4. melanochir (Fig. 341), 4. paniscus of Guiana, 
A. geoffroyi of Central America, 4. afer of Eastern Peru, and 
al. hybridus of Colombia. 

Eriodes.2—Form slender ; limbs very long; fur woolly ; inter- 
nasal septum narrower than usual in the family; pollex rudimentary ; 
tail naked beneath distally ; nails exceedingly compressed laterally, 
and pointed. 

This genus is represented by three species from South-East 
Brazil, which, while closely allied to the true Spider Monkeys, 
differ by their woolly hair, the narrow internasal septum, the 
presence of a rudimentary pollex, and the great compression of the 


1 Geoffroy, Ann. du Muséum, vol. vii. p. 260 (1806). 
2 TI. Geoffroy, Dict. Class. vol. xv. p. 443 (1829). 


716 PRIMATES 


nails. The species are £. arachnoides, E. hemidactylus, and E. 
hypoxanthus. 

Lagothriz..—Form rather robust; limbs moderate ; fur woolly ; 
pollex well developed ; tail distally naked beneath. 

The Woolly Monkeys differ from the preceding genera by the 
presence of a well-developed pollex. They resemble Eriodes in 
their fur and compressed nails, but differ in the more widely 


Fic. 341.—The Black-handed Spider Monkey (Ateles melanochir). 
From Proc. Zool. Soc. 1871, pl. 15. 


separated nares. The tail resembles that of the preceding genera. 
Speaking of these Monkeys Mr. Bates observes that “the Barrigudos 
are very bulky animals, whilst the Spider Monkeys are remarkable 
for the slenderness of their bodies and limbs. I obtained specimens 
of what have been considered two species, one (L. olivaceus ?) 
having the head clothed with gray, the other (L. humboldti, Fig. 
342) with black fur. They both live together in the same places, 


1 Geoffroy, Ann. du Muséum, vol. xix. p. 106 (1812). 


CEBIDA 717 


and are probably only differently coloured individuals of one and 
the same species. I sent home a very large male of one of these 
kinds, which measured 27 inches in length of trunk, the tail being 
26 inches long ; it was the largest monkey I saw in America, with 
the exception of a black Howler, whose body was 28 inches in 
height. The skin of the face in the Barrigudo is black and 
wrinkled, the forehead is low, with the eyebrows projecting. . . . 
In the forests the Barrigudo is not a very active animal; it lives 
exclusively on fruits, and is much persecuted by the Indians on 


Fic. 342.—Humboldt’s Lagothrix (Lagothriz humboldti). From Proc. Zool. Soc. 1863, pl. 31. 


account of the excellence of its flesh as food.” Five species are 
usually recognised, viz. L. canus, L. humboldti, L. castelnam, L. 
ischudiit, and L. geoffroyt. 

Cebus.1\—Form rather robust ; limbs moderate ; fur not woolly ; 
pollex well developed ; tail not naked beneath distally. 

This, the typical, genus includes the Sapajous or Capuchins 
(Fig. 343), which are so commonly seen in this country in captivity, 
being the favourite Monkeys of itinerant musicians. They are 
smaller and stouter in build than the Spider Monkeys, from which 
they are readily distinguished by the well-developed pollex, and 
the absence of a naked under surface to the extremity of the tail. 


1 Erxleben, Syst. Regne Animal, p. 44 (1777). 


718 PRIMATES 


At least twenty species have been described (C. fatuellus, C. lunatus, 
C. capucinus, C. albifrons, C. hypoleucus, ete.), but it is probable that 
some of these are not entitled to stand, since there is a large 


Fic. 343,--The White-cheeked Sapajou (Cebus lwnatus). From Proc. Zool. Soc. 1865, pl. 45. 


amount of individual variation. Fossil remains of species of Cebus 
have been described from the Pleistocene cavern-deposits of Brazil. 


Family CERCOPITHECIDA, 


Dentition: ¢ 2, ¢ 4, p 2, m 8; total 32. Crowns of molars elon- 
gated antero-posteriorly, with the tubercles forming a pair of 
imperfect transverse ridges, and the last lower molar usually with 
a hind talon. A bony external auditory meatus. A narrow inter- 
narial septum. Tail non-prehensile. Ischiatic callosities present. 
Cheek-pouches present or absent. Pollex, when present, opposable. 
Pelvic limbs never much longer than pectoral. 
Czcum without vermiform appendage. 

This family includes all the Old World Apes, with the excep- 
tion of the Stmiide, and may be divided into the subfamilies Cerco- 
pithecinee and Semnopithecina. 

Subfamily Cereopithecinze.—Pelvic and pectoral limbs approxi- 


mately equal; tail variable; cheek-pouches present; stomach 
simple. 


Sternum narrow. 


CERCOPITHECIDE 719 


This subfamily comprises the African Baboons, the common 
Indian Monkeys constituting the genus Macacus, together with the 
African Cercopithecus and Cercocebus and a few allied types. 

‘Cynocephalus..—Muzzle much elongated (Fig. 344), with the 
nostrils terminal; ischial callosities very large; tail more or less 
short; muzzle swollen by enlargement of the maxille. Now con- 
fined to Africa and Arabia. 

This genus comprises the typical Baboons, and we may select 
the well-known Mandrill (C. maimon), of tropical West Africa, as a 
good illustrative example. It may be mentioned in passing that 
the name Mandrill appears to have been first introduced into 
English literature by William Smith in his New Voyage to Guinea, 


Fic. 344.—Skeleton of the Chacma Baboon (Cynocephalus porcarius). From De Blainville. 


published in 1744, wherein he mentions among the animals of 
Sierra Leone one “called by the white men in this country Man- 
drill,” but adds, “why it is so called I know not.”? Smith gives 
sufficiently accurate details to show that his animal is not that now 
called Mandrill, but the Chimpanzee. Buffon, however, while 
quoting Smith’s description, transferred the name to the very 
1 Lacépede, ‘‘ Nouv. tabl. méth.” (1799) in Mém. de ? Institut, vol. iii. p. 490 
1801). 
2 : ‘Mandrill’ seems to signify a ‘man-like Ape,’ the word ‘ Drill’ or ‘ Dril’ 
having been anciently employed in England to denote an Ape or Baboon. Thus 
in the fifth edition of Blount’s ‘ Glossographia, or a dictionary interpreting the 
hard words of whatsoever language now used in our refined English tongue . . . 
very useful for all such as desire to understand what they read,’ published in 
1681, I find ‘Dril, a stonecutter’s tool wherewith he bores little holes in marble, 
ete. Also a large overgrown Ape and Baboon, so called.’ ‘Drill’ is used in the 
game sense in Charlton’s Onomasticon Zoicon, 1668. The singular etymology 
of the word given by Buffon seems hardly a probable one.”—Huxley’s Man's 
Place in Nature, p. 10, 1863. 


720, PRIMATES 


different species now under consideration, and to that it has been 
attached ever since. 

The Baboons generally are distinguished from most other 
Monkeys by the comparative equality of the length of their limbs, 
which with the structure of the vertebral column adapts them 
rather for quadrupedal progression on the ground than for climbing 
among the branches of trees; and some of them, like the South 
African Chacma (C. porcarius), of which the skeleton is shown 
in Fig. 344, live habitually among rocks, and are much less 
completely frugivorous than other Apes. They are also remark- 
able for the great size of their face and jaws as compared with 
the part of the skull which encloses the brain. The Mandrill, 
in addition to these characters, is distinguished by the heaviness 
of its body, stoutness and strength of its limbs, and exceeding 
shortness of its tail, which is a mere stump, not 2 inches long, 
and usually carried erect. Jt is, moreover, remarkable for the 
prominence of its brow ridges, beneath which the small and 
closely approximated eyes are deeply sunk; the immense size of 
the canine teeth; the great development of a pair of oval bony 
prominences on the maxillary bones in front of the orbits, rising on 
each side of the median line of the face, and covered by a longi- 
tudinally ribbed naked skin; and more especially for the extra- 
ordinarily vivid colouring of some parts of the skin. The body 
generally is covered with a full soft coating of hair of a light olive- 
brown above and silvery-gray beneath, and the chin is furnished 
underneath with a small pointed yellow beard. The hair of the 
forehead and temples is directed upwards so as to meet in a point 
on the crown, which gives the head a triangular appearance. The 
ears are naked and of a bluish-black colour. The hands and feet 
are naked and black. A large space around the greatly developed 
ischial callosities, as well as the upper part of the insides of the 
thighs, are naked and of a crimson colour, shading off on the sides to 
lilac or blue, which, depending not upon pigment but upon injec- 
tion of the superficial blood-vessels, varies in intensity according to 
the condition of the animal—increasing under excitement, fading 
during sickness, and disappearing after death. But it is in the face 
that the most remarkable disposition of vivid hues occur, more 
resembling those of a brilliantly coloured flower than what might 
be expected in the cutaneous covering of a mammal. The cheek- 
prominences are of an intense blue, the effect of which is heightened 
by deeply sunk longitudinal furrows of a darker tint, while the 
central line and termination of the nose are a bright scarlet. Not- 
withstanding the beauty of these colours in themselves, the whole 
combination, with the form and expression of features, quite 
justifies Cuvier’s assertion that “il serait difficile de se figurer un 
-6tre plus hideux que le Mandrill.” 


CERCOPITHECIDA 721 


It is only to fully adult males that this description applies. 
The female is of much smaller size, and of more slender make; 
and, though the general tone of the hairy parts of the body is 
the same, the prominences, furrows, and colouring of the face are 
very much less marked. The young males have black faces. At 
the age of three the blue of the cheeks begins to appear, but it is 
not until they are about five, when they cut their great canine 
teeth, that they acquire the characteristic red of the end of the 
nose. 

The Mandrills, especially the old males, are remarkable for the 
ferocity of their disposition, as well as for other disagreeable quali- 
ties, which are fully described in Cuvier’s account of the animal in 


Fig. 345.—The Yellow Baboon (Cynocephalus babuin). From Archives du Muséum, 
vol. ii. pl. 34. 


La Ménagerie du Muséum d'Histoire Naturelle (1801), but when 
young they can easily be tamed. Like the rest of the Baboons, 
they appear to be rather indiscriminate eaters, feeding upon fruit, 
roots, reptiles, insects, scorpions, etc., and inhabit open rocky 
ground rather than forests. Not much is known of the Mandrill’s 
habits in the wild state, nor of the exact limits of its geographical 
distribution. The specimens brought to Europe all come from the 
west coast of tropical Africa, from Guinea to the Gaboon. 

An allied species, the Drill (C. lewcophieus), which resembles the 
Mandrill in size, general proportions, and shortness of tail, but 
wants the bright colouring of the face which makes that animal so 
remarkable, inhabits the same district. Other well-known species 
are the Yellow Baboon (C. babwin), of West Africa (Fig. 345); the 
Arabian Baboon (C. hamadryas), of Arabia and Abyssinia ; and the 
Anubis Baboon (C. anubis), of West Africa. 

46 


722 PRIMATES 


It is very noteworthy from a distributional point of view, as 
showing the former intimate connection between the faunas of the 
Oriental and Ethiopian regions, that fossil remains of Baboons have 
been found in the Pleistocene cavern-deposits of Madras, and also 
in the older Pliocene beds of the Siwalik Hills in Northern India ; 
the two species from the latter deposits having been described as 
C. subhimalayanus and C. faleoneri. 

Theropithecus.\—Distinguished from C'ynocephalus by the nostrils 
not being terminal, but situated as in J/acacus. This genus is 
represented by the Abyssinian Gelada (7. geluda) and the allied 
TL. obscurus. 

Cynopithecus.°—The Black Ape of Celebes (C. niger) forms a 


Fic. 346.—The Tibetan Macaque (Macacus tibetanus). Froin Milne-Edwards, Recherches des 
Mammiferes, pl. 34. 


connecting link between the Baboons and the genus Macacus ; the 
skull differing from that of the latter in the development of longi- 
tudinal ridges on the sides of the upper surface of the maxille, as 
in some of the species of Cynocephalus. The muzzle is also more 
produced than in Macacus. 

Macacns.’—Muzzle considerably produced ; nostrils not terminal; 
cheek-pouches and ischial callosities well developed ; tail long, short, 
or absent ; a distinct talon to the third lower molar. 

With the exception of the Barbary Ape (JL. inuus) of Northern 
Africa and Gibraltar, the Macaques are now exclusively Asiatic, 
one species (Fig. 346) occurring in Tibet, and another (JV. speciosus) 


1 T. Geoflroy, Arch. du Muséum, vol. ii. p. 576 (1841). 
* I. Geoffroy, Voyage de Belanger, p. 66 (1834), 
* Lacépede, Mem. de Institut, vol. iii. p, 450 (1801). Amended. 


CERCOPITHECID.E 723 


being found in Japan. All these Monkeys are of stout build, and 
it is chiefly by the greater production of the muzzle, the larger 
ischiatic callosities, and the frequent shortness of the tail that they 
are distinguished from the under-mentioned African genera. The 
transition from the longer-tailed to the short-tailed forms is so 
complete that the proposed generic separation of the latter as Inuus 
is impracticable. In AM. inwus the tail is wanting ; in JL tibetanus 
(Fig. 346) and MW. nemestrinus of Tenasserim it is short; in the 
common Bengal Monkey (J. rhesus) it is about one-half the length 
of the head and body, while in MZ. cynomolgus and its allies it is 
still longer. In the Indian Lion-tailed Monkey (Jf. silenus) it is 
tufted at the end. 

The following summary of the habits of the Macaques is taken 
from Mr. W. T. Blanford’s Mammals of British India: “The species 
of the present genus resemble each other in their habits; they are 
found in flocks, often of considerable size, and generally composed 
of individuals of both sexes and of all ages. They are active 
animals, though less rapid in their movements, whether on trees 
or on the ground, than the Semnopithect. Their food is varied, 
most of the species, if not all, eating insects as well as seeds, fruits, 
etc., and one kind feeding partly on crustacea. They have occa- 
sionally been known to devour lizards, and, it is. said, frogs also. 
All have the habit of cramming food into their cheek-pouches for 
mastication at leisure—a practice that must be familiar to any one 
who has fed monkeys in confinement. The voice and gestures of 
all the species are similar, and differ entirely from those of both 
the Gibbons and Semnopitheci. . . . The majority of the species are 
very docile when young. They thrive well, and several of them 
have bred in confinement. The period of gestation is about seven 
months, only a single young one, as a rule, being produced at a 
birth. They become adult at the age of four or five years, but 
breed earlier.” 

The Common Indian Jf. rhesus is found in the Himalaya at an 
elevation of over 8000 feet. 

Fossil remains of AMacacus are found in India in the Pleistocene 
of Madras and the Pliocene of the Punjab ; and they also occur in 
the Pliocene of France and Italy, those from the latter deposits 
having been incorrectly separated as Aulaxinwus. Part of the jaw 
of a Monkey from the Pleistocene of Essex has been described as 
Macacus pliocenus, and is very interesting as showing the presence 
of Apes in Europe at that late period. 

