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AN  INTRODUCTION 


TO    THE 


STUDY    OF    MAMMALS 


AN   INTBODUCTION 


TO   THE    STUDY    OF 


MAMMALS 


LIVING   AND   EXTINCT 


BY 


WILLIAM  HENEY  FLOWEE 

C.B.,  F.B.8,  D.C.L.,  LL.D.,  P.Z.8,  F.L.8.,  F.G.S.,  &c. 

DIRECTOR  OF  THE  NATURAL  HISTORY  DEPARTMENTS,   BRITISH   MUSEUM 

AND 

EICHAED  LYDEKKEE 

B.A.,  F.G.8.,  F.Z.S.,  &c." 


ILLUSTRATED 


LONDON:    ADAM  AND   CHAELES  BLACK 

MDCCCXCI 


'JUN  3     1958 


PKEFACE 

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  Encyclopaedia  Britannica.  The 
account  of  the  orders  Kodentia,  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,"  "Eat,"  "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,  filling  up  the  gaps,  and  adding  new  matter 
Avhere  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  vii 

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. 

LONDON,  March  1891. 


CORRIGENDA. 

Page  280,  for  Chseropsis  read  Chceropsis. 

Page  292,  for  Chaeropotamidae  and  Chaeropotamus  read  Chceropotamidae  and 
Chceropotanms. 

Page  590,  for  Paecilogale  read  Poecilogale. 


CONTENTS 

CHAPTEE   I 

PAGE 

INTRODUCTORY  REMARKS  .....  1 

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      ....  7 

I.  Tegumentaiy  Structures  ....  7 

Hair,  7  ;  "Colour,  8  ;  Scales,  etc.,  11  ;  Nails,  claws,  and 
hoofs,  12  ;  Odour-secreting  glands,  12. 

II.   Dental  System .  .  .  .  .  .13 

Teeth,  13  ;  Structure  of  teeth,  13  ;  Development  of  teeth, 
15  ;  Forms  of  teeth,  17  ;  Succession  of  teeth,  19  ;  Arrangement, 
homologies,  and  notation  of  teeth,  21  ;  Dental  formulae,  25  ; 
Modifications  of  teeth  in  relation  to  function,  28  ;  Taxonomy, 
30  ;  Trituberculism,  30. 

III.  The  Skeleton    ......          33 

Definition,  33  ;  Axial  skeleton,  34  ;  Skull,  34  ;  Vertebral 
column,  39  ;  Cervical  vertebra,  41  ;  Dorsal  vertebrae,  42 ; 
Lumbar  vertebrae,  42  ;  Sacral  vertebrae,  43  ;  Caudal  vertebrae, 
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  girdle,  50  ;  Thigh  and 
Leg,  51  ;  Pes,  52. 

IV.  The  Digestive  System  .  .  .  .  .53 

General  considerations,  53  ;  Mouth,  54  ;  Salivary  glands, 
55  ;  Stomach,  57  ;  Intestinal  canal,  59  ;  Liver,  60. 

V.   Circulatory,  Absorbent,  Respiratory,  and  Urinary  Systems         63 

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. 


CONTENTS 


PAGE 

VI.   Ni-rvous  System  and  Organs  of  Sense     .  .  .69 

Brain,  69  ;  Nerves,  71  ;  Sense  of  touch,  72  ;  Taste  and 
smell,  7-2  :  Sight,  72  ;  Hearing,  73. 

VII.   Reproductive  Organs     .  .  .  .  .74 

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  .  .         82 

Origin,  82  ;  Classification,  84  ;  Table  of  orders  and 
families,  88. 

CHAPTER   IV 

GEOGRAPHICAL  AND  GEOLOGICAL  DISTRIBUTION  .  .         93 

I.  Geographical  Distribution          .  .  .  .93 

Zoological  regions,  96  ;  Palsearctic  region,  97  ;  Ethiopian 
region,  98;  Oriental  region,  100 ;  Celebes,  102;  Nearctic  region, 
102;  Neotropical  region,  103;  Aquatic  mammals,  104. 

II.   Geological  Distribution  .  .  .  .107 

Sequence  of  strata,  107  ;  Mesozoic  mammals,  109  ;  Multi- 
tuberculata,  109  ;  Polyprotodont  types,  113 ;  Tertiary  mammals, 
115. 

CHAPTER   V 

THE  SUBCLASS  PROTOTHERIA  OR  ORNITHODELPHIA         .  .       117 

General  characters,  117.  Family  ORNITHORHYNCHIDJE, 
119  ;  Ornithorhynchus,  119.  Family  ECHIDNIDJE,  124  ; 
Echidna,  125 ;  Proechidna,  126  ;  Fossil  species,  127. 


CHAPTER   VI 

THE  SUBCLASS  METATHERIA  OR  DIDELPHIA       .  .  .128 

General  characters,  128  ;  Distribution,  131  ;  Classification, 
131. 