Cercocebus.\—An African genus agreeing with J/ucacus in the 
presence of a hind talon to the third lower molar, but with the 
other characters of Cercopithecus. The species of this genus are 
known as Mangabeys, or White-eyelid Monkeys, and include 

1 Geoffroy, dann. du Muséwin, vol. xix. p. 97 (1812), 


74 PRIMATES 


C. eollarvis, CL fuliginosus, C. aethiops, and C. allagena + all being 
from West Africa. 

Cercopithecus..—Muzzle more or less short ; ischial callosities 
moderate ; tail long; no talon to third lower molar. Build more 
slender than in Afweucus. Confined to Africa. 

The members of this and the last genus include those Monkeys 
which in their comparative slender build and length of tail make 
the nearest approach 
to the next subfamily. 
There are numerous 
species, among which 
the Green Monkey (C- 
callitrichus), the Grivet 
(CL qriseo- viridis), the 
Vervet (C. lalandi), the 
Pluto Monkey (C’ pluto, 
Fig. 347), the Patas 
(C. ruber), the Diana 
Monkey ((’. diana), and 
the Mona Monkey (C. 
mona) are well-known 
types. 

Subfamily Semno- 
pithecine. >? — Pelvic 
limbs longer than the 
pectoral; tail very 
long; no cheek- 
pouches; stomach sac- 
culated. Buildslender. 

This subfamily is 
represented by three 
genera, of which one is 


Fic, 347.—The Pluto Monkey (Cercopithecus pluto). From African and two are 
Gray, Proc. Zool. Soc. 1848, p. 57. Asiatic. Mr W. T. 


Blanford, in his J/am- 
mals of British India, observes that “the members of this subfamily 
are readily distinguished by their slender form, and by the absence 
of cheek-pouches. They are more purely herbivorous than the 
Macaque Monkeys, and a considerable portion of their food consists 
of leaves and young shoots. In consequence probably of the nature 
of their food, these Monkeys are more delicate than the species of 
Macacus, and are thus less easily kept in captivity. They are con- 
sequently far less well represented in European museums, and have 
been less studied by European naturalists. Very little is known of 
their general life-history or of their feeding habits.” 


1 Erxleben, Syst. Regne. Animal, p. 22 (1777). 2 Or Colobine. 


CERCOPITHECID.E 725 


Their digestive organs are much modified, the stomach attaining 
an extraordinary complexity, which may be described as follows. 
An ordinary stomach must be supposed to be immensely elongated, 
and gradually tapering from the 
cardiac end to a very prolonged, 
narrow, pyloric extremity. Then 
twolongitudinal muscular bands, 
corresponding in situation to the 
greater and lesser curvatures of 
anordinary stomach—the former 
commencing just below the fun- 
dus, and the latter at the cardiac 
orifice, and both proceeding \ 
towards the pylorus—are de- 
veloped, so as to pucker up the 
cavity into a number of pouches, 
exactly on the same principle as 
the human colon is puckered up 
by its three longitudinal bands. 
These pouches are largest and 
most strongly marked at the 
wesophageal end, and becoming 
less and less distinct, quite cease 
several inches before the pylorus 
is reached, the last part of the 
organ being a simple smooth- 
walled tube. The fundus, or 
cardiac end of the stomach, is 
formed by a single large sac, 
slightly constricted on its under 
surface by the prolongation of ; 
the inferior longitudinal band, Fia. 348.—Lateral view OF the skull and palatal 

‘ aspect of the cranium of Semnopithecus nemeus. 
or that corresponding to the (rom De Blainville.) 
great curvature. The cesophagus 
enters into the upper part of the left, or pyloric end of this sac, or 
rather at the point of junction between it and the second (also a 
very large) sacculus. Furthermore, the whole of this elongated 
sacculated organ is, by the brevity, as it were, of the lesser curva- 
ture, coiled upon itself in an irregularly spiral manner, so that 
when in situ the pylorus comes to be placed very near the esophageal 
entrance. 

Nasalis!1—Skull resembling that of the Cercopithecie in that 
the lower border of the nasal bones extends considerably below the 
lower border of the orbits, whereas in the other Semapitherinw the 
aperture of the nares extends upwards between the orbits, Nose 

1 Geoffroy, Ann. du Muséum, vol. xix. p. 90 (1812). 


726 PRIMATES 


produced into a large proboscis. Other characters as in Semmo- 
pithecus. 

This genus includes only the Proboscis Monkey (WN. larvatus) of 
Borneo, remarkable for the great prolongation of the nose in the 
adult. In young animals the nose is relatively much shorter, and 


Fic. 349.—Semnopithecus rovellane. (From Milne-Edwards, Recherches des 
Mammiferes, pl. 36.) 


bent upwards after the manner of that of Semnopithecus roxellane 
(Fig. 349). 

Semnopithecus.'—Pollex small; narial aperture extending up- 
wards between the orbits. Now confined to Asia. 

This genus is characteristic of South-Eastern Asia from the 
Himalaya southwards, the Oriental region being its head-quarters. 
The development of the muzzle is less than in the Macaques, and 
the facial angle is higher, but it does not appear that this indicates 
greater intellectual capacity. The outlying S. roaellane 2 (Fig. 349), 

1 F, Cuvier, His/, Nat. des Mammiferes (1821), * Semno-pithéque.” 
* Separated generically by some writers as Rhénopithecus. 


CERCOPITHECID.E viata 


of the highlands of Eastern Tibet and Kansu, is remarkable for the 
peculiar upturned nose, in which respect, as already mentioned, 
it recalls the young of Vasilis lurratus. The genus is represented 
in India and Burma by no less than fourteen species, of which the 
common Indian Langur, or Hanuman Monkey (8. ente//us) and 
the larger Himalayan Langur (8. schisfoceus) are two of the best 
known. In the former the length of the head and body is about 
24, and that of the tail 38 inches in adult males. This monkey, 
owing to the veneration in whieh it is held by the Hindus, is a 
great pest’ in many parts of India, frequently pilfering grain from 
the shops in the native bazaars. According to Mr. Blanford, 
it “is usually found in smaller or larger communities, composed 
of individuals of both sexes and of all ages, the youngest clinging 
to their mothers and being carried by them, especially when 
alarmed, An old male is occasionally found solitary, as with so 
many other mammals... . Apart from villages, the high trees 
on the banks of streams or of tanks, and, in parts of Central 
India, rocky hills are the favourite haunts of these monkeys. 
Whether on trees, on rocks, or on the ground, they are exceedingly 
active.” The closely allied 5. schisfaveus attains a larger average 
size, full grown males attaining a length of 30 inches, the tail 
measuring 36 inches. In the spring and winter this species may 
be observed in the Kashmir Himalaya leaping among the snow-laden 
trees of the forest. In a fossil state Semmnopitheens occurs in the 
Pleistocene and Pliocene of India, and it has also been recorded 
from the Pliocene of France and Italy. 

olohtts 1— This African genus differs from Scmnopithecus in that 
the pollex is absent or reduced to a small tubercle, which may or may 
not carry a nail, About eleven species have been described, some 
of which are remarkable for the beautitul mantle of long silky 
hair which hangs down from each side of the body, and for their 
tufted tails. In CL guerest from Abyssima these are white, and the 
rest of the body and limbs black. Others (as C. sufanas) ave entirely 
black. The skins of the long-haired species are largely imported 
into Europe for the manufacture of ladies’ muffs, ete. 

Extinct. Genere.—Certain types of Apes from the European 
Tertiavies indicate genera’ referable to the Cercopithecida, but 
distinet from any of those now living. Of these MWes«pithecus,? from 
the Lower Pliocene Pikermi beds of Attica, is known by almost 
complete skeletons, and resembles Jacacus in the shertness and 
stoutness of the limbs, but agrees with semnepificens in the characters 
of the skulland teeth. An allied Monkey from the Lower Pliocene 
of Perpignan, in France, differs from WVWesrjiitheens peitelict by its 
superior size, proportionately more produced muzzle, and larger 

1 Miger, Prodronivs Syst. Mai, et Avis. p. 69 (1811). 
2 Wagner, Gelehrte Aizeigen, vol. vili, No. 38. p. 310 (183%. 


728 PRIMATES 


hind talon to the last lower molar; it has been described under 
the name of Dolichopithecus. 

The genus Oreopithecus* was founded upon the remains of an 
Ape from the Middle Miocene of Monte Bamboli, in Tuscany, of 
somewhat larger size than a Gibbon, and apparently presenting 
characters connecting the Cercopithecide and Simiide. According 
to Dr. Ristori,’ it resembles the former, especially Cynocephalus and 
Semnopithecus, in the long dental series and the elongation of the 
last molars ; but in the shortness of the face, rounding of the chin, 
and the diagonal arrangement of the molar tubercles, it approximates 
to the Similw, of which it may have been an ancestral type. 


Fonily SIMuUDz. 


* Crowns of molars relatively wide, with the angles more or 
less rounded off, the tubercles not forming tranverse ridges, and 
the last lower molar without a hind talon. No tail. No cheek- 
pouches. Ischiatic callosities, if present, small. Pectoral limbs 
much longer than pelvic. Sternum broad. Ccum with vermiform 
appendage. Centrale of carpus sometimes absent. Other characters 
as in Cercopithecida, 

This family contains the true Anthropoid Old World Apes, 
namely the Gibbons, Orangs, Chimpanzees, and Gorillas, which 
are the most highly organised of all the Apes, and thus make the 
nearest approach to Man. 

ITylobates.A—Skull not produced at the vertex ; body and limbs 
slender, the pectoral limbs being so elongated that the hands reach 
the ground when walking upright ; hallux well developed ; a centrale 
in the carpus; and small ischiatic callosities. Size smaller than in 
the following genera, the height of the largest species (JZ. syndactylus) 
not much exceeding 3 feet. Now confined to Asia. 

The Gibbons, or Long-armed Apes (Figs. 350, 351), are readily 
distinguished from the remaining members of the family by the 
characters given above, as well as by the circumstance that they 
are the only Apes which habitually walk in an upright position. 
It is in these animals that we meet with the last traces of the 
ischial callosities so largely developed in the Cercopithccide. The 
species are now restricted to South-Eastern Asia, being especially 
abundant in the Malay Archipelago and adjacent regions. 

The largest species is the Sumatran Niamang (11. syiductylus), 
which attains a height of 3 feet, and has been generically 
separated by some writers as Sivmenga. It is remarkable as 

1 Depéret, Comptes Rendus, vol. cix, p. 982 (1889); see also Afén. Soc. Géol. 
France, ‘ Paleontologie,” vol. i. (1890). * Gervais, Comptes Rendus, vol. 
Ixxiv. p. 1217 (1872). * Scimmie Fossili Italiane, Bol’. Conum. Geol. 1890. 

4 Illiger, Prodromus Syst. Maman. ct Avium, p. 67 (1811), 


SIMIMD.E 729 


having a better developed chin and wider sternum than any other 
Ape, and differs from the other members of the genus by the 
circumstance that the second and third digits of the pes are united 
by skin as far as 
their last joints. 
Exclusive of this 
species, the Gibbons 
differ but little from 
“one another in size 
and general con- 
formation, and since 
the colour of indi- 
viduals undoubtedly 
referable to a single 
species is remarkably 
variable, there is 
much uncertainty 
about the number of 
species, and much 
confusion in the 
nomenclature. 
Among well-marked 
species we may 
mention the Hoolock 
(H. hoolock), ranging 
from the South of 
Assam through 
Sylhet and Cachar to 
the Ivawadi Valley 
near Bhamo, the 
White-handed Gib- 
bon (H. lar, Fig. 


350), which is found 


in Tenasserim and Fic. 350.—The White-handed Gibbon (Hylobates lar). From 
Blanford, Mammals of British India, p. 8. 


throughout Malay- 
ana, the Dun-coloured Gibbon (H. entelloides, Fig. 351) of Malayana, 
and the Tufted Gibbon (H. pileatus) of Siam and Cambogia. 

The following account of the habits of the Gibbons is taken 
from Mr. W. T. Blanford’s Mammals of British India. “Gibbons 
are thoroughly arboreal, and Hoolocks are almost, if not entirely, 
confined to hill-forest. They move chiefly by means of their long 
arms, by which they swing themselves for prodigious distances from 
branch to branch and from tree to tree. They descend hillsides 
at a surprising pace, their descent being accomplished by grasping 
tbamboos or branches that bend beneath their weight, and allow 
them to drop until they can seize the ends of other bamboos or 


730 PRIMATES 


branches lower on the slope, and take another mighty swing down- 
wards. They also ascend with great rapidity, swinging themselves 
from tree to tree. When walking on the ground the Hoolock rests 
on its hind feet alone, with the sole flat on the ground, and the 
great toe widely separated from the other digits. The arms are 
usually held upwards, sometimes horizontally, their great length 
giving the animal a very peculiar aspect. Gibbons walk rather 


Fic. 351.—The Dun-coloured Gibbon (Hyloletes exteloides). From Archiv. dy Mustum, 
vol. ii. pl. 29. 


quickly, with a waddling gait, and can easily be overtaken by men 
when on the ground. The food of these Apes consists of fruit, 
leaves, young shoots, spiders (of which they are very fond), insects, 
birds’ eges, and almost certainly of young birds, if not of any birds 
they can capture. Anderson found that small birds were killed 
and devoured by Hoolocks in confinement with a method and 
eagerness that showed this prey to be the natural food of the Apes. 
The Hoolock drinks with its lips, putting its head down to the 
water as Monkeys do. All species of Hylobates have a powerful 


SIMIIDA 731 


voice, and the common name of the Hoolock is taken from its 
peculiar double call, which is repeated several times. At a distance 
the sound much resembles a human voice; it is a peculiar wailing 
note, audible from afar, and in the countries inhabited by these 
animals is one of the most familiar forest sounds. The calls com- 
mence at daybreak, and are continued till 9 or 10 A.M, several of 
the flock joining in the cry, like hounds giving tongue. After 9 or 
10 o'clock in the morning the animals feed or rest, and remain 
silent throughout the middle of the day, but recommence calling 
towards evening, though to a less extent than in the earlier part of 
the day.” 

The skull of the Gibbons, although agreeing with that of other 
Apes in its prognathism, presents a somewhat human appearance, 
and the molar teeth are also very like diminutive human molars. 
In the anterior inward inclination of the two series of cheek-teeth 
and the inward 
position of the 
upper premolars 
the Gibbons make 
an approach to the 
human type un- 
known in other 
Apes. 

The figure of 
the liver of one 
species of this 
genus is introduced 


to show the general Fic. 352.—Under surtace of the liver of Hylobates lar. u, Umbili- 
absence of lateral cal fissure; p, portal fissure; ve, vena cava; 1, left lobe; 7, right 
i lobe; s, Spigelian lobe; ¢, caudate lobe; g, gall-bladder. 


fissures and the 
small size of the caudate lobe (c) characteristic of the liver of all the 
Simiide, except Gorillu (see p. 706), as well as that of Man. Another 
specimen of the liver of the same species showed scarcely any trace 
of a caudate lobe. 