Suborder  POLYPROTODONTIA        .  .  133 

Family  DIDELPHYID.E,  133  ;  Chironectes,  134  ;  Didelphys, 
135.  Family  DASYTJRID.E,  136  ;  Subfamily  Dasyurinse,  136  ; 
Thylacinus,  136  ;  Sarcophilus,  137  ;  Dasyurus,  138  ;  Phascolo- 
gale,  139  ;  Sminthopsis,  139  ;  Aiitechinomys,  139  ;  Subfamily 
MyrmecobiiniE,  140;  Myrmecobius,  140.  Family  PERAMELID.E, 
141  ;  Perameles,  142  ;  Peragale,  143  ;  CJueropus,  143. 


CONTENTS 


PAGE 

* H harder  UIPKOTODONTIA  .....        144 

Family  PHASCOLOMYIDA;,  144  ;  Pkascolomys,  145  ;  Phascol- 
onus,  146.  .Fami7i/PHALANGERiD.E,147;  Subfamily  Tarsipedinse, 

148  ;  Tarsipcs,  148  ;  Subfamily  Phalangerinse,  149  ;  Phalanger, 

149  ;  Trichosurus,  150  ;  Pseudoehirus,  151  ;  Petauroides,   152  ; 
Dactylopsila,    152  ;    Petaurus,     153  ;     Gfymnobelideus,     154  ; 
Dromicia,  154  ;  Distcechurus,  155  ;  Acrobates,  155  ;  Subfamily 
Phascolarctinae,  155 ;    Phascolarctus,   156.      EXTINCT    PHAL- 
ANGEROIDS,  157  ;    Thylacoleo,  157.      Family  MACROPOPID-E, 
158  ;  Subfamily  Hypsiprymnodontinae,  162  ;  Hy2>siprymnodon, 
162  ;  7'ricfts,  162  ;  Subfamily  Potoroinse,  162  ;  Potorous,  163  ; 
Bett&ngia,  163  ;  Caloprymnus,  164  ;  jEpyprymnus,   164  ;  /Swfi- 
family  Macropodinse,  164  ;  Lagostrophus,  165  ;    Deiidrolagus, 
165  ;    Dorcopsis,    166  ;    Lagorchestes,    166 ;     Onychogale,    166  ; 
Peiroyale,  167  ;  Macropus,  167  ;  Extinct  genera,  170.    EXTINCT 
FAMILIES,  171  ;  Diprotodon,  171  ;  Nototherium,  171. 

CHAPTER   VII 

THE  SUBCLASS  EUTHERIA  AND  THE  ORDER  EDENTATA  .  .173 

General  characters  and  classification  of  Eutheria,  173. 
ORDER  EDENTATA         .  .  .  .  .  .176 

Family  BRADYPODIIXE,  179  ;  Bradypus,  181  ;  Cholcepus, 
182  ;  Nothropus,  183.  Family  MEGATHERIID^E,  183  ;  Mega- 
therium, 185  ;  Scelidotherium  and  Mylodon,  188  ;  Promega- 
therium,  189.  Family  MYRMECOPHAGID.E,  190 ;  Myrmecophaga, 
190  ;  Tamandua,  192  ;  Cycloturus,  193.  Family  DASYPODID^;, 
194  ;  Subfamily  Chlamydophorinre,  196  ;  Chlamydophorus,  196  ; 
Subfamily  Dasypodinffi,  197  ;  Dasypus,  197  ;  Xcmirus,  198  ; 
Priodon,  198  ;  Tolypeutes,  199  ;  Subfamily  Tatusiiuae,  200  ; 
Tatusia,  200  ;  Extinct  genera,  201.  Family  GLYPTODONTID*, 
202.  Family  MANID^:,  204  ;  Manis,  204  ;  Palceomanis,  208. 
Family  ORYCTEROPODID.E,  208  ;  Orycteropus,  208.  Biblio- 
graphy, 211. 

CHAPTER  VIII 

THE  ORDERS  SIRENIA  AND  CETACEA    .  .  .212 

ORDER  SIRENIA  .  .  .  .  .  .212 

Family  MANATID.S,  215  ;  Manat-us,  215.  Family  HALI- 
CORID^!,  220  ;  Halicore,  220.  Family  RHYTINID^:,  221  ; 
Rhytina,  221.  EXTINCT  SIRENIANS,  222 ;  Halilherium,  222  ; 
Other  forms,  223.  Bibliography,  224. 

ORDER  CETACEA  .  .  .  .  .  .225 

234 

Family  BAL.fiNiD.fi,  234  ;  Balccna,  236  ;  Neobalcena,  241  ; 
Rhachiaiiectes,  241 ;  Megaptera,  241 ;  Balcenoptera,  242 ;  Extinct 
genera,  245. 


CONTENTS 


PAGE 

Suborder  ARCHJEOCETI    . 

Family  ZEUGLODONTIDJS,  246  ;  Zeuglodon,  246. 

Suborder  ODONTOCETI    .  .  •  •  .247 

Family  PHYSETERID.E,  247  ;  Subfamily  Physeteriuse,  248  ; 
Physcter,  248  ;  Coffia,  250  ;  Extinct  physeteroids,  251  ;  Knit- 
family  Ziphiinse,  251  ;  Hyperoodon,  252  ;  ZipUus,  254  ;  Mcso- 
plodon,  254;  Berardius,  256;  Choneziphius,  257.  Famif;/ 
SQUALODONTID.E,  257 ;  Squalodon,  257.  Family  PLATANIS-TID^E, 
257  ;  Platanista,  258  ;  Inia,  259 ;  Pontoporia,  259  ;  Fossil  forms, 
259.  Family DELPHINIUM,  260;  Monodon,26Q;  Delphinapterus, 
262  ;  Plwccena,  263  ;  Cephalorhynchus,  266  ;  Orcella,  267 ;  Orca, 
267  ;  Pseudorca,  268  ;  Globicephalus,  268  ;  Grampus,  270  ; 
Feresia,  270  ;  Lagenorhynchus,  270  ;  Delphinus,  271  ;  Tursiops, 
271 ;  Prodclphinus,  271  ;  tffewo,  271  ;  Sotaliq,  272.  Biblio- 
graphy, 272. 