A fossil Ape from the Middle Miocene of France, originally 
described as Pliopithecus, indicates an extinct Gibbon which does 
not appear to be generically separable from Hylobates. 

Simia1—Skull (Fig. 353) produced at the vertex; body and 
limbs massive ; the pectoral limbs reaching to the ankle ; a centrale 
in the carpus ; hallux very small ; sixteen dorso-lumbar vertebrae, and 
twelve pairs of ribs; no ischiatic callosities. Oriental. 

This genus includes the large red-haired Apes from Sumatra 
and Borneo commonly known as Orangs, or Orang-Utans,” of which 
there is probably only a single species (S. satyrus). These animals 

1 Linn. Syst. Nat. 12th ed. vol. i. p. 34 (1766). 
2 A Malay word, signifying “ Man of the Woods.” 


732 PRIMATES 


inhabit the swampy forests near the coasts; and the males attain a 
height of about 4 feet 4 inches. The body is very bulky and the 
legs exceedingly short, but the arms are very long, reaching in the 
erect posture down to the 
ankles. The Orang walks 
resting on the knuckles of 
the hands and the outer 
sides of the feet, with the 
soles of the latter turned 
mainly inwards, as in Fig. 
354. Its movements 
appear to be slow and 
deliberate, and in those 
specimens which have been 
kept in captivity in this 
country the demeanour is 
languid and melancholy, 
although this is far from 
being the case with those 
shown in the more congenial 
climate of the Zoological 
Fig. 353.—Side view of the skull of adult Orang (Simia Gard srs at Calcutta. The 
satyrus). From Trans. Zool. Soc. vol. i. pl. 53. habits of these animals are 
arboreal, and they build a 

kind of shelter or nest of boughs and leaves; their food appears 
to consist mainly of fruits, and is exclusively of a vegetable nature. 
The whole of the body is clothed with long hair of a reddish-brown 
colour, and full-grown males have a well-developed beard; the 
males not unfrequently also develop a large warty protuberance, 
formed of fibro-cellular tissue, on either side of the face. The 
hands are long, and are characterised by the small size of the 
pollex, which does not reach to the end of the metacarpal of 
the index finger. The feet have a similar structure, the hallux 
only reaching to the middle of the proximal phalange of the 
adjacent toe, and being often destitute not only of a nail, but 
likewise of the terminal phalange. The presence of a centrale in 
the carpus is a feature in which Simia agrees with Hylobates and 
the lower Apes, and differs from the two following genera and Man. 
With very rare exceptions the number of dorso-lumbar vertebra is 
sixteen, of which twelve carry ribs, and therefore belong to the 
dorsal series, while the remaining four are lumbar. The distinction 
between the last lumbar and the first sacral vertebrae is clearly 
marked in young skeletons by the additional pleurapophysial 
ossifications (sacral ribs) in the transverse processes of the latter. 
Thus though Simia presents a closer resemblance to Man than does 
Anthropopithecus in the number of ribs, it differs in the more 


SIMUDE 733 


important characters of that of the whole series of trunk-vertebre.? 
The hemispheres of the brain are much convoluted; the whole 
brain being more human-like than in any other Ape. The larynx is 
remarkable for having a prolongation from each ventricle, which in 
the adult become of enormous dimensions, and unite in front of 


Fig. 354.—The Orang-Utan (Simia satyrus). From Mr. Wolf's sketch at the 
Zoological Gardens. 


the trachea to form one large sac extending downwards between 
the muscles to the axilla. 

The skull of the Orang (Fig. 353) is characterised by its highly 
vaulted cranial portion, which is comparatively short (brachy- 
cephalic). The sagittal crest is well developed on the vertex, and 
has a highly convex contour; the superciliary ridges are but 
moderately developed, and do not stand out in the prominent 
manner so characteristic of the Gorilla. The aperture of the nares 
in the skull is more pear-shaped than in the two following genera. 

The canines of the male Orang attain a great development: 


1 One skeleton in the Museum of the Royal College of Surgeons has five 
lumbar vertebre, and has thus given rise to the statement that the number of 
vertebre in the Orang is the same as in Man. 


734 PRIMATES 


and the molars are characterised by the complex structure of théir 
cusps and the numerous rugosities on the crown surface. The 
outer border of the upper premolars is placed in the same line as 
that of the molars. 
The broken canine tooth of a large Anthropoid Ape from the 
= Lower Pliocene of the Siwalik Hills 
probably indicates the existence at 
that period of a species of Simia in 
Northern India. 

Gorilla..—Skull not produced at the 
vertex ; body and limbs massive, the 
pectoral limb not reaching below the 
middle of the lower leg (Fig. 355) ; 
no centrale in the carpus; hallux well 
developed ; seventeen dorso-lumbar 
vertebrae, of which thirteen carry ribs ; 
‘jf no ischiatic callosities. Male much 
| larger than female, and with very 
strongly marked cranial ridges, which 
| are wanting in the latter. Mandibular 
symphysis long. Ethiopian. 

The well-known Gorilla (Fig. 356), 
of which there seems to be only one 
species (G. savagei), is found in Western 
Equatorial Africa, chiefly or entirely 
in the district enclosed by the 
Cameroon and Congo rivers. It is 
the largest of all the Apes, its bulk 
considerably exceeding that of man, 
although from the shortness of the 
legs it appears never to attain a greater 

; height than 51 feet. The first intro- 

Fie. 355.—Skeleton of the Gorilla. duction of this animal to the notice 
iii Di cuiieaec: of zoologists was made in 1847 by 

Dr. Thomas Savage, but it was not fully known till many years later. 
The skin of the Gorilla is entirely black, the hair being blackish, 
but turning more or less gray in old individuals. The arms reach 
down as far as the middle of the lower leg; while the pollex 
extends only a short distance beyond the base of the first phalange 
of the index finger, and the hallux reaches nearly as far as the 
distal extremity of the corresponding digit of the foot. The digits 
of both the hand and foot are united together by integument as 
far as the distal extremities of the first phalanges. The larynx 
has very capacious air-sacs, which meet in front of the trachea and 
communicate with the ventricles; and in advanced age these sacs 


' I. Geoffroy, Comptes Rendus, vol. xxiv. p. 84 (1852), 


SLM DA 735 


may extend to the axilla. The ears are relatively small. The 
skull is of an elongated or dolichocephalic type; that of the adult 
male being characterised by the enormous development of the 
supraorbital ridges, which form a kind of penthouse over the eyes, 
and contribute to the peculiarly ferocious appearance of the animal. 
The sagittal crest is also very large. The canine teeth of the male 
are very large, and are inclined outwards in both jaws. In the 


Fic. 356.—The Gorilla (Gorilla savagei). From Trans. Zool. Soc. vol. iv. pl. 43. 


cheek-teeth the upper premolars are of considerable antero-posterior 
extent, with their outer border placed in the same line as that of 
the molars; and the third upper molar is larger than either of the 
others. : 

The posterior cervical vertebre are characterised by the great 
height of their neural spines, which thus form a strong basis for 
the powerful cervical muscles supporting the massive skull. In 
some instances the fourth lumbar vertebra becomes ankylosed to 
the sacrum, as is occasionally found to be the case in some of the 
lower human races. 

In the absence of a centrale to the carpus, and also in the 


736 PRIMATES 


number of the dorso-lumbar vertebrxe, the present and following 
genus resemble man; although they both differ in having thirteen 
in place of twelve pairs of ribs. 

The brain of the Gorilla, according to Dr. Hartmann, resembles 
that of the Orang in the complexity of its convolutions, and is 
thereby distinguished from that of the Chimpanzee. In form it is 
of the long oval characteristic of Man; the brain of the Chimpanzee 
and Orang being more rounded. 

Gorillas live in family parties in the depths of the dense forests 
of Western Equatorial Africa, seeking their food during the day, 
while at night it is said that the female and young ascend a tree 
at the foot of which the male sleeps. They walk with the backs 
of their closed hands and the flat soles of the feet placed on the 
ground. Although there has been much exaggeration on this 
point, it appears certain that the male Gorilla is an extremely 
ferocious and dangerous animal when brought to bay, but the 
statements as to its making unprovoked assaults on men do not 
appear authentic. They utter deep guttural sounds, which on 
some occasions may be described as grunts and at others as a 
roar. 

Anthropopithecus1—One of the most important differences of 
this genus from the preceding is the absence of any marked 
disparity between the two sexes, either in the size or the con- 
formation of the skull, although the male can always be dis- 
tinguished by the larger size of the canine teeth. The mandibular 
symphysis is also much shorter. Differences in the characters of 
the teeth are described below. The genus is confined at the present 
day to the Ethiopian region. 

The Chimpanzees (Fig. 357) inhabit Western and Central 
Equatorial Africa; and there has been much discussion whether 
they should all be included under one specific name (d. troglodytes), or 
whether there are really two or more species. A female specimen 
now living in the London Zoological Gardens, characterised among 
other distinctive features by the nearly bald head, clearly indicates, 
however, a second species, which probably corresponds to the 
imperfectly defined .f. calrus of Du Chaillu. 

The region inhabited by the Chimpanzees extends from the 
Gambia to the Benguela, reaching as far inland as 28° E. long. 
The Common Chimpanzee is a smaller animal than the Gorilla, its 
height not exceeding 5 feet. In colour it is darker than the 
latter, and the ears are relatively larger. In the upright position 


1 De Blainville, Lecons Orules (1839). The Chimpanzees have been very 
generally described under the name of Zroglodytes, but since this name is 
preoccupied for a genus of birds, it is incumbent to follow the strict rule, and 
adopt the name Anthropopitheens, although both the present witers have 
elsewhere expressed the opposite opinion. 


SIMIIDAE 737 


the arms reach only a short distance below the knee, in which 
respect the Chimpanzee is more human-like than any of the other 
Apes. The face is furnished with distinct whiskers, eyebrows, and 
eyelashes. The pollex reaches nearly or quite to the base of the 
first phalange of the index finger, and the hallux to the base of 
the second phalange of the corresponding digit of the foot. The 
laryngeal sacs are as largely developed as in the Gorilla. 

Although the skull of the Chimpanzee has distinct superciliary 


/ 


/ 


By, 
j 


\ \ 
‘ 


\’ 


Fia. 357.—The Chimpanzee (Anthropopithecus troglodytes). From Mr. Wolf's drawing of a young 
individual in the Zoological Society’s Gardens. 


‘ridges, yet the high bony crests of the calvarium of the male 
Gorilla are wanting, and the whole coronal region of the skull is 
more rounded and far less rugged. 

The canine teeth of the male Chimpanzee are relatively much 
smaller than in the Gorilla and Orang. The upper molars are 
characterised by the third one being smaller than either of the 
other two, as well as by the presence of an indistinct cingulum on 
their inner surfaces. The upper premolars differ from those of the 
other genera of the family by the shortness of their antero-posterior 

47 


738 PRIMATES 


diameter, and also by the larger size of their external as compared 
with their internal cusps; while the outer border of these teeth is 
placed internally to that of the upper molars. In all these respects 
the teeth of the Chimpanzee make a decided approximation to the 
human type. 

Many young individuals of the Chimpanzee have been brought 
to Europe, but they appear to succumb sooner or later to the effects 
of an unsuitable climate. All these examples show that the dis- 
position of this Ape is gentle, lively, and intelligent, and in all 
respects markedly opposite to that of the Orang. In a wild state 
these Apes are essentially forest-dwellers, and are more arboreal in 
their habits than the Gorilla. They live either in families, or in 
small parties of several families. Frequently at least they construct 
a kind of nest in the trees as a sleeping-place ; the male being said 
to sleep on a forked branch below the level of this nest. In walk- 
ing the Chimpanzee usually supports himself on the backs of his 
closed fingers, and either on the soles of the feet or on the closed 
toes. 

From a distributional point of view the discovery of a fossil 
Ape in the Pliocene of the Punjab, apparently closely allied to the 
Chimpanzee, is of great interest. This determination rests upon 
the evidence of an imperfect palate originally described under the 
name of Palwopithecus, but subsequently referred to the present 
genus. The teeth of this jaw present all the essential characters 
of those of the Chimpanzee, but the two series of cheek-teeth have 
a slight anterior convergence, the premolars are shorter in the 
antero-posterior direction than is usually the case in that species, 
and the outer incisor is relatively narrower than in the latter. In 
these features the extinct 4. sivalensis makes a nearer approxima- 
tion to the human type than is the case with its living congeners. 

Dryopithecus.'—The extinct Dryopithecus of the Middle Miocene 
of France is represented by a single species of the approximate 
size of the Chimpanzee, and appears to be the most generalised 
member of the family. According to the recent observations of 
Professor Gaudry,? while it resembles the Gorilla in that the two 
series of lower cheek-teeth diverge anteriorly and the penultimate 
premolar is larger than the last of that series, it differs in having a 
much longer and narrower mandibular symphysis, and thus indicates 
a transition to the Cercopithecide. A gradual transition in the form 
of the mandible may, indeed, be traced from Dryopithecus, through 
Gorilla, to Anthropopithecus ; the latter having a short and wide 
symphysis, with the two series of cheek-teeth slightly converging 
anteriorly, and the penultimate premolar being not larger than the 
last. In all these specialised characters the jaw of the Chimpanzee 

1 Lartet, Comptes Rendus, vol. xliii. p. 219 (1856). 
2 Mém. Soc. Géol. France, ‘‘ Paleontologie,” vol. i. Mém. No. 1 (1890). 


HOMINIDE . 739 


approximates to that of Man, in which the symphysis is still further 
shortened and widened, and the anterior convergence of the cheek- 
teeth so much increased as to produce a horse-shoe-like form in the 
whole dental series. 


Family HoMInip&. 


In the Systema Nature of Linneus Man was separated only 
generically from the Apes, but in the next great work which exer- 
cised a wide-spread influence over the progress of zoological science, 
the Regne Animal of Cuvier, he forms a distinct order under the 
name of Bimana, the Monkeys and Lemurs being associated together 
as Quadrumana. This has been the prevailing arrangement in the 
zoological systems of the present century, though in the classifica- 
tion of Owen lis position is still farther removed from that of the 
Monkeys, as in it the genus Homo forms one of the four primary 
divisions or subclasses of the Mammalia, called Archencephala, the 
Quadrumana being united with the Carnivora, Ungulata, and others 
in another division called Gyrencephala. On the other hand, the 
tendency of most modern systematists, for reasons which have been 
fully stated by Professor Huxley,! is to revert towards the Linnzan 
position. 