CHAPTEK  IX 

THE  ORDER  UNGULATA                        .  273 

UNGULATA  VERA          .  .275 

Suborder  ARTIODACTYLA               .              .  .                            .275 

SUINA,  278.  Family  HIPPOPOTAMID^:,  278  ;  Hippopotamus, 
278.  Family  SUID.E,  281  ;  Sus,  281  ;  BaUrusa,  287  ;  Phaco- 
choRrus,  288.  Family  DICOTYLID^E,  289 ;  Dicotyles,  289  ; 
ffyotherium,  etc.,  291 .  EXT-INCT  TRANSITIONAL  ARTIODACTYLES, 
292  ;  Chreropotamidse,  292  ;  Anthracotheriidfe,  292  ;  Meryco- 
potamus,  293  ;  Cotylopidse,  293  ;  Anoplotheriidse,  293  ;  Caeno- 
theriidse,  294  ;  Dichodontidse,  294.  TYLOPODA,  295.  Family 
CAMELID.E,  295 ;  Camclus,  296 ;  Auchenia,  298 ;  Extinct 
Cameloids,  303.  TRAGULINA,  305.  Family  TRAGULID^:,  305  ; 
Traguhis,  305  ;  Dorcatherium,  306  ;  Extinct  Traguloids,  306. 
PECORA,  307j  Antlers,  308;  Horns,  310;  Teeth,  310;  Stomach, 
312.  Family  CERVID.S:,  313 ;  Subfamily  Moschinse,  314  ; 
Moschus,  314  ;  Subfamily  Cerviuse,  316  ;  Plesiometacarpalia, 
316  ;  Cervulus,  316  ;  Elaphodus,  318  ;  Certms,  319  ;  Telemeta- 
carpalia,  323  ;  Rangifer,  324  ;  Alces,  326  ;  Cervalces,  327  ; 
Caprcolus,  327  ;  Hydropotes,  328  ;  Cariacus,  329  ;  Pudua,  330  ; 
Extinct  genera,  330.  Family  GIRAFFIDJE,  330  ;  Giraffa,  331  ; 
Allied  extinct  types,  332.  Family  ANTILOCAPRID^E,  333  ; 
Antilocapra,  333.  Family  BOVID^E,  334  ;  Alcelaphus,  334 ; 
Connochoctes,  336  ;  CepJialophus,  338  ;  Tetraceros,  338  ;  Neo- 
tragus,  338  ;  Naiwtragus,  339  ;  Pelea,  339  ;  Cobus,  339  ;  Cervi- 
capra,  340  ;  Antilope,  340 ;  ^Epyceros,  341  ;  Saiga,  341  ; 
Paniholops,  341 ;  Gazella,  341  ;  Hippotragus,  343  ;  Oryx,  343  ; 
Addax,  345  ;  Boselaphus,  345  ;  Tragelaphus,  346  ;  Strepsiceros, 
347  ;  Oreas,  348 ;  Extinct  types,  348  ;  Eupicapra,  349  ;  Nemo- 
rhcedus,  350  ;  Haploceros,  351  ;  Budorcas,  351  ;  Capra,  352  ; 
Ovis,  354  ;  Ovibos,  357  ;  Bos,  360. 


CONTENTS 


PAGE 

Suborder  PERISSODACTYLA  .  .  .  .  .368 

Family  TAPIRID.E,  370  ;  Tapirus,  370  ;  Palceotapirus,  373. 
Family  LOPHIODONTID^,  373.  Family  PAL^OTHERIID^E,  375. 
Family  EQUID.E,  376  ;  Protohippus,  380  ;  Hipparion,  380  ; 
Equus,  381.  Family  RHIXOCEROTID.E,  402  ;  Rhinoceros,  402  ; 
Extinct  types,  411.  Families  LAMBDOTHERIID.E,  CHALICO- 
THERIID.S:,  and  TITANOTHERIID^E,  412.  Family  MACRAU- 
CHENIID.E,  414.  Family  PROTEROTHEIIIIT)^,  414. 

SUBUNGDLATA     .  .  .  .  .  .  .414 

Suborder  HYRACOIDEA     .  .  .  .  .  .415 

Family  HYKACIDJE,  415  ;  Hyrax,  417  ;  Dendrohyrax,  418. 

Suborder  PROBOSCIDEA    .  .  .  .  .  .418 

Family  ELEPHANTID.E,  423  ;  Elephas,  424  ;  Mastodon,  431. 
Family  DINOTHERIID.E,  435  ;  Dinotherium,  435. 

Suborder  AMBLYPODA      ......  436 

Uintatherium,  436  ;  Coryphodon,  437. 

Suborder  CONDYLARTHRA             .....  438 

Suborder  TOXODONTIA     ......  439 

Ncsodon,  439  ;  Toxodon,  439  ;  Typotherium,  440. 

Group  TILLODONTIA         ......  441 

Bibliography  of  Ungulates  .  .  .  .  .442 

CHAPTER   X 

THE  ORDER  RODENTIA   ......       443 

Suborder  SIMPLICIDENTATA  ...  .  .  .       448 

Section  SCIUROMORPHA,  448.  Family  ANOMALURID.E,  449  ; 
Anomalurus,  449.  Family  SCIURID.S:,  450  ;  Sciurus,  450  ; 
Rhithrosciurus,  452  ;  Xerus,  452  ;  Tamias,  452  ;  Pteromys  and 
Sciuroplerus,  453  ;  Eupetaurus,  454 ;  Extinct  genera,  454  ; 
Arclomys,  454  ;  Cynomys,  455  ;  Spermophilus,  456  ;  Extinct 
genera,  457-  Family  HAPLODONTID*:,  457  ;  Haplodon,  457. 
Family  CASTORID^:,  457  ;  pastor,  457.  Section  MYOMORPHA, 
459.  Family  MYOXID^E,  459  ;  Myoxus,  459  ;  Eliomys,  459  ; 
Graphiurus,  459  ;  Claviglis,  460  ;  Muscardinus,  460.  Family 
LOPHIOMYID.E,  460  ;  Lophiomys,  460.  Family  MURID^;,  461  ; 
Hydromys,  461 ;  Xeromys,  461 ;  Platacanthomys,  462 ;  Gcrbilltts, 
462  ;  Pachyuromys,  462 ;  Mystromys,  462  ;  Otomys  and  Dasymys, 
462 ;  Malacomys,  462 ;  Phlceomys,  462  ;  Dendromys,  463  ; 
Cricetus,  463  ;  Holochilus,  464 ;  Sigmodon,  464  ;  Rhitltrodon 
and  Odwtodon,  464  ;  Neotoma,  464  ;  Jfypogcomys,  465  ;  Nesomys, 
465  ;  Brachytarsomys,  465  ;  Hallomys,  465  ;  Eliurus,  465  ; 
Phenacomys,  466  ;  Arvicola,  466  ;  Synaptomys,  467  ;  Mi/odes, 
467  ;  Cuniculus,  470  ;  Fiber,  470  ;  Neofiber,  472  ;  Ellobius, 


CONTENTS 


472;  Siphnfux,   472;  Deomys,  473;  Mus,  473:  !75  ; 

Oolunda,  476 ;  Uromys,  476 ;  Chincromys,  476  ;  Hapalotis, 
476  ;  Mastacomys,  476  ;  Acanthomys,  476  ;  Echinothrix,  477  : 
Typklomys,  477  ;  Cricetomys  and  Saccostomus,  477  ;  Pithechirus, 
477.  Family  SPALACIDJE,  477  ;  Spalax,  477  ;  Ehizomys,  477  ; 
Bathyergus,  478 ;  Georychus  and  Myoscalops,  478 ;  Hetero- 
cephalus,  478.  Family  GEOHYID^E,  478 ;  Geomys,  478  ; 
Thonwmys,  478  ;  Dipodomys,  479  ;  Perognathus  and  Hetcromys, 
479.  Family  DIPODID^:,  479  ;  Sminthus,  479  ;  Zapus,  •  480  ; 
Z>i/ws,  480  ;  Alactaga,  480  ;  Platycercomys,  480  ;  Pedetes,  480. 
Section  HYSTRICOMORPHA,  480.  Family  OCTODOXTID^E,  480. 
Ctenodactylus,  481  ;  Pectinator,  481  ;  Octodon,  481  ;  Habrocoma, 
482 ;  Schizodon,  482  ;  Ctenomys,  482 ;  Spalacopus,  482  ; 
Petromys,  482  ;  Afyopotamus,  482  ;  Capromys,  482  ;  Aulacodus, 
483  ;  Plagiodon,  483  ;  Lonchcres  and  Echinomys,  483  ;  Mesomys, 
483 ;  Dactylomys,  483  ;  Cercomys,  483  ;  Carterodon,  484  ; 
Fossil  forms,  484.  Family  THERIDOMYID.E,  484.  Family 
HYSTRICID.E,  484  ;  Erethizon,  484  ;  Synetheres,  485  ;  CJuetamys, 
486  ;  Hystrix,  486  ;  Atherura,  487  ;  Trichys,  487.  Family 
CHINCHILLID.E,  487  ;  Chinchilla,  487  ;  Lagidium  and  Lagosto- 
mus,  488  ;  Extinct  genera,  488.  Family  CASTOROIDIDJ;,  488  ; 
Castoraides,  488.  Family  DASYPROCTID^,  488  ;  Dasyprocta, 
488  ;  Ccelogenys,  489.  Family  DIXOMYID^E,  489 ;  Dinomys,  489. 
Family  CAVIID^B,  489  ;  Cltma,  489  ;  Dolichotis,  490  ;  Hydro- 
choerus,  490  ;  Extinct  genera,  491. 