Considering solely the facts of Man’s bodily structure, it can be 
clearly demonstrated that the points in which he differs from the 
Ape most nearly resembling him are not of greater importance than 
_ those by which that Ape differs from other universally acknowledged 
members of the group; and therefore, in any natural system, if 
Man is to be made a subject of zoological classification upon the 
same principles as those applied elsewhere, he must be included in 
the order which comprises the Monkeys. We say upon the same 
principles as are applied elsewhere, since zoological classification has 
never taken into consideration the psychological characteristics 
which distinguish the subjects of its investigations, but only their 
tangible and physical structure, otherwise endless confusion would 
result, at all events with our very imperfect knowledge of animal 
psychology. The essential attributes which distinguish Man, and 
give him a perfectly isolated position among living creatures, are 
not to be found in his bodily structure, and should therefore either 
be left entirely out of consideration, or have such weight given to 
them as would remove him completely out of the region of zoological 
classification. To profess to classify Man as if he were one of the 
animals (as in all points of the structure and functions of his organs 
he undoubtedly is), to place him in the class Mammalia, and then 


1 Man's Place in Nature, 1863, and Anatomy of Vertebrated Animals, 1871. 
See also the more recent investigations of Broca into the comparative structure 
of Man and the higher Apes, published mostly in the Revue @ Anthropologie. 


740 PRIMATES 


to allow other considerations to influence the judgment as to the 
particular position he should occupy in the class, is most illogical. 

Man, therefore, considered from a zoological point of view, must 
be included in the order Primates, even if the Lemurs be removed 
from it, since his structural affinities with the Monkeys are far 
closer than are those of the so-called “‘Half-Apes.” We may, without 
treading upon debatable ground, go farther, and say that the 
differences between Man and the Anthropoid Apes are really not 
so marked as those which separate the latter from the American 
Monkeys. This being admitted, perhaps the best exposition relating 
to the present condition of the order will be to regard Man as 
representing a fifth family of the Anthropoidea, which should be 
known as the Hominide. In thus ranking Man as one of the five 
principal families or sections of the suborder it should, however, be 
observed that this course does not in the least degree imply that 
such families are precisely equivalent to one another, or that the 
intervals by which they are separated are of equal importance ; all 
that we commit ourselves to being that they are five perfectly 
distinct groups, all branches from a common stem, and in the 
present state of nature not united by any intermediate types. 

The distinctions between the Hominide and Simiide are chiefly 
relative, being greater size of brain and of brain-case as compared 
with the facial portion of the skull, smaller development of the 
canine teeth of the males, complete adaptation of the structure of the 
vertebral column to the vertical position, greater length of the lower 
as compared with the upper extremities, and greater length of the 
hallux or great toe, with almost complete absence of the power of 
bringing it in opposition to the other four toes. The last feature 
together with the small size of the canine teeth are perhaps the 
most marked and easily defined distinctions that can be drawn 
between the two groups. 

Man is universally admitted to form a single genus, Homo of 
Linnzus, but a question of considerable importance in treating of 
him from a zoological point of view, and one which has been a sub- 
ject of much controversy, is whether all men should be considered 
as belonging to a single or to several species. This question is 
perhaps of less importance now than formerly, when those who 
maintained a plurality of species associated with the hypothesis 
plurality of origin. One of the strongest arguments against the 
view that the various races of Man represent more than one species 
is that none of those who have maintained it have been able to 
agree as to how many distinct specific modifications can be defined, 
almost every number from three to twenty or more having been 
advocated by different authors. If the distinguishing characters of 
the so-called species had been so marked, there could not be such a 
remarkable diversity of opinion upon them. Again, the two facts 


HOMINID: 741 


—(1) that, however different the extremes of any two races may 
be in appearance (and it must be admitted that, as advocated by 
many polygenists, the differences are greater than many which are 
considered specific among other animals), every intermediate grada- 
tion can be found through which the one passes into the other, 
and (2) that all races are fertile inter se—are quite conclusive in 
favour of considering Man as representing a single species in the 
ordinary sense in which the word is now used, and of treating of 
all his various modifications as varieties or races. 

The great problem at the root of all zoology, the discovery of a 
natural classification which shall be an expression of our knowledge 
of the real relationship or consanguinity of different forms, is also 
applicable to the study of the races of Man. When we can satis- 
factorily prove that any two of the known groups of mankind are 
descended from the same common stock, a point is gained. The 
more such points we have acquired the more nearly shall we be 
able to picture to ourselves, not only the present, but also the past 
distribution of the races of Man upon the earth, and the mode and 
order in which they have been derived from one another. But the 
difficulties in the way of applying zoological principles to the classi- 
fication of Man are vastly greater than in the case of most animals. 
When groups of animals become so far differentiated from each 
other as to represent separate species, they remain isolated; they 
may ,break up into further subdivisions—in fact, it is only by 
further subdivision that new species can be formed; but it is of 
the very essence of species, as now universally understood by 
naturalists, that they cannot recombine, and so give rise to new 
forms. With the varieties of Man it is otherwise. They have 
never so far separated as to answer to the physiological definition 
of species. All races, as said above, are fertile with one another, 
though perhaps in different degrees. Hence new varieties have 

_ constantly been formed, not only by the segmentation of portions 
of one of the old stocks, but also by various combinations of those 
already established. 

Without entering into the difficult question of the method of 
Man’s first appearance upon the world, we must assume for it vast 
antiquity,—at all events as measured by any historical standard. 
Of this there is now ample proof. During the long time Man 
existed in a savage state—a time compared to which the dawn of 
our historical period is as yesterday—he was influenced by the 
operation of those natural laws which have produced the variations 
seen in other regions of organic nature. The first Men may very 
probably have been all alike ; but when spread over the face of 
the earth and subjected to all kinds of diverse external conditions, 
—climate, food, competition with members of their own species or 
with wild animals,—racial differences began slowly to be developed 


742 PRIMATES 


through the potency of various kinds of selection acting upon the 
slight variations which appeared in individuals in obedience to the 
tendency planted in all living things. These differences manifested 
themselves externally in the colour of the skin, the colour, quality, 
and distribution of the hair, the form of the head and features, and 
the proportions of the limbs, as well as in the general stature. 

Geographical position must have been one of the main elements 
in determining the formation and permanence of races. Groups of 
Men isolated from their fellows for long periods, such as those 
living on small islands, to which their ancestors may have been 
accidentally drifted, would naturally, in course of time, develop a 
new type of features, of skull, of complexion, or hair. A slight set 
in one direction in any of these characters would constantly tend 
to intensify itself, and so new races would be formed. In the same 
way different intellectual or moral qualities would be gradually 
developed or transmitted in different groups of Men. The longer 
a race thus formed remained isolated the more strongly impressed 
and the more permanent would its characteristics become, and less 
liable to be changed or lost when the surrounding circumstances 
were altered or under a moderate amount of intermixture from 
other races—the more “true,” in fact, would it be. On the other 
hand, on large continental tracts, where no mountain ranges or 
other natural barriers form obstacles to free intercourse between 
tribe and tribe, there would always be a tendency towards uni- 
formity, from the amalgamation of races brought into close relation 
by war or by commerce. Smaller or feebler races would be 
destroyed or absorbed by others impelled by superabundant popu- 
lation or other causes to spread beyond their original limits; or 
sometimes the conquering race would itself disappear by absorption 
into the conquered. 

Thus for untold ages the history of Man has presented a shift 
ing kaleidoscopic scene: new races gradually becoming differenti- 
ated out of the old elements, and, after dwelling a while upon the 
earth, becoming either suddenly annihilated or gradually merged 
into new combinations ; a constant destruction and reconstruction ; 
a constant tendency to separation and differentiation, and a tendency 
to combine again into a common uniformity—the two tendencies 
acting against and modifying each other. The history of these 
processes in former times, except in so far as they may be inferred 
from the present state of things, is a difficult study, owing to the 
scarcity of evidence. If we had any approach to a complete 
paleontological record, the history of Man could be reconstructed ; 
but nothing of the kind is forthcoming. Evidence of the anatomi- 
cal characters of Man as he lived on the earth during the time 
when the most striking racial characteristics were being developed, 
during the long ante-historic period in which the Negro, the Mon- 


HOMINID 743 


golian, and the Caucasian were being gradually fashioned into their 
respective types, is entirely wanting, or if any exists it is at present 
safely buried in the earth, perhaps to be revealed at some unex- 
pected time and in some unforeseen manner. Even the materials 
from which a history of the modifications of the human species as 
known to our generation must be constructed are rapidly passing 
away, since the age in which we live is an age in which, in a far 
greater degree than any previous one, the destruction of races, both 
by annihilation and absorption, is going on. Owing to the rapid 
extension of maritime discovery and commerce, changes such as 
have never been witnessed before are now taking place in the 
ethnology of the world—changes especially affecting the island 
populations among which, more than elsewhere, the solution of 
many of these problems may be looked for. The subject is, how- 
ever, attracting the attention of observers of all countries to a 
greater degree than it ever has before, and such progress has been 
made in perfecting the methods of investigation of racial character- 
istics that we are beginning to learn what lines of research are 
profitable and what are barren, so that we may hope the time is 
not far distant when we may get some clear insight into the know- 
ledge of the natural classification and relationships of the races of 
Man. 

The following is a brief summary of the principal results 
which appear to have been attained up to the present time by the 
study of this somewhat difficult subject.’ 

The most ordinary observation is sufficient to demonstrate the 
fact that certain groups of men are strongly marked from others by 
definite characters common to all members of the group, and trans- 
mitted regularly to their descendants by the laws of inheritance. 
Thus the Chinaman and the Negro, the native of Patagonia and the 
Andaman Islander, are as structurally distinct from each other as 
are many of the so-called species of any natural group of animals. 
Indeed, it may be said with truth that their differences are even 
greater than those which mark the groups called genera by many 
naturalists of the present day. Nevertheless the difficulty of 
parcelling out all the individuals composing the human species into 
certain definite groups, and of saying of each man that he belongs 
to one or other of such groups, is insuperable. No such classifica- 
tion has ever been, or, indeed, can ever be obtained. There is not 
one of the most characteristic and most extreme forms, like those 
just named, from which transitions cannot be traced by almost 
imperceptible gradations to any of the other equally characteristic 
and equally extreme forms. Indeed, a large proportion of mankind 

1 “On the Classification of the Varieties of the Human Species,” by W. H. 
Flower, Journal of the Anthropological Institute of Great Britain and Ireland, 
May 1885. 


744 PRIMATES 


is made up, not of extreme or typical, but of more or less general- 
ised or intermediate forms, the relative numbers of which are 
continually increasing, as the long-existing isolation of nations and 
races breaks down under the ever-extending intercommunication 
characteristic of the period in which we live. 

The difficulties of framing a natural classification of Man, or 
one really representing the relationship of the various minor groups 
to each other, are well exemplified by a study of the numerous 
attempts which have been made from the time of Linneus and 
Blumenbach onwards. Even in the first step of establishing certain 
primary groups of equivalent rank there has been no accord. Thus 
four primitive types were sketched out by Linnzus—the European, 
Asiatic, African, and American. These were expanded into five 
by Blumenbach by the addition of the Malay,’ and reduced by 
Cuvier to three by the suppression of the last two. Many later 
writers have largely increased the number of these so-called primary 
divisions, but the conclusion, so often arrived at by various anthro- 
pologists, and so often abandoned for some more complex system, 
that the primitive man, whatever he may have been, has in the 
course of ages divaricated into three extreme types, represented by 
the Caucasian of Europe, the Mongolian of Asia, and the Ethiopian 
of Africa, and that all existing individuals of the species can be 
ranged around these types, or somewhere or other between them, 
seems, on the whole, to give the clearest view of the facts of the 
case. Large numbers are doubtless the descendants of direct 
crosses in varying proportions between well-established extreme 
forms; for, notwithstanding opposite views formerly held by some 
authors on this subject, there is now abundant evidence of the 
wholesale production of new races in this way. Others may be 
the descendants of the primitive stock before the strongly marked 
existing distinctions had taken place, and therefore present, though 
from a different cause from the last, equally generalised characters. 
In these cases it can only be by most carefully examining and 
balancing all characters, however minute, and finding out in what 
direction the preponderance lies, that a place can be assigned to 
them. It cannot be too often insisted on that the various groups 
of mankind, owing to their probable unity of origin, the great 
variability of individuals, and the possibility of all degrees of 
intermixture of races at remote or recent periods of the history of 
the species, have so much in common that it is extremely difficult 
to find distinctive characters capable of strict definition by which 
they may be differentiated. It is more by the preponderance of 
certain characters in a large number of members of a group, than 
by the exclusive or even constant possession of these characters 


1 The Malay of Blumenbach was a strange conglomeration of the then little 
known Australian, Papuan, and true Malay types. 


HOMINID 745 


in each of its members, that the group as a whole must be 
characterised. : 

Bearing these principles in mind, we may endeavour to formu- 
late, as far as they have as yet been worked out, the distinctive 
features of the typical members of each of the three great divisions, 
and then show into what subordinate groups each of them seems to 
be divided. 

We begin with the Ethiopian, Negroid, or Melanian, or “ black ” 
type. It is characterised by a dark, often nearly black, complexion; 
black hair, of a kind called “frizzly ” or, incorrectly, “woolly,” i.e. 
each hair is closely rolled up on itself, a condition always associated 
with a more or less flattened or elliptical transverse section; a 
moderate or scanty development of beard; an almost invariably 
dolichocephalic skull ; small and moderately retreating jugal bones 
(mesopic face); a very broad and flat nose, platyrhine in the 
skeleton ; moderate or low orbits; prominent eyes; thick, everted 
lips; prognathous jaws ; large teeth (macrodont) ; a narrow pelvis 
(index in the male 90 to 100); a long forearm (humero-radial 
index 80); and certain other proportions of the body and limbs 
which are being gradually worked out and reduced to numerical 
expression as material for so doing accumulates. 

The most characteristic examples of the second great type, the 
Mongolian or Xanthous, or “yellow,” have a yellow or brownish 
complexion ; black coarse straight hair, without any tendency to curl, 
and nearly round in section, on all other parts of the surface except 
the scalp scanty and late in appearing; a skull of variable form, 
mostly mesocephalic (though extremes both of dolichocephalism and 
brachycephalism are found in certain groups of this type) ; a broad 
and flat face, with prominent, anteriorly-projecting jugal bones 
(platyopic: face); nose small, mesorhine or leptorhine ; orbits high 
and round, with very little development of glabella or supraciliary 
ridges ; eyes sunken, and with the aperture between the lids narrow ; 
in the most typical members of the group with a vertical fold of 
skin over the inner canthus, and with the outer angle slightly 
elevated ; jaws mesognathous; teeth of moderate size (mesodont). 
The proportions of the limbs and form of the pelvis have yet to be 
worked out, the results at present obtained showing great diversity 
among different individuals of what appear to be well-marked races 
of the group, but this is perhaps due to the insufficient number of 
individuals as yet examined with accuracy. 

The last type, which, for want of a better name, we must still 
call by the misleading one that has the priority, Caucasian, or 
“white,” has usually a light-complexioned skin (although in some, in 
so far aberrant cases, it is as dark as in the Negroes); hair fair or 
black, soft, straight, or wavy, in section intermediate between the 
flattened and cylindrical form; beard fully developed; form of 


746 PRIMATES 


cranium variable, mostly mesocephalic ; jugal bones retreating ; face 
narrow and projecting in the middle line (pro-opic); orbits moderate ; 
nose narrow and prominent (leptorhine); jaws orthognathous ; teeth 
small (microdont) ; pelvis broad (pelvic index of male 80); forearm 
short, relatively to humerus (humero-radial index 74). 