Suborder  DUPLICIDENTATA  .  .  .  .  .491 

Family  LAGOMYID.E,  491  ;  Lagmnys,  491.     Family  LEPO- 
RID.E,  492  ;  JSejnts,  492. 


CHAPTER   XI 

THE  ORDER  CARNIVORA  .  .  .  .       496 

Suborder  CARXIVORA  VERA         .  .  .  .  .497 

Section  ^ELTTROIDEA,  501.  Family  FELID.E,  502 ;  /'e^is, 
502  ;  Cyncelurus,  523  ;  Extinct  genera,  523.  Family  VIVER- 
RID^E,  525  ;  Cryptoprocta,  525  ;  Viverra,  526  ;  /bssa,  527  ; 
Genelta,  528  ;  Prionodon,  530  ;  Poiana,  531  ;  Paradoxurus, 
532  ;  Arctogalc,  533 ;  Hemigale,  533  ;  Arctictis,  534  ;  Nandinia, 
534  ;  Cynogale,  534  ;  Herpestes,  535  ;  Helogale,  537  ;  Bdeogale, 
537  ;  Cynictis,  537 ;  Rhinogale,  537  ;  Crossarchus,  537  ;  Suricata, 
538 ;  Galidictis,  Galidea,  and  Hemigalidea,  538 ;  JSupleres, 
538 ;  Extinct  genera,  539.  Family  PROTELEID.E,  539  ;  Proteles, 
539.  Family  HY^NID^:,  540 ;  Hyaena,  540.  Section  CYNOIDEA, 
544.  Family  CANIDJE,  544 ;  Cams,  546 ;  Lycaon,  553 ;  Icticyon, 
553;  Otocyon,  554;  Extinct  genera,  555.  £ecW0«ARCTon>EA,  556. 
Family  URSIDJ:,  557  ;  tfams,  557  ;  Melursus,  560  ;  ^Eluropus, 
560 ;  Extinct  genera,  561.  Family  PROCYONID.E,  562  ; 
dUlurus,  562  ;  Procyon,  564  ;  Bassaris,  566  ;  Bassaricyon,  566  ; 
Nasua,  566  ;  Cercoleptcs,  567.  Family  MUSTELID.E,  567  ; 


CONTENTS 


PAGE 

Lutra,  567  ;  Extinct  Otters,  570  ;  Latax,  570  ;  Mephitis,  572  ; 
Conepatus,  574  ;  Arctonyx,  574  ;  Mydaus,  575  ;  Meles,  575  ; 
Taxidea,  576  ;  Mellivara,  576  ;  Helictis,  578  ;  Ictonyx,  579  ; 
Galidis,  579  ;  Muslela,  579  ;  Extinct  Mustelines,  590  ;  Pcecilo- 
gale,  590  ;  Lyncodon,  590  ;  GWo,  591. 

Suborder  PINNIPEDIA      ......        592 

Family  OTAKIID^;,  593  ;  Otaria,  593.  Family  TRICHE- 
CHiDjE,  596 ;  Trichechus,  597.  Family  PHOCID^:,  600  ; 
Halichcerus,  601  ;  Phoca,  601  ;  Moiuichus,  604  ;  Ogmorhinus, 
605 ;  Lobodon,  605  ;  Pcecilophoca,  605 ;  Ommatophoca,  605  ; 
Cystophora,  605  ;  Macrorhimis,  606  ;  Extinct  seals,  606 

Suborder  CREODONTA  .....       606 

Hyaenodontidse,  608  ;  Proviverridae,  608  ;  Aretocyonidae  and 
Mesonychidse,  609. 

CHAPTEE   XII 

THE  ORDER  IXSECTIVORA          .  .  .  .  .610 

Suborder  DERMOPTERA  .  .  .  .  .  .614 

Family  GALEOPITHECID^E,  614  ;  Galeojnthecus,  614. 
Suborder  INSECTIVORA  VERA      .  .  .  .  .616 

Family  TUPAIID.S:,  617  ;  Tupaia,  617  ;  Ptilocercus,  618  ;  Ex- 
tinct genera,  618.  Family  MACROSCELIDID.E,  618  ;  Macroscel- 
idfs,  618 ;  Rhynchocyon,  618.  Family  ERINACEID.E,  619  ; 
Gymnura,  619  ;  Erinaceus,  620  ;  Extinct  genera,  621  ;  Family 
SORICIDJ3,  621  ;  Sorex,  622  ;  Soriculus,  624  ;  Notiosorex,  624  ; 
Marina,  624  ;  Crossopus,  625  ;  Myosorex,  625  ;  Crocidura,  626  ; 
Diplomesodon,  626;  Anurosorex,  626;  Chimarrogale,  626;  Necto- 
gale,  627  ;  Fossil  Soricidse,.  627.  Family  TALPID*:,  628  ; 
Myogale,  628  ;  Urotrichus,  629  ;  Uropsilus,  629  ;  Scalops,  630  : 
Scapanus,  630  ;  Condylura,  630  ;  Scaptonyx,  630  ;  Talpa,  630  ; 
Extinct  genera,  634.  Family  ADAPISORICID.E,  634.  Family 
POTAMOGALID.E,  634  ;  Potamogale,  635 ;  Geogale,  635.  Family 
SOLENODONTID.E,  635  ;  Solenodon,  636  ;  Centetes,  637  ;  Hemi- 
centetes,  637  ;  Ericulus,  638  ;  Microgale,  638  ;  Oryzorictes,  638  ; 
Chrysochloris,  639.  EXTINCT  TYPES,  640.  Bibliography,  640. 