In endeavouring to subdivide into minor groups the numerous 
and variously-modified individuals which cluster around one or 
other of these great types—a process quite necessary for many 
practical or descriptive purposes—the distinctions afforded by the 
study of physical characters are often so slight that it becomes 
necessary to take other considerations into account, among which 
geographical distribution and language hold an important place. 


I. The Ethiopian or Negroid races may be primarily arranged as 
follows :— 

A. African or Typical Negroes.—Inhabitants of all the central 
portion of the African continent, from the Atlantic on the west to 
the Indian Ocean on the east, greatly mixed all along their 
northern frontier with Hamitic and Semitic Melanochroi, a mixture 
which, taking place in various proportions and under varied con- 
ditions, has given rise to many of the numerous races and tribes 
inhabiting the Sudan. 

A branch of the African Negroes are the Bantu—distinguished 
chiefly, if not entirely, by the structure of their language. Physic- 
ally indistinguishable from the other negroes with whom they come in 
contact in the Equatorial regions of Africa, the Southern Bantu, or 
Kaffirs, as they are generally called, show a marked modification of 
type, being lighter in colour, having a larger cranial capacity, less 
marked prognathism, and smaller teeth. Some of these changes 
are probably due to crossing with other races. 

B. The Negrillos—diminutive sub-brachycephalic tribes, inhabit- 
ing the dense forests of Central and Western Equatorial Africa— 
represent a distinct section of the Negro race. They form the 
only exceptions to the general dolichocephaly of the African branch 
of the Negroid division, and when found in a pure state are the 
smallest of all known human races, averaging scarcely more than 
4 feet in height. The colour of their skin is yellowish rather than 
black. 

C. The Bushmen (Bosjesmen, men of the woods, of the Dutch 
colonists of South Africa) constitute a very distinct modification of 
the Negro type. The hair shows the extreme of the frizzly 
character ; being shorter and less abundant than that of the 
ordinary Negro, it has the appearance of growing in separate tufts, 
which coil up together into rounded balls compared to “ pepper- 
corns.” In their yellow complexion, wide cheek-bones, and 
peculiar form of the eyes they so much resemble some of the 


HOMINIDE 747 


Mongolian races that anthropologists have been inclined to trace 
affinities to or admixture with them, although the character of the 
hair makes such a supposition almost inadmissible. The width of 
the cheek-bones and the narrowness of the forehead and chin give 
a lozenge shape to the front view of the face. The forehead is 
prominent and straight; the nose extremely flat and broad, more 
so than in any other race; the lips prominent and thick, although 
the jaws are less prognathous than in the true Negro races. The 
cranium has many special characters by which it can be easily 
distinguished from that of any other race. The average height of 
the males is about 4 feet 8 inches. There is every reason to 
believe that the Bushmen represent the earliest race of which we 
have any knowledge inhabiting the southern part of the African 
continent, but that long before the advent of Europeans upon the 
scene they had been invaded from the north by Negro tribes, who, 
being superior in size, strength, and civilisation, had taken posses- 
sion of the greater part of their territories, and, mingling freely 
with the aborigines, had produced the mixed race called Hottentots, 
who retained the culture and settled pastoral habits of the Negroes, 
with many of the physical features of the Bushmen. These in 
their turn, encroached upon by the Kaffirs from the north and by 
Europeans from the south, are now greatly diminished, and 
threatened with the same fate which will surely soon befall the 
scanty remnant of the early inhabitants who still retain their 
primitive type. 

D. Oceanic Negroes or Melanesians.—These include the Papuans 
of New Guinea and the majority of the inhabitants of the islands of 
the Western Pacific, and form also a substratum of the population, 
greatly mixed with other races, of regions extending far beyond 
the present centre of their area of distribution. 

They are represented, in what may be called a hypertypical 
form, by the extremely dolichocephalic Kai Colos, or mountaineers 
of the interior of the Fiji Islands, although the coast population of 
the same group has lost the distinctive characters by crossing. In 
many parts of New Guinea and the great chain of islands extending 
eastwards and southwards ending with New Caledonia they are 
found in a more or less pure condition, especially in the interior and 
more inaccessible portions of the islands, almost each of which 
shows special modifications of the type recognisable in details of 
structure. Taken altogether, their chief physical distinction from 
the African Negroes lies in the fact that the glabella and supra- 
orbital ridges are generally well developed in the males, whereas in 
Africans this region is usually smooth and flat. The nose also, 
especially in the northern part of their geographical range, New 
Guinea, and the neighbouring islands, is narrower (often mesorhine) 
and prominent. The cranium is generally higher and narrower. 


748 PRIMATES 


It is, however, possible to find African and Melanesian skulls quite 
alike in essential characters. 

The now extinct inhabitants of Tasmania were probably pure, 
but aberrant, members of the Melanesian group, which had 
undergone a modification from the original type, not by mixture 
with other races, but in consequence of long isolation, during which 
special characters had been gradually developed. Lying completely 
out of the track of all civilisation and commerce, even of the most 
primitive kind, they were little liable to be subject to the influence 
of any other race; and there is in fact nothing among their 
characters which could be accounted for in the way above 
suggested, as they were intensely, even exaggeratedly, Negroid 
in the form of nose, projection of mouth, and size of teeth, 
typically so in character of hair, and aberrant chiefly in the width 
of the skull in the parietal region. A cross with any of the 
Polynesian or Malay races sufficiently strong to produce this 
would, in all probability, have also left some traces on other parts 
of their organisation. 

On the other hand, in many parts of the Melanesian region 
there are distinct evidences of large admixture with Negrito, Malay, 
and Polynesian elements in varying proportions, producing numerous 
physical modifications. In many of the inhabitants of the great 
island of New Guinea itself and of the islands lying around it this 
mixture can be traced. In the people of Micronesia in the north 
and New Zealand in the south, although the Melanesian element 
is present, it is completely overlaid by the Polynesian, but there 
are probably few, if any, of the islands of the Pacific in which 
it does not form some factor in the composite character of the 
natives. 

The inhabitants of the continent of Australia have long been 
a puzzle to ethnologists. Of Negroid complexion, features, and 
skeletal characters, yet without the characteristic frizzly hair, their 
position has been one of great difficulty to determine. They have, 
in fact, been a stumbling-block in the way of every system proposed. 
The solution, supported by many considerations too lengthy to enter 
into here, appears to lie in the supposition that they are not a 
distinct race at all, that is, not a homogeneous group formed by the 
gradual modification of one of the primitive stocks, but rather a 
cross between two already-formed branches of these stocks. Accord- 
ing to this view, Australia was originally peopled with frizzly-haired 
Melanesians, such as those who still do, or did before the European 
invasion, dwell in the smaller islands which surround the north, 
east, and southern portions of the continent, but that a strong 
infusion of some other race, probably a low form of Caucasian 
Melanochroi, such as that which still inhabits the interior of the 
southern parts of India, has spread throughout the land from the 


HOMINID A 749 


north-west, and produced a modification of the physical characters, 
especially of the hair. This influence did not extend across Bass’s 
Straits into Tasmania, where, as just said, the Melanesian element 
remained in its purity. It is more strongly marked in the northern 
and central parts of Australia than on many portions of the southern 
and western coasts, where the lowness of type and more curly hair, 
sometimes closely approaching to frizzly, show a stronger retention 
of the Melanesian element. If the evidence should prove sufficiently 
strong to establish this view of the origin of the Australian natives, 
it will no longer be correct to speak of a primitive Australian, or 
even Australoid, race or type, or look for traces of the former 
existence of such a race anywhere out of their own land. Absolute 
proof of the origin of any race is, however, very difficult, if not 
impossible, to obtain, and there is nothing to exclude the possibility 
of the Australians being mainly the direct descendants of a very 
primitive human type, from which the frizzly-haired Negroes may 
be an offset. This character of hair is probably a specialisation, 
for it seems very unlikely that it was the attribute of the common 
ancestors of the human race. 

E. The fourth branch of the Negroid race consists of the 
diminutive round-headed people called Negritos, still found in a 
pure or unmixed state in the Andaman Islands, and forming a 
substratum of the population, though now greatly mixed with in- 
vading races, especially Malays, in the Philippines, and many of 
the islands of the Indo-Malayan Archipelago, and of some parts 
of the southern portion of the mainland of Asia. They also con- 
tribute to the varied population of New Guinea, where they appear 
to merge into the taller, longer-headed, and longer-nosed Melanesians 
proper. They show in a very marked manner some of the most 
striking anatomical peculiarities of the Negro race, such as the 
frizzly hair, the proportions of the limbs, especially the humero- 
radial index, and the form of the pelvis; but they differ in many 
cranial and facial characters, both from the African Negroes on the 
one hand, and the typical Oceanic Negroes, or Melanesians, on the 
other, and thus form a very distinct and well-characterised group. 
Wherever they are still found they are obviously holding their 
own with difficulty, if not actually disappearing, and there is much 
about their condition of civilisation and the situations in which 
they occur to induce us to look upon them, as in the case of the 
Negrillos of Central and the Bushmen of South Africa, as the 
remains of a population which occupied the land before the incom- 
ing of the present dominant races. 


II. The principal groups that can be arranged round the Mon- 


golian type are as follows :— 
A. The Eskimo appear to be a branch of the typical North 


750 PRIMATES. 


Asiatic Mongols, who in their wanderings northwards and east- 
wards across the American continent, where they have been isolated 
almost as perfectly as an island population would be, hemmed 
in on one side by the eternal Polar ice, and on the other by hostile 
tribes of American Indians, with which they rarely, if ever, mingled, 
have gradually developed characters, most of which are strongly- 
expressed modifications of those seen in their allies who still remain 
on the western side of Behring Strait. It has also been shown 
that these special characteristics gradually increase from west to 
east, and are seen in their greatest perfection in the inhabitants 
of Greenland, at all events in those where no crossing with the 

Danes has taken place. A typical Eskimo skull presents a com- 
bination of characters by which it can be at once distinguished 
from that of any other of the groups of mankind. Such scanty 
remains as have yet been discovered of the earliest inhabitants of 
Europe do not present any structural affinities to this type, and 
there is therefore no justification for the supposition that they 
belonged to the same race, although it is not unlikely that similar 
external conditions may have led them to adopt similar modes of life. 

B. The typical Mongolian races constitute the present popula- 
tion of Northern and Central Asia. They are not very distinctly, 
but still conveniently for descriptive purposes, divided into a 
Northern and a Southern group. 

_ @ The members of the former, Mongolo-Altaic or Sibiric group, 
are united by the affinities of their language. These people, from 
the cradle of their race in the great plateau of Central Asia, have 
at various times poured out their hordes upon the lands lying to the 
west, and thence penetrated almost to the heart of Europe. The 
Lapps, Finns, the Magyars, and the Turks are each the descendants 
of one of these waves of incursion, but they have for so many genera- 
tions intermingled with the peoples through whom they have passed 
in their migrations, or whom they have found in the countries in 
which they have ultimately settled, that their original physical 
characters have been completely modified. Even the Lapps, that 
diminutive tribe of nomads inhabiting the most northern parts of 
Europe, supposed to be of Mongolian descent, show so little of the 
special attributes of that branch that it is difficult to assign them 
a place in it in a classification based upon physical characters. 
The Japanese are said by their language to be allied rather to the 
Northern than to the following branch of the Mongolian stock. 

b. The southern Mongolian or Sinitic group, divided from ‘the 
former chiefly by language and habits of life, includes the greater 
part of the population of China, Tibet, Burma, and Siam. 

C. The next great division of Mongoloid people is the Malay, 
forming the bulk of the population of the Indo-Malayan Archipelago 
and (mixed with the Negro) of Madagascar, subtypical it is true, 


HOMINID 751 


but to which an easy transition can be traced from the most char- 
acteristic members of the type. 

D. The brown Polynesians, Malayo-Polynesians, Mahoris, 
Sawaioris, or Kanakas, as they have been variously called, seen 
in their greatest purity in the Samoan, Tongan, and Eastern Poly- 
nesian Islands, are still more modified, and possess less of the 
characteristic Mongolian features; but yet it is difficult to place 
them anywhere else in the system. The large infusion of the 
Melanesian element throughout the Pacific must never be forgotten 
in accounting for the characters of the people now inhabiting the 
islands—an element in many respects so diametrically opposite to 
the Mongolian that it would materially alter the characters, especi- 
ally of the hair and beard, which has been with many authors a 
stumbling-block to the affiliation of the Polynesian with the Mongolian 
stock. This mixture is physically a fine one, and in some propor- 
tions produces a combination, as seen, for instance, in the Maories 
of New Zealand, which in all definable characters approaches quite 
as near, or nearer, to the Caucasian type than to either of the 
stocks from which it may be presumably derived. This resemblance 
has led some ethnologists to infer a real extension of the Caucasian 
element at some very early period into the Pacific Islands, and to 
look upon their inhabitants as the product of a mingling of all the 
three great types of men. Though this is a very plausible theory, 
it rests on little actual proof, since the combination of Mongolo- 
Malayan and Melanesian characters in different degrees, together 
with the local variations certain to arise in communities so isolated 
from each other and exposed to such varied conditions as the in- 
habitants of the Pacific Islands, would probably account for all the 
modifications observed among them. 

E. The native population (before the changes wrought by the 
European conquest) of the great continent of America, excluding 
the Eskimo, present, considering the vast extent of the country 
they inhabit and the great differences of climate and other sur- 
rounding conditions, a remarkable similarity of essential characters 
with much diversity of detail. 

The construction of the numerous American languages, of which 
as many as twelve hundred have been distinguished, is said to point 
to unity of origin, as, though widely different in many respects, 
they are all, or nearly all, constructed on the same general gram- 
matical principle—that called polysynthesis—which differs from that 
of the languages of any of the Old World nations. The mental 
characteristics of all the American tribes have much that is in 
common ; and the very different stages of culture to which they 
had attained at the time of the conquest, as that of the Incas and 
Aztecs and the hunting or fishing tribes of the north and south, 
which have been quoted as evidence of diversities of race, were not 


752 PRIMATES 


greater than those between different nations of Europe, as Gauls and 
Germans on the one hand, and Greeks and Romans on the other, in 
the time of Julius Casar. Yet all these were Aryans, and in treat- 
ing the Americans as one race it is not intended to imply that they 
are more closely allied than the different Aryan peoples of Europe 
and Asia. The best argument that can be used for the unity of 
the American race—using the word in a broad sense—is the great 
difficulty of forming any natural divisions in it founded upon physical 
characters. Thus there is no difference throughout the whole con- 
tinent in the important character of the hair, this beingalways straight 
and lank, long and abundant on the scalp, but sparse elsewhere. 
The colour of the skin, notwithstanding the enormous differences of 
climate under which many members of the group exist, varies but 
little. It is true that in the features and cranium certain special 
modifications prevail in different districts, but the same forms 
reappear at widely separated parts of the continent. Thus skulls 
almost undistinguishable from one another may be met with from 
Vancouver's Island, from Peru, and from Patagonia. 