CHAPTER   XIII 

THE  ORDER  CHIROPTERA          .  .  .  .  .641 

Suborder  MEGACHIROPTERA         ...  .       650 

Family  PTEROPODID.E,  650  ;  Epomoplwrus,  650  ;  Pteropus, 
651  ;  Xantharpyia,  652 ;  Boncia,  653  ;  Cynoptcrus,  653  : 
Harpyia,  653  ;  Cephalotes,  653  ;  Pteralopex,  654  ;  Notopteris, 
654  ;  Eonycteris,  654  ;  Carpmiycteris  and  Melonyctcris,  654  ; 
Xesonycteris,  655  ;  Callinycteris,  655  ;  Trygcnyctcris,  655. 


CONTENTS 


PAGE 

Suborder  MICROCHIROPTERA        .  •        655 

Section  VESPERTILIONINA,  655.  Family  RHINOLOPHID^E, 
656  ;  JRhinolophus,  656  ;  Hipposiderus,  657  ;  Anthops,  657  ; 
Uli  iiionyctcris  and  Tricenops,  658  ;  Ccelops,  658  ;  Megaderma, 
658.  Family  VESPERTILIONIDJE,  660  ;  Plccotus,  660  ;  Synotus, 
661  ;  Otonycteris,  661  ;  Nyctophilus,  661  ;  Antrozous,  661  ; 
Vesperugo,  661  ;  Chalinolobus,  662  ;  Scotophilus,  662  ;  Nydi<;  - 
jus,  663  ;  Atalapha,  663  ;  Harpyiocephalus,  663  ;  Vespertilio, 
663  ;  Cerivoula,  664  ;  Natalus,  664  ;  Minioptcrus,  664  ;  Thyrop- 
tera,  665 ;  Myxopoda,  665  ;  Fossil  Vespertilionidse,  665. 
Section  EMBALLONURINA,  666.  Family  EMBALLONURID^:,  666 ; 
Furiptcrus  and  Amorphochihis,666;  Emballonura,  667;  Coleura, 
667  ;  Rhynchonycteris,  667  ;  Saccopteryx,  667  ;  Taphozous,  667  ; 
IHclidurus,  668  ;  Nodilio,  668  ;  Khinopoma,  669  ;  Chiromeles, 
669  ;  Molossus,  670  ;  Nyctinomm,  670  ;  Mystacops,  671.  Family 
PHYLLOSTOMATID^E,  672  ;  Chilonycteris,  672  ;  Alormops,  672  ; 
Lonchorhina,  Otopterus,  and  Dolichophyllum,  673  ;  Vampyrus, 
etc.,  673  ;  Desmodus,  677  ;  Diphylla,  678. 


CHAPTEE   XIV 

THE  ORDER  PRIMATES  ......       680 

Suborder  LEMUROIDEA    .  .  .  .  .  .682 

Family  LEMURIDJ;,  683  ;  Indris,  684  ;  Propithecus,  684  ; 
Avahis,  686 ;  Lemur,  687 ;  Hapalemur,  689 ;  Lepidolemur, 
689  ;  Chirogaleus,  689  ;  Galago,  690  ;  Nycticebus,  691  ;  ioris, 
692 ;  Perodicticus,  693.  Family  TARSIID.E,  694 ;  Tarsius, 
694.  Family  CHIROMYIDJS,  694  ;  Chiromys,  695.  EXTINCT 
LEMUROIDS,  696. 

Suborder  ANTHROPOIDEA   .  .  .  .  .         .       699 

Family  HAPALIDJE,  709;  Hapale,  710;  Midas,  710.  Family 
CEBIDJE,  711  ;  Mycetcs,  711  ;  Pitheda,  712 ;  Uacaria,  712 ; 
Callithrix,  713  ;  C'hrysothrix,  714  ;  Nydipithecus,  714  ;  Ateles, 
715  ;  Eriodes,  715 ;  Lagothrix,  716 ;  Ceitts,  717.  Family 
CERCOPITHECID.E,  718  ;  Cynocephalus,  719  ;  Theropithecus,  722  ; 
Cynopithecus,  722 ;  Macaciis,  722 ;  Cercocebus,  723 ;  Cerco- 
jiif/tccus,  724 ;  Nasalis,  725  ;  Semnopithecus,  726 ;  Colobus, 

727  ;  Extinct  genera,  727.     Family  SIMIID.E,  728  ;  Hylobates, 

728  ;  Simia,  731 ;  Gorilla,  734  ;  Aitihropopithecus,  736.     Family 
HOMIXIDJE,  739  ;  Homo,  740.     Classification  of  the  varieties  of 
.Man,  743. 


AN   INTRODUCTION 

TO 

THE   STUDY  OF  MAMMALS 

LIVING  AND   EXTINCT 
CHAPTER   I 

INTRODUCTORY  REMARKS 

MAMMALIA  (French,  Mammifkres;  German,  Sdugethiere)  is  the  name 
invented  by  Linnaeus  (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 


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  Palaeozoic  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  branchiae  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 


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 


INTRODUCTORY 


protection  the  mother,  in  a  large  number  of  cases,  has  a  special 
pouch  enclosing  the  mammae,  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  fcetal  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  Kodents,  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  Eorqual, 
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 


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  Keindeer  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  Eeindeer.  The  Kodents,  as  represented  by  Hares  and  Kabbits, 
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 


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  Hyperoodon  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. 


CHAPTEE   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  (Cholcepus)  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  vibiissce  or 
whiskers,  and  endowed,  through  the  abundant  nerve  supply  of  their 
basal  papilla?,  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  (CJirysochloris),  the  Desmans  or  Water  Moles  (Myogale), 
and  Potamogale  velox,  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  (Brachyurm)  has  its  whole  face  of 


TEG  UMENTA  RY  STR  UC TURKS 


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  (Diclidurus  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,  dSlurus, 
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 


io  GENERAL  ANATOMICAL  CHARACTERS 

albinism  :i  rondition  in  \vhi<li  the  pigment  <>r  colouring  matter 
<if  in  the  tissues  constituting  the  external  cuvrrings  of 
I!M'  ho.lv,  :HM!  which  gives  them  their  characteristic  hue,  is  absent. 
When  it,  occurs  the  hair  is  of  :m  opaque  white,  the  daws,  hoofs,  etc.,  of 
n  |i.i|c  liMi  ii  . •,.|.)iu-,  :ind  the  skin  and  eyes  pink,  in  con>e(|uenee  of  the 
colour  of  tin-  hi ood  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 
illy  as  an  individual  peculiarity  among  wild  animals  of 
many  Kimls ;  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  :•  -  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  Due,  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  u 

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  moonlight1  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 
Manidce  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,  Eats,  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.  ]87. 


i:  GENERAL  ANATOMICAL   CHARACTERS 

Equidtc,  the  grasping  under  surface  of  the  tail  of  the  prehensile-tailed 

keys,  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  Hoof$. — 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,  Ornith&rhynchvs,  Proeehidna,  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  Taphozaus  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  Saceopteryx,  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. 


II.    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.1 

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  sul>- 
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 

1  L.  F.  E.  Rousseau,  Anatomic  comparie  du  Systeme  dentaire  chez  THommt  et 
chez  les principaux  Animaux,  2d  ed.,  1839  ;  F.  Cuvier,  Des  Dents  des  Mammiferts 
considertes  comme  caracteres  zooloyiques,  1822-25  ;  R.  Owen,  Odontography, 
1840-45  ;  C.  G.  Giebel,  Odontographie,  1855  ;  C.  S.  Tomes,  Manual  of  Dental 
Anatomy,  Human  and  Comparative,  3d  ed.,  1889. 


14  GENERAL  ANATOMICAL   CHARACTERS 

gums  or  fibre-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.1  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.2 

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. 

2  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  Megatherium,  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  dis^in- 
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  lacunae  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 


1 6  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 
thi|f  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,  I.)  It  is  a 
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 

TV  2 


i8 


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 

the   pulp -cavity,   which    is 
very    different 
widely     open 


from     the 
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,  IV.)  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 

Fio.  1.— Diagrammatic  Sections  of  various  forms  of    SO    in   old  age,  it  being  Only 

_.3th.    i.  incisor  or  tusk  of  Elephant,  with  pulp-  in  consequence  of  absorption 

cavity  persistently  open  at  base.     II.  Human  incisor         d  retrocession  of    tne   SUr- 
during  development,  with   root  imperfectly  formed,  . 


Teeth, 


a  smaii  aperture  at  tne  end  01  tne  root.    iv.  Human  r 

molar,  with  broad  crown  and  two  roots.    V.  Molar  of  S 

the  Ox,  with  the  enamel  covering  the  crown  deeply  tication,  it  is  never  renewed. 

folded,  and  the  depressions  filled  up  with  cement.    The  T'he  Ot)6n  cavitv  at  the  base 

surface  is  worn  by  use ;  otherwise  the  enamel  coating       <•    i       •  £     ±1     J        i         J 

would  be  continuous  at  the  top  of  the  ridges,   in  all  of  the  imperfectly  developed 

the  figures  the  enamel  is  black,  the  pulp  white,  the  tooth  (Fig.  1,  II.)  CaUSBS  it 
dentine  represented  by  horizontal  lines,  and  the  cement  tQ  resemble  tne  persistent 

condition    of    the    rootless 

tooth.  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  hit, 
German  milchzahne)  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.1 

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.  iii.  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  Dasyuridse,"  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  (Monodori),  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  correspondingdepressions 
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 
palaeontology  fully  confirm  this  view,  as  by  tracing  back  many 
groups  now  widely  separated  in  dental  characters  we  find  a 
gradual  approximation  to  a  common  type.  In  this  generalised  form 
of  mammalian  dentition  (which  is  best  exemplified  in  the  genera 
Anoplotherium  and  Homalodontotherium)  the  entire  number  of  teeth 
present  is  44,  or  11  above  and  1 1  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  premaxilla, 
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  a  rule  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  Cot //lops,  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  formulce. — 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 : — 