Naturalists who have admitted but three primary types of the 
human species have always found a difficulty with the Americans, 
hesitating between placing them with the Mongolian or so-called 
“yellow” races, or elevating them to the rank of a primary group. 
Cuvier, indeed, does not seem to have been able to settle this point 
to his own satisfaction, and leaves it an open question. Although 
the large majority of Americans have in the special form of the 
nasal bones, leading to the characteristic high bridge of the nose of 
the living face, in the well-developed superciliary ridge and retreat- 
ing forehead, characters which distinguish them from the typical 
Asiatic Mongol, yet in many other respects they resemble them so 
closely that, while still admitting the difficulties of the case, we are 
inclined to include them as aberrant members of the Mongolian 
type! It is, however, quite open to any one adopting the Negro, 
Mongolian, and Caucasian groups as primary divisions to place the 
Americans apart as a fourth. 

Now that the high antiquity of man in America—perhaps as 
high as that which he has in Europe—has been discovered, the 
puzzling problem, from which part of the Old World the people of 
America have sprung, has lost its significance. It is, indeed, quite 
as likely that the people of Asia may have been derived from 
America as the reverse. However this may be, the population of 
America, except at the extreme north, was, before the time of 
Columbus, practically isolated from the rest of the world. Such 
visits as those of the early Norsemen to the coasts of Greenland, 

1 No one can have seen a group of Botocudos from Brazil or of natives of 


Tierra del Fuego without being struck by their markedly Mongolian external 
characteristics. 


HOMINIDE 753 


Labrador, and Nova Scotia, or the purely accidental stranding 
of a canoe containing survivors of a voyage across the Pacific or 
the Atlantic, can have had little appreciable effect upon the char- 
acteristics of the people. It is difficult, therefore, to look upon the 
anomalous and special characters of the American people as the 
effects of crossing, as was suggested in the case of the Australians 
—a consideration which gives more weight to the view of treating 
them as a distinct primary division. 


III. The Caucasian, Eurafrican, or white division, includes the 
two groups called by Professor Huxley Xanthochroi and Melano- 
chroi, which, though differing in colour of eyes and hair, agree so 
closely in all other anatomical characters, so far, at all events, as has 
at present been demonstrated, that it seems preferable to consider 
them as modifications of one great type than as primary divisions 
of the species. Whatever their origin may have been, they are now 
intimately blended, though in different proportions, throughout the 
whole of the region of the earth they inhabit; and it is to the 
rapid extension of both branches of this race that the great changes 
now taking place in the ethnology of the world are mainly due. 

A. The Xanthochroi, or blonde type, with fair hair, eyes, and 
complexion, chiefly inhabit Northern Europe (Scandinavia, Scotland, ~ 
and North Germany), but, although much mixed with the next 
group, they also extend as far as Northern Africa and Afghanistan. 
Their mixture with Mongoloid people has given rise to the Lapps, 
Finns, and some of the tribes of Northern Siberia. 

B. Melanochroi, with black hair and eyes, and skin of almost 
all shades from white to black. They comprise the great majority 
of the inhabitants of Southern Europe, Northern Africa, and South- 
West Asia, and consist mainly of the Aryan, Semitic, and Hamitic 
families. The Dravidians of India, the Veddahs of Ceylon, and 
probably the Ainos of Japan, and the Maoutze of China, also 
belong to this race, which may have contributed something to the 
mixed character of some tribes of Indo-China and the Polynesian 
Islands, and, as before said, have given at least the characters of 
the hair to the otherwise Negroid inhabitants of Australia. In 
Southern India they are largely mixed with a Negrito element, 
and in Africa, where their habitat becomes coterminous with that 
of the Negroes, numerous cross-races have sprung up between them 
all along the frontier line. The ancient Egyptians were nearly pure 
Melanochroi, though often showing in their features traces of their 
frequent intermarriages with their Ethiopian neighbours to the 
south. The Copts and fellahs of modern Egypt are their little- 
changed descendants. 

In offering this scheme of classification of the varieties of the 
human species, it is not suggested that it is one universally accepted 

48 


754 PRIMATES 


by anthropologists, or that it is likely to be final. Whatever care 
be bestowed upon the arrangement of already acquired details, or 
whatever judgment be shown in their due subordination one to 
another, the acquisition of new knowledge may at any time call for 
a complete or partial rearrangement of the system. The difficulties 
which encompass the subject have, indeed, been already indicated, 
and will be found abundantly illustrated in the writings of those 
authors who have specially devoted themselves to its elucidation. 


Bibliography.—P. Topinard, Eléments @’ Anthropologie Générale, 1885; A. de 
Quatrefages, Histoire Générale des Races Humaines (1. Questions Générales, 1887 ; 
2. Classification des Races Humaines, 1889); Quatrefages and Hamy, Crania 
Ethnica (1873-1879) ; D. G. Brinton, Races and Peoples, 1890. 


AaRbD-WoLr, 540 
Aard-Vark, 211: 
Absorbent system, 63 
Acanthoglossus, 125 
Acanthomys, 476 
Aceratherium, 411 
Achenodon, 292 

_ Achyrodon, 114 
Acrobates, 155 
Acrotheriwm, 440 
Adapis, 697 
Adapisorex, 634 
Adapisoricida, 634 
Adapisoriculus, 634 
Addax, 345 
Adenota, 339 
Adinotheriwm, 440 
Aslurictis, 524 
Ajlurodon, 562 
Asluroidea, 501 
aliluropus, 560 
Ailurus, 562 
pyceros, 341 
Aipyprymnus, 164 
Agabelus, 260 
Agouti, 488 
Agriocherus, 293 
Ai, 182 

Air-sacs, 68 
Alactaga, 480 
Albinism, 10 
Alcelaphus, 334 
sllces, 326 
Allantois, 77 
«ldlodon, 111 
Allops, 413 
Allotheria, 109 
Alpaca, 303 
Amblotherium, 114 
Amblypoda, 436 
Amorphochilus, 666 
Amphictis, 539 
Amphicyon, 555 


INDEX 


Amphidozotherium, 6384 
Amphilestes, 114 
Amphiperatherium, 135 
Amphisorex, 628 
Amphitheriwm, 114 
Amphitragulus, 330 
Amynodon, 412 
Anaptomorphus, 697 
Anchilophus, 376 
Anchippodus, 441 
Anchitherium, 376 
Ancylopoda, 413 
Ancylotherium, 413 
Anoa, 361 
Anomaluride, 449 
Anomalurus, 449 
Anoplotheriida, 293 
Anoplotherium, 294 
Anteater, 191 

Scaly, 205 
Antebrachium, 47 
Antechinomys, 139 
Antelopes, 334 
Anthops, 657 
Anthorhina, 674 
Anthracotheriide, 292 
Anthracotherium, 292 
Anthropoidea, 699 
Anthropopithecus, 736 
Antilocapra, 333 
Antilocapride, 333 
Antilope, 340 
Antlers, 308 
Antrozous, 661 
lnurosorex, 626 
Aoudad, 356 
Apar, 199 
Ape, 699 
alphelops, 411 
Aphelotherium, 697 
alrcheelurus, 524 
Archeoceti, 246 
Archeomys, 484 


Archizonurus, 157 
Arctictis, 584 
Arctocebus, 693 
Arctocephalus, 595 
Arctocyon, 609 
Arctocyonide, 609 
Arctogale, 533 
Arctoidea, 556 
Arctomyine, 454 
Arctomys, 454 
Arctonyx, 574 
Arctotherium, 561 
Argali, 355 
Armadillo, 195 
Artibeus, 676 
Artiodactyla, 275 
Arvicola, 466 
Arvicoline, 465 
Ass, 383 
Atalapha, 663 
Ateles, 715 
Atherura, 487 
Auchenia, 298 
Aulacodus, 483 
Aulaxinuus, 723 
Aurochs, 367 
Australasian region, 102 
Avahis, 686 

Axis, 320 
Aye-aye, 695 


Babirusa, 287 
Baboon, 719 
Bachitherium, 807 
Badger, 575 
American, 576 
Sand, 575 
Balena, 236 
Balenide, 234 
Balenodon, 251 
Balenoidea, 234 
Balenoptera, 242 
Balenotus, 240 


756 


INDEX 


Bandicoot, 141 
Banteng, 365 
Bassaricyon, 566 
Bassaris, 566 
Bats, 641 
Bathyergus, 478 
Bdeogale, 537 
Bear, 558 

Beaver, 458 

Beisa, 343 

Beluga, 262 
Berardius, 256 
Bettongia, 163 
Bharal, 356 
Bibos, 360 
Bighorn, 355 
Binturong, 534 
Bison, 362 
Black-Fish, 269 
Bladder, 69 
Blarina, 624 
Blastomeryx, 330 
Blaubok, 343 
Blessbok, 335 
Blood, 63 
Bolodon, 111 
Boneta, 653 
Bontebok, 384 
Bosch-Vark, 286 
Boselaphus, 345 
Bothriolabis, 291 
Bottlenose, 253, 270 
Bovide, 334 
Brachium, 47 
Brachyphylla, 675 
Brachytarsomys, 465 
Brachyurus, 712 
Bradypodide, 179 
Bradypus, 181 
Brain, 69 
Bramatherium, 333 
Brocket, 330 
Brontotherium, 413 
Bruta, 176 
Bubalus, 361 
Budorcas, 351 
Buffalo, 361 
Bush-dog, 553 


CacHALot, 249 
Cadurcotherium, 412 
Caecum, 59 
Coelogenys, 489 
Ceenopithecus, 696 
Ceenothertide, 294 
Ceenotherium, 294 
Callinycteris, 655 
Callithrix, 718 
Callophoca, 606 
Calomys, 463 
Caloprymnus, 164 


Calotragus, 339 
Camel, 296 
Camelidee, 295 
Camelus, 296 
Canidae, 544 
Canis, 546 

Capra, 852 
Capreolus, 327 
Capromys, 482 
Capybara, 491 
Caracal, 518 
Cardiatherium, 491 
Cardiomys, 491 
Cariacus, 829 
Caribou, 324 
Carnivora, 496 
Carollia, 674 
Carponycteris, 654 
Carpus, 48 
Carterodon, 484 
Castor, 457 
Castoride, 457 
Castorotdide, 488 
Castoroides, 488 
Cat, 517 

Cavia, 489 
Caviidee, 489 
Cavy, 490 
Cayluxotherium, 621 
Cebidee, 711 
Cebocherus, 292 
Cebus, 717 
Cement, 15 
Centetes, 637 
Centetide, 637 
Centurio, 676 
Cephalogale, 562 
Cephalophus, 338 
Cephalorhynchus, 266 
Cephalotes, 653 
Cercocebus, 723 
Cercoleptes, 567 
Cercomys, 483 
Cercopithecide, 718 
Cercopithecus, 724 
Cerivoula, 664 
Cervalces, 327 
Cervicapra, 340 
Cervide, 313 
Cervinee, 316 
Cervulus, 316 
Cervus, 319 
Cetacea, 225 
Cetotherium, 245 
Chenohyus, 291 
Cheetomys, 486 
Chalcochloris, 639 
Chalicomys, 458 
Chalicothertide, 418 
Chalicotherium, 418 
Chalinolobus, 662 


Chamois, 349 
Champsodelphis, 259 
Cheeta, 523 
Chevrotain, 305 

Water, 306 
Chilonycteris, 672 
Chimarrogale, 626 
Chimpanzee, 736 
Chinchilla, 487 
Chinchillida, 487 
Chirogaleus, 689 
Chiromeles, 669 
Chiromyide, 694 
Chiromys, 695 
Chironectes, 184 
Chiroptera, 641 
Chiru, 341 
Chiruromys, 476 
Chlamydophorine, 196 
Chlamydophorus, 196 
Chlamydotherium, 201 
Cheeronycteris, 674 
Cheropotumidc, 292 
Cheropotamus, 292 
Cheeropsis, 280 
Cheropus, 148 
Cholepus, 182 
Chorion, 77 
Chrysochloride, 638 
Chrysochloris, 639 
Chrysothrix, 714 
Cimoliomys, 118 
Circulation, 63 
Civet, 526 

Palm, 532 
Classification, 84, 88 
Claviglis, 460 
Claws, 12 
Coati, 566 
Cobus, 339 
Colodon, 184 
Colops, 658 
Cogia, 250 
Coleiira, 667 
Colobus, 727 
Colour, 8 
Comphotherium, 621 
Condylarthra, 438 
Condylura, 630 
Conepatus, 574 
Connochectes, 336 
Contracavia, 491 
Coryphodon, 437 
Coryphodontide, 488 
Cotylophora, 307 
Cotylopide, 293 
Cotylops, 293 
Coypu, 482 
Cranium, 35 
Crassitherium, 223 
Creodonta, 606 


INDEX 


757 


Cricetodipus, 479 
Cricetodon, 464 
Cricetomys, 477 
Cricetus, 463 
Criotherium, 349 
Crocidura, 626 
Crossarchus, 537 
Crossopus, 625 
Crusta Petrosa, 15 
Cryptophractus, 201 
Cryptopithecus, 699 
Cryptoprocta, 525 
Ctenacodon, 112 
Ctenodactylus, 481 
Ctenomys, 482 
Cuniculus, 470 
Cuscus, 149 
Cyclopidius, 293 
Cycloturus, 1938 
Cyncelurus, 523 
Cynictis, 537 
Cynocephalus, 719 
Cynodictis, 555 
Cynogale, 534 
Cynohycenodon, 608 
Cynoidea, 544 
Cynomys, 455 
Cynonycteris, 652 
Cynopithecus, 722 
Cynopterus, 653 
Cyon, 551 
Cystophora, 605 


Dacrytherium, 294 
Dactylomys, 483 
Dactylopsila, 152 
Damalis, 334 
Dapheenus, 555 
Dasymys, 462 
Dasypodide, 194 
Dasypodine, 197 
Dasypotherium, 201 
Dasyprocta, 488 
Dasyproctide, 488 
Dasypus, 197 
Dasyuride, 136 
Dasyurus, 138 
Daubentonia, 695 
Deer, 317, 319 
Delphinapterus, 262 
Delphinide, 260 
Delphinoidea, 247 
Delphinus, 271 
Dendrohyraz, 418 
Dendrolagus, 165 
Dendromys, 463 
Dental system, 13 
Dentine, 14 
Deomys, 473 
Dermoptera, 614 
Desman, 629 


Desmodus, 677 
Desmotylus, 223 
Diaphragm, 67 
Diceratherium, 411 
Dichobunus, 294 
Dichodon, 294 
Dichodontide, 294 
Diclidurus, 668 
Dicolpomys, 484 
Dicotyles, 289 
Dicotylide, 289 
Didelphia, 128 
Didelphyide, 133 
Didelphys, 184 
Didymictis, 5389 
Digestive system, 53 
Dinictis, 523 
Dinoceras, 487 
Dinocyon, 556 
Dinomyide, 489 
Dinomys, 489 
Dinotheriide, 435 
Dinotherium, 435 
Dinoziphius, 251 
Diobroticus, 458 
Dioplotherium, 223 
Diphylla, 678 
Diphyodont, 20 
Diplarthra, 275 
Diplomesodon, 626 
Dipodide, 479 
Dipodomys, 479 
Dipodops, 479 
Diprotodon, 171 
Diprotodontia, 144 
Diprotodontide, 171 
Dipus, 480 
Disteechurus, 155 
Dedicurus, 203 
Dog, 551 
Dolichophyllum, 673 
Dolichopithecus, 728 
Dolichotis, 490 
Dolphin, 270 
Dorcatherium, 306 
Dorcopsis, 166 
Dormouse, 459 
Douroucoli, 714 
Dremotherium, 330 
Dromatheriwm, 113 
Dromicia, 154 
Drypolestes, 114 
Dryopithecus, 738 
Duck-bill, 120 
Ductless glands, 65 
Dugong, 221 
Duikerbok, 338 
Duplicidentata, 491 


Echidna, 125 
Echidnide, 124 


Echinogale, 634 
Echinomys, 483 
Echinothrix, 477 
Edentata, 176 
Effodientia, 178 
Eland, 348 
Elaphodus, 318 
Elasmognathus, 371 
Elasmothertum, 411 
Eleotragus, 340 
Elephant, 424 
Elephantide, 423 
Elephas, 424 
Eleutherocercus, 203 
Eliomys, 459 
Eliurus, 465 

Elk, 326 

Ellobius, 472 
Llotherium, 292 
Emballonura, 667 
Emballonuride, 666 
Enamel, 15 
Enhydra, 570 
Enhydriodon, 57) 
Emhydrocyon, 562 
Entomophaga, 178 
Eohippus, 374 
Eomys, 464 
Eonycteris, 654 
Eotherium, 224 
Epiblema, 488 
Epiglottis, 67 
Lpihippus, 374 
Epomophorus, 650 
Eporeodon, 293 
Liquide, 376 
Lquus, 381 
Erethizon, 484 
Ericulus, 638 
Erinaceide, 619 
Erinaceus, 620 
Eriodes, 715 
Ermine, 590 
Eschatius, 808 
Ethiopian region, 98 
Eucastor, 458 
Eucetus, 251 
Lupetaurus, 454 
ELupleres, 538 
Euryceros, 346 
Euryurus, 208 
Eusmilus, 524 
Lutatus, 201 
Eutheria, 173 
Evotomys, 467 
Eye, 72 


Fatiow Derr, 323 
Felidae, 502 

Felis, 502 
Felsinotherium, 223 


758 


INDEX 


Fennec, 553 
Fennecus, 553 
Feresia, 270 
Fiber, 470 
Flying Fox, 651 
Lemur, 615 
Squirrel, 453 
Foot, 52 
Fossa, 527 
Foussa, 525 
Fox, 552 
Fox-Bat, 651 
Furia, 666 
Furipterus, 666 


Galago, 690 
Galeopithecide, 614 
Galeopithecus, 614 
Galera, 579 
Galictis, 579 
Galidea, 538 
Galidictis, 538 
Gaur, 365 
Gayal, 365 
Gazella, 341 
Gelocus, 294 
Gemsbok, 343 
Genet, 528 
Genetta, 528 
Geogale, 635 
Geographical distribution, 
9 


Geological 
107 
Geomyide, 478 
Geomys, 478 
Georychus, 478 
Gerbillinz, 462 
Gerbillus, 462 
Gibbon, 728 
Giraffa, 331 
Girafide, 330 
Glands, 12 
Glauconycteris, 662 
Globicephalus, 268 
Glossonycteris, 674 
Glossophaga, 674 
Glutton, 591 
Glyptodon, 203 
Glyptodontide, 202 
Gnu, 336 
Golunda, 476 
Goat, 352 
Gopher, 478 
Goral, 351 
Gorilla, 734 
Grampus, 267 
Grampus, 270 
Graphiurus, 459 
Greenland Whale, 236 
Grimmia, 338 


distribution, 


Grisonia, 579 
Ground Sloth, 184 
Gryphoca, 606 
Grypotherium, 189 
Guanaco, 301 
Guib, 347 
Guinea-Pig, 490 
Gulo, 591 
Gymnobelideus, 154 
Gymnoptychus, 454 
Gymnura, 619 


Habrocoma, 482 
Habrothrix, 464 
Hair, 7 
Halicherus, 601 
Halicore, 220 
Halicoride, 220 
Halitheriide, 222 
Halitherium, 222 
Tallomys, 465 
Hamster, 463 
Hapale, 710 
Hapalemur, 689 - 
Hapalidee, 709 
Hapalotis, 476 
Haploceros, 351 
Haplodon, 457 
Haplodontide, 457 
Hare, 492 
Harpyia, 653 
Harpyiocephalus, 663 
Harte-beest, 335 
Hearing, 73 

Heart, 63 
Hedgehog, 620 
Helicophora, 340 
Helictis, 578 
Heliophobius, 478 
Helladotherium, 333 
Helogale, 537 
Hemiuuchenia, 303 
Hemicentetes, 637 
Iemiderma, 674 
Hemigale, 533 
Hemigalidea, 588 
Hemitragus, 354 
Herpestes, 535 
Herpetocetus, 245 
Herpetotherium, 135 
Heterocephalus, 478 
Heterocetus, 245 
Heterodont, 23° 
Heterohyrax, 418 
Heteromys, 479 
ITipparion, 380 
LTippodactylus, 381 
ITippohyus, 291 
LTippopotamide, 278 
THippopotamus, 278 
Hipposiderus, 657 


Hippotigris, 384 
Hippotragus, 348 
Holochilus, 464 
Holomeniscus, 303 
Homalodontotherium, 412, 
414 
Hominide, 740 
Homo, 739 
Homodont, 22 
Hoofs, 12 
Hoolock, 729 
Hoplocetus, 251 
Hoplophoneus, 524 
Hoplophorus, 202 
Horns, 310 
Horse, 382 
Hunting dog, 553 
Hyena, 540 
Hycenarctus, 561 
Hycenide, 540 
Hycenocyon, 562 
Hycenodon, 608 
Hyenodontide, 608 
Hydaspitherium, 333 
Hydrocherus, 490 
Hydromyine, 461 
Hydromys, 461 
Hydropotes, 328 
Hylobates, 728 
Hylomys, 619 
Hyoid, 39 
Hyomoschus, 306 
Hyopotanvus, 292 
Hyopsodus, 698 
Hyotherium, 291 
Hypertragulus, 307 
Hypogeomys, 465 
Hypsiprymnodon, 162 
Hypsiprymnodontine, 162 
Hypsiprymnopsis, 111 
THypsiprymnus, 163 
Hyrachyus, 373 
Hyracide, 415 
Hyracodon, 412 
Hyracodontotherium, 439 
Hyracoidea, 415 
Hyracotherium, 373 
Hyves, 417 
ITystricide, 484 
Hystricomorpha, 480 
Hystrix, 486 


IBex, 353 
Ichneumon, 535 
Icticyon, 553 
Ictitherium, 539 
Ictonyx, 579 
Ictops, 640 
Indris, 684 
Indrodon, 699 
Inia, 259 


INDEX 


759 


Insectivora, 610 
Intestine, 59 
Inuus, 723 
Ischnoglossa, 674 
Isectolophus, 374 
Issiodoromys, 491 
Ivory, 14 
Txacanthus, 259 


JACKAL, 550 
Jaguar, 521 
Jerboa, 480 


Kanearoo, 159 
Kerivoula= Cerivoula, 
Kidney, 69 
Killer, 267 
Kinkajou, 567 
Koala, 156 
Koalemus, 157 
Kobus = Cobus, 
Kogia = Cogia, 
Kudu, 348 
Kusimanse, 538 


Lagenorhynchus, 270 
Lagidium, 488 
Lagomyide, 491 
Lagomys, 491 * 
Lagorchestes, 166 
Lagostomus, 488 
Luagostrophus, 165 
Lagothriz, 716 
Lambdotheritde, 413 
Lambdotherium, 4138 
Langur, 727 
Lantanotherium, 618 
Larynx, 67 
Lasionycteris, 661 
Lata, 570 

Leg, 51 

Lemming, 467 
Lemur, 687 
Lemuride, 683 
Lemuroidea, 682 
Leopard, 514 
Lepidolemur, 689 
Leporide, 492 
Leptictide, 640 
Leptictis, 640 
Leptobos, 367 
Leptomeryx, 307 
Leptonycteris, 674 
Leptonyx, 605 
Leptotragulus, 304 
Lepus, 492 
Lestodon, 189 
Leucocyon, 553 
Limnosyops, 413 
Linsang,i530 

Lion, 504 


Liotomus, 113 
Listriodon, 291 
Liver, 60 

Llama, 299, 302 
Lobodon, 605 
Loncheres, 483 
Lonchoglossa, 674 
Lonchorhina, 673 
Lophiodon, 373 
Lophiodontide, 373 
Lophiomeryx, 294 
Lophiomyide, 460 
Lophiomys, 460 
Lophiotherium, 374 
Lophocetus, 259 
Lophostoma, 673 
Loricata, 179 
Loris, 692 
Loxolophodon, 437 
Lungs, 68 

Lutra, 567 
Lycalopex, 552 
Lycaon, 553 
Lymphatics, 65 
Lyncodon, 590 
Lynx, 518 


Macacus, 722 
Macherodus, 524 
Muacrauchenia, 414 
Macrauchentide, 414 
Macroglossus, 654 
Macrophyllum, 673 
Macropodide, 158 
Macropodine, 164 
Macropus, 167 
Macrorhinus, 606 
Macroscelides, 618 
Macroscelidide, 618 
Macrotherium, 418 
Muacrotus, 673 
Malacomys, 462 
Mammary glands, 75 


Mammoth, 428 


Man, 739 
Manatee, 215 
Manatide, 215 
Manatus, 215 
Mandrill, 719 
Manidee, 204 
Manis, 204 
Manus, 48 
Maral, 322 
Markhoor, 354 
Marmoset, 709 
Marmot, 454 
Prairie, 456 
Marsupialia, 128 
Marten, 580 
Martes, 580 
Mastacomys, 476 - 


Mastodon, 481 
Megachiroptera, 650 
Megaderma, 658 
Megaloglossus, 655 
Megamys, 488 
Megaptera, 241 
Megatheriide, 183 
Megatherium, 185 
Melanism, 9 
Meles, 575 
Mellivora, 576 
Melonycteris, 654 
Melursus, 560 
Menacodon, 115 
Meniscoéssus, 113 
Meniscomys, 454 
Meniscotherium, 439 
Menodus, 413 
Mephitis, 572 
Merychippus, 380 
Merycocherus, 293 
Mesodectes, 640 
Mesohippus, 376 
Mesomys, 483 
Mesonychide, 609 
Mesonyx, 609 
Mesopithecus, 727 
Mesoplodon, 254 
Mesotaria, 606 
Mesotherium, 440 
Mesozoic mammals, 108 
Metacarpus, 49 
Metamynodon, 412 
Metatheria, 128 
Metriotherium, 294 
Miacidee, 539 
Aiacis, 539 
Microcavia, 491 
Microcebus, 690 
Microcherus, 696 
Microchiroptera, 655 
Microconodon, 113 
Microgale, 638 
Microlestes, 111 
Micromeryx, 330 
Micronycteris, 673 
Microsorex, 624 
Microsyops, 698 
Microtus, 466 
Midas, 710 
Milk-teeth, 20 © 
Mimon, 674 
Miniopterus, 664 
Mink, 586 
Miohippus, 376 
Miosiren, 223 
Mizxodectes, 699 
Mole, 630 

Golden, 639 

Star-nosed, 630 
Mole-Rat, 477 


760 


INDEX 


Molossus, 670 
Monachus, 604 
Monatheriwm, 606 
Monkey, 699 
Monodelphia, 173 
Monodon, 260 
Monophylla, 674 
Monophyodont, 20 
Moose, 326 
Morenia, 484 
Mormops, 672 
Moropus, 413 
Morotherium, 413 
Morse, 597 
Moschine, 314 
Moschus, 314 
Moufflon, 356 
Mouse, 475 
Mouth, 54 
Mulita, 201 
Multituberculata, 109 
Mungoose, 535 
Muntjac, 316 
Muride, 461 
Mus, 473 
Muscardinus, 460 
Musk Deer, 314 

Ox, 358 

Rat, 470, 626 
Musquash, 470 
Mustela, 579 
Mustelidce, 567 
Mycetes, 711 
Mydaus, 575 
Mylodon, 189 
Myodes, 467 
Myogale, 628 
Myolagus, 492 
Myomorpha, 459 
Myopotamus, 482 
Myoscalops, 478 
Myosorex, 625 
Myoxide, 459 
Myoxus, 459 
Myrmecobiine, 140 
Myrmecobius, 140 
Myrmecophaga, 190 
Myrmecophagide, 190 
Mysurachne, 634 
Mystacina, 671 
Mystacoceti, 234 
Mystacops, 671 
Mystromys, 462 
Myzxocebus, 689 
Myzxopoda, 665 


Nalizs, 12 
Nakong, 346 
Nandinia, 534 
Nanotragus, 339 
Nares, 66 


Narwhal, 261 
Nasalis, 725 
Nasua, 566 
Natalus, 664 
Nearctic region, 102 
Necrogymnurus, 621 
Necrolemur, 696 
Necromantis, 679 
Nectogale, 627 
Nectomys, 464 
Nemorhedus, 350 
Neobalena, 241 
Neofiber, 472 
Neomeris, 266 
Neoplagiaulax, 113 
Neosorex, 624 
Neotoma, 464 
Neotragus, 338 
Neotropical region, 103 
Nerves, 71 
Nesocerodon, 491 
Nesocia, 475 
Nesodon, 439 
Nesomys, 465 
Nesonycteris, 655 
Nesotragus, 339 
Neurotrichus, 629 
Nilghai, 345 
Nimravus, 524 
Noctilio, 668 
Nostrils, 66 
Notharctus, 698 
Nothropus, 188 
Nothrotherium, 184 
Notiosorex, 624 
Notopteris, 654 
Nototheriide, 172 
Nototherium, 171 
Nyctereutes, 552 
Nycteride, 658 
WNycteris, 659 
Nycticebus, 691 
Nycticejus, 662 
Nyctilestes 665 
Nyctinomus, 670 
Nyctipithecus, 714 
Nyctitherium, 665 
Nyctophilus, 661 


Ocetot, 521 
Ochetodon, 464 
Octodon, 481 
Octodontide, 480 
Odobcenus, 597 
Odontoceti, 247 
Ogmorhinus, 605 
Ommatophoca, 605 
Onotragus, 339 
Onychogale, 166 
Onychomys, 463 
Opossum, 133 