.     .  3-3  •          1-1  i         4-4  3-3      11-11 

incisors   -  — ,   canines  - — -,   prernolars   -  — ,   molars    -  —  =  — — —  : 

o  —  o  X  —  i.  4  —  4  o  —  o       11  —  11 

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 — 

*  f  >  c  T.  P  I,  m  f  =  TT  ;  total  44- 

The  individual  teeth  of  each  group  are  always  enumerated  from 
before  backwards,  and  by  such  a  formula  as  the  following — 
i  1,  i  2,  i  3,  c,  p  1,  p  2,  p  3,  p  4,  m  1,  m  2,  m  3 
i  1,  i  2,  i  3,  c,  p  1,  p  2,  p  3,  p  4,  m  1,  m  2,  m  3 
or  more  briefly — 

.  1,  2,  3        l_        1,  2,  3,  4         1,  2,  3 
1  1T2T3'  C  I'  P  1,  2,  3,  4'  m  17273' 

A  special  numerical  designation  is  thus  given  by  which  each  one 
can  be  indicated.  In  mentioning  any  single  tooth,  such  a  sign  as  — 
will  mean  the  first  upper  molar,  ^l  the  first  lower  molar,  and  so  on. 
The  use  of  such  signs  saves  much  time  and  space  in  description.1 

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,  Ch/mnura,  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  i  1 ;  when  two,  they  are  i  1  and  i  2.  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  /,  0,  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 


771.2, 


m.  3 


PIG.  3.— Milk  and  Permanent  Dentition  of  Upper  (I.)  and  Lower  (II.)  Jaw  of  the  Dog  (Canis 
familiaris),  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  f,  dc  \,  dm  -f,  =  -f,  total  28.  In  a  few 
Ungulates,  however,  such  as  the  Hyrax  and  Tapir,  and  in  some 
instances  the  Rhinoceros  and  the  extinct  Palceotherium,  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,1  and  Mr.  0.  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  i^ 

m  ~~>  ky  the  l°ss  °f  the  fifth  lower  incisor, 


i'      i  2  3  4'      i  2  3 
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          1,  0,  3,  4  1,  2,  3,  4  .,        ,  ,         ,     ., 

*  i,  2,  3,  4,  o>  c  I'  P  170737!'   m  ITvsTV  or  the  formula  of   the 
Opossum  (Didelphys),  usually  written  i  J,  c^,p  ^,  m  £.      Again, 
in  the  same  formula  the  loss  of  the  fourth  and  fifth  incisors  in 

12300        1 
both  jaws,  and  also  of  the  fourth  molars,  gives  us  i  1'  2'  3'  Q'  Q>  c  -, 

P  !'  »'  Q'  !>  m.  r^1!.  or  the  formula  of  a  typical  Eutherian,  like  the 

I  .    —  .    -'.   4  lj  4)    O 

1  According  to  Mr.  G.  E.  Dobson  there  are  four  upper  incisors  in  some  of 
the  Soriddce. 


28  GENERAL   ANATOMICAL   CHARACTERS 

Pig,  which  we  generally  write  as  i  f ,  c  £,  p  £,  m  f .  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  Ornithwhynchus,  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  Felidce.  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  Eodents. 

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  Eodents,  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. 

Tritubercidism. — 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  tritubercular  type  j 
these  three  cusps  being  arranged  in  a  triangle,  with  the  apex 
directed  inwardly  in  the  upper  teeth  (Fig.  4,  e),  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 


3> 


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 


FIG.  4. — Molar  teeth  of  Mesozoic  Mammals  (enlarged).  Triconodont  type — 1,  Dromatherium ; 
2,  Microconodon  ;  3,  Amphilestes ;  4,  Phascolotherium ;  5,  Triconodon.  Tritubercular  type — 6,  7, 
Spalacotkerium ;  10,  Asthenodon.  Tubercular  sectorial  type — 8,  Amphitherium  ;  9,  Peramiw;  11- 
13,  Amblotherium ;  14  (?)  Amblotherium.  pr,  Protocoue  ;  hy,  hypocone ;  pa,  paracone ;  me, 
metacone,  in  the  upper  teeth ;  and  protoconid,  hypoconid,  paraconid,  and  metaeonid  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,  i)  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,  e)  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,  e  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'  bv  the  SUP' 
pression  of  one  of  the 

is  j  •*• 

Itte          'j  \L    /\  three  primitive  cusps,  it 

"  develops  into  the  quadri- 

•Fio.  5.— Diagram  of  two  upper  and  two  lower  left  tubercular    type     of     the 

quadritubercular  molars  in  mutual  apposition.    The  cusps  Vni  n  nH  nn  t  «Ari  PO 
and  ridges  of  the  upper  molars  in  double  lines,  and  those 

of  the  lower  in  black  lines.    The  lower  molars  are  looked  In    the   Upper  molars 

at  from  below,  as  if  transparent,    pr,  Protocone ;  hy,  hypo-  the   primitive  triangle  in 

cone  ;  pa,  paracone  ;  me,  metacone  ;  ml,  protoconule  •  pi  ,  i 

metaconule  ;  prd,  protoconid ;  hyd,  hypoconid  ;  pad,  para-  tjle    s6COdont    Series  may 

conid  ;  med,  metaconid  ;