Orang, 731 
Orca, 267 
Orcella, 267 
Oreas, 348 
Oreodon, 293 
Oreopithecus, 728 
Oreotragus, 339 
Oriental region, 100 
Ornithodelphia, 117 
Ornithorhynchide, 119 
Ornithorhynchus, 119 
Orohippus, 374 
Orotherium, 374 
Orthaspidotherium, 634 
Orthomys, 484 
Orycteropodide, 208 
Orycteropus, 208 
Oryx, 343 
Oryzomys, 463 
Oryzorictes, 688 
Otaria, 593 
Otariida, 593 
Otocyon, 554 
Otomys, 462 
Otonycteris, 661 
Otopterus, 673 
Otter, 568 

Sea, 571 
Ounce, 517 
Ovaries, 75 
Ovibos, 357 
Oviduct, 75 
Ovis, 354 
Oxen, 360 
Oxhycena, 608 
Oxymycterus, 464 


Paca, 489 
Pachyacanthus, 224 
Pachydermata, 87 
Pachynolophus, 374 
Pachyuromys, 462 
Paciculus, 465 
Palearctic region, 97 
Paleocastor, 458 
Paleocetus, 245 
Paleoerinaceus, 621 
Paleolemur, 697 
Paleomanis, 208 
Paleomeryx, 330 
Palconycteris, 657 
Paleophoca, 606 
Paleopontoporia, 259 
Paleoprionodon, 539 
Paleeoreas, 348 
Paleoryx, 344 
Paleospalax, 629 
Paleosyops, 413 
Paleotapirus, 373 
Paleotheriide, 375 
Paleotherium, 375 


INDEX 


761 


Paleotragoceros, 349 
Palauchenia, 303 
Pathyena, 544 
Palla, 341 
Palm-Civet, 532 
Paloplotherium, 375 
Palorchestes, 170 
Panda, 562 
Pangolin, 205 
Panochthus, 203 
Panther, 514 
Pantholops, 341 
Paradoxurus, 532 
Paramys, 457 
Parasorex, 618 
Peccary, 289 
Pecora, 307 
Pectinator, 481 
Pedetes, 480 
Pediotragus, 339 
Pelea, 339 
Pellegrinia, 484 
Pelvis, 50 
Pelycodus, 699 
Peragule, 143 
Peralestes, 115 
Peramelide, 141 
Perameles, 142 
Peratherium, 135 
Periptychus, 439 
Perissodactyla, 368 
Perodicticus, 693 
Perognathus, 479 
Pes, 52 
Petauroides, 152 
Petaurus, 153 
Petrodromus, 618 
Petrogale, 167 
Petromys, 482 
Phacocherus, 288 
Phalanger, 149 
Phalangeride, 147 
Phalangerine, 149 
Phalanges, 49 
Phalangista, 149 
Phascolarctine, 155 
Phascolarctus, 156 
Phascologale, 139 
Phascolomyide, 144 
Phascolomys, 145 
Phascolonus, 146 
Phascolotherium, 114 
Phenacodus, 439 
Phenacomys, 466 
Phleomine, 462 
Phleomys, 462 
Phloramys, 484 
Phoca, 601 
Phocena, 263 
Phocanella, 606 
Phocidee, 600 


Phylloderma, 674 
Phyllonycteris, 674 
Phyllophaga, 178 
Phyjllorhina, 657 
Phyllostoma, 674 
Phyllostomatide, 672 
Physeter, 248 
Physeteride, 247 
Physeterina, 248 
Physeterula, 251 
Physetodon, 251 
Physodon, 251 
Pica, 492 
Pichiciago, 196 
Pig, 282 
Pilosa, 179 
Pinnipedia, 592 
Pithanotomys, 484 
Pithechirus, 477 
Pithecia, 712 
Placenta, 75 
Plagiaulacide, 113 
Plagiaulax, 111 
Plagiodon, 483 
Platacanthomyine, 461 
Platacanthomys, 462 
Platanista, 258 
Platanistidee, 257 
Platycercomys, 480 
Platygonus, 291 
Platyonyx, 188 
Platyphoca, 606 
Platypus, 120 
Plecotus, 660 
Plesiadapis, 698 
Plesiarctomys, 457 
Plesictis, 590 
Plesiocetus, 245 
Plesiometacarpalia, 316 
Plesiosorex, 634 
Plesispermophilus, 457 
Pleuraspidotherium, 684 
Pleurolichus, 479 
Plexocherus, 491 
Pliauchenia, 304 
Pliolagostomus, 488 
Pliolophus, 374 
Pliopithecus, 731 
Poébrotherium, 304 
Peecilogale, 590 
Pecilophoca, 605 
Poéphagus, 360 
Pogonodon, 524 
Poiana, 531 
Polecat, 587 
Polymastodon, 118 
Polyprotodontia, 133 
Pontistes, 259 
Pontoporia, 259 
Porcupine, 486 
Tree, 485 


Porpoise, 263 
Potamarchus, 488 
Potamocherus, 286 
Potamogale, 635 
Potamogalide, 634 
Potamophilus, 534 
Potamotherium, 570 
Potoroine, 162 
Potoroo, 163 
Potorous, 163 
Pouched-Rat, 478 
Praopus, 201 
Prehallux, 49 
Prepollex, 49 
Primates, 680 
Priodon, 198 
Prionodon, 5380 
Priscodelphinus, 259 
Proclurus, 523 
Proboscidea, 418 
Probubalus, 361 
Procamelus, 304 
Procapra, 341 
Procavia, 417 
Procoptodon, 170 
Procyon, 564 
Procyonide, 562 
Prodelphinus, 271 
Prodremotherium, 807 
Proechidna, 126 : 
Prohalicore, 223 
Prohyoena, 562 
Prolagostomus, 488 
Promegatherium, 189 
Promylodon, 190 
Prongbuck, 333 
Prophoca, 606 
Propithecus, 684 
Prorastomatide, 224 
Prorastomus, 224 
Protechinomys, 484 
Proteleide, 539 
Proteles, 539 
Proterothertide, 414 
Proterotherium, 414 
Protoadapis, 698 
Protohippus, 380 
Protolabis, 304 
Protoreodon, 293 
Prototheria, 117 
Protoxodon, 440 
Protragelaphus, 349 
Protragoceros, 349 
Proviverra, 608 
Proviverride, 608 
Prox, 317 
Pseudelurus, 523 
Pseudalopex, 552 
Pseudochirus, 151 
Pseudots, 355 
Pseudorca, 268 


762 


INDEX 


Pseudorhinolophus, 657 
Pseudosciurus, 454 
Psittacotherium, 442 
Pteralopex, 654 
Pterodon, 608 
Pteromys, 453 
Pteropodide, 650 
Pieropus, 651 
Ptilocercus, 618 
Ptilodws, 118 
Pudua, 330 

Puma, 520 
Putorius, 585 


Quaaaa, 384 


Rassit, 494 
Bandicoot, 143 
Racoon, 565 
Rangifer, 824 
Rasse, 527 
Rat, 474 
Ratel, 576 
Rat-Kangaroo, 163 
Red Deer, 322 
Rehbok, 339 
Reitbok, 349 
Reproductive organs, 74 
Respiratory system, 63 
Rhabdosteus, 259 
Rhachianectes, 241 
Rhinoceros, 402 
Rhinocerotidee, 402 
Rhinogale, 537 
Rhinolophidee, 656 
Rhinolophus, 656 
Rhinonycteris, 658 
Rhinophylla, 674 
Rhinopithecus, 726 
Rhinopoma, 669 
Rhipidomys, 463 
Rhithrodon, 464 
Rhithrosciurus, 452 
Rhizomys, 477 
Rhizoprion, 257 
Rhogeésa, 661 
Rhynchocyon, 618 
Rhynchonycteris, 667 
Rhytina, 221 
Rhytinide, 221 
Ribs, 44 
River-Hog, 286 
Rock-Wallaby, 167 
Rodentia, 443 
Roe, 327 
Rorqual, 242 
Rosmarus, 597 
Ruminants, 307 
Rupicapra, 349 
Rytiodus, 223 


SABLE, 584 
Saccomys, 479 
Saccopteryx, 667 
Saccostomus, 477 
Sacrum, 43 
Saiga, 341 
Saki, 712 
Salivary glands, 55 
Sambur, 320 
Samotherium, 333 
Sapajou, 717 
Sarcophilus, 137 
Scaldicetus, 251 
Scales, 11 
Scalops, 630 
Scapanus, 630 
Scapteromys, 464 
Scaptochirus, 633 
Scaptonyx, 630 
Scelidotherium, 188 
Schizodelphis, 259 
Schizodon, 482 
Schizostoma, 673 
Sciuravus, 457 
Sciuridc, 450 
Sciurodon, 454 
Sciuroides, 454 
Sciuromorpha, 448 
Sciwropterus, 453 
Sciwrus, 450 
Scopophorus, 339 
Scotophilus, 662 
Scotozous, 661 
Sea-Leopard, 605 
Sea-otter, 571 
Seal, 600 

Eared, 594 
Selenacodon, 113 
Semnopithecus, 726 
Sense organs, 69 
Serow, 351 
Sheep, 354 
Shoulder-girdle, 4 
Shrew, 622 ? 

Tree, 617 

Water, 625 
Siamang, 728 
Siamanga, 728 
Sight, 72 
Sigmodon, 464 
Simia, 731 
Simtidee, 728 
Simocyon, 562 
Simplicidentata, 448 
Siphneus, 472 
Sirenia, 212 
Sivatherium, 322 
Skeleton, 33 
Skull, 34 
Skunk, 572 
Sloth, 180 


Sloth, Ground, 184 
Smell, 72 
Sminthopsis, 139 
Sminthus, 479 
Solenodon, 636 
Solenodontide, 635 
Sorex, 622 
Soricide, 621 
Soriculus, 624 
Sotalia, 272 
Souslik, 456 
Spalacide, 477 
Spalacopus, 482 
Spalacothertwm, 115 
Spalax, 477 
Spantotherium, 294 
Spermophilus, 456 
Sperm Whale, 249 
Spider Monkey, 715 
Spilogale, 574 
Spiny Anteater, 124 
Spleen, 65 
Squalodon, 257 
Squalodontide, 257 
Squamata, 179 
Squirrel, 451 
Stegodon, 427 
Steneofiber, 458 
Steno, 271 
Stenoderma, 676 
Stenoplesictis, 539 
Stenops, 691 
Stenorhynchus, 605 
Stereognathus, 110 
Sternum, 44 
Sthenurus, 170 
Stoat, 590 
Stomach, 57 
Strepsiceros, 847 
Sturnira, 676 
Stylacodon, 114 
Stylinodon, 442 
Styloceros, 317 
Stylodon, 114 
Stypolophus, 608 
Subungulata, 414 
Suide, 281 

Suina, 278 
Suricata, 538 

Sus, 281 
Syllophodus, 484 
Symborodon, 413 
Synaptomys, 467 
Synetheres, 485 
Synotus, 661 
Systemodon, 374 


Taxin, 351 
Talpa, 630 
Talpide, 628 
Tamandua, 192 


INDEX 


763 


Tamias, 452 
Taphozous, 667 
Tapir, 371 
Tapiride, 370 
Tapirulus, 294 
Tapirus, 370 
Tardigrada, 178 
Tarsiide, 694 
Tarsipedine, 148 
Tarsipes, 148 
Tarsius, 694 
Taste, 72 
Tatouay, 198 
Tatusia, 200 
Tatusiine, 200 
Taxidea, 576 
Tayra, 579 
Teetee, 713 

Teeth, 13 
Tegument, 7 
Teledu, 575 
Telemetacarpalia, 323 
Temnocyon, 555 
Tenrec, 637 
Terphone, 338 
Tertiary mammals, 115 
Tetraceros, 338 
Tetraconodon, 292 
Tetracus, 634 
Tetrastylus, 488 
Theridomyide, 484 
Theridomys, 484 
Theropithecus, 722 
Thigh, 51 
Thomomys, 478 
Thoracophorus, 203 
Thylacine, 137 
Thylacinus, 136 
Thylacoleo, 157 
Thymus gland, 66 
Thyroid body, 66 
Thyroptera, 665 
Tiger, 511 
Tillodontia, 441 
Tillotherium, 441 
Tinoceras, 437 
Titanomys, 492 
Titanotheriidee, 413 
Titanotherium, 413 
Tolypeutes, 199 
Tomitherium, 698 


Toxodon, 439 
Toxodontia, 489 
Touch, 72 
Trachea, 67 
Trachyops, 674 
Trachytherium, 224 
Tragelaphus, 346 
Tragoceros, 349 
Tragops, 841 
Tragulide, 305 
Tragulina, 305 
Tragulus, 305 
Trechomys, 484 
Triacanthodon, 113 
Tricenops, 658 
Trichechidee, 596 
Trichechus, 597 
Trichosurus, 150 
Trichys, 487 
Triclis, 162 
Triconodon, 113 
Trilodon, 484 
Trituberculism, 30 
Tritylodon, 111 
Trochictis, 570 
Troglodytes, 736 
Trogontherium, 458 
Trygenycteris, 655 
Tubulidentata, 179 
Tupaia, 617 
Tupatidee, 617 
Tursiops, 271 
Tylopoda, 295 
Tylostoma, 674 
Typhlomys, 477 
Typotherium, 440 


Uacaria, 712 
Uakari, 712 
Uintatheriide, 437 
Uintatherium, 436 
Umbilical vesicle, 77 
Unau, 183 
Ungulata, 273 
Urinary organs, 69 
Urocyon, 553 
Uromys, 476 
Uropsilus, 629 
Urotrichus, 629 
Urside, 557 
Ursus, 557 


THE END 


Urus, 367 
Uses of mammals, 4 
Uterus, 75 


VAMPYRE, 676 
Vampyrus, 673 
Vertebra, 39 
Vesperimus, 463 
Vespertiliavus, 666 
Vespertilio, 663 
Vespertilionide, 660 
Vesperugo, 661 
Vicugna, 300 
Viscacha, 488 
Vishnutherium, 832 
Viverra, 526 
Viverricula, 527 
Viverridee, 525 
Vole, 465 

Vulpes, 552 
Vulpine Phalanger, 150 


WALLABY, 169 
Walrus, 597 
Wapiti, 322 
Wart-Hog, 288 
Weasel, 589 
Whale, 225 
White Whale, 262 
Wolf, 548 
Wolverene, 591 
Wombat, 145 


Xantharpyia, 652 
Xenurus, 198 
Aeromys, 461 
Xerus, 452 
Xiphodon, 294 


YAK, 364 
Yapock, 184 
Yolk-sac, 77 


Zapus, 480 

Zebra, 385 
Zeuglodon, 246 
Zeuglodontide, 246 
Ziphiine, 251 
Ziphius, 254 
Zoological regions, 96 


Printed by R. & R. CLark, Edinburgh 


PORES 


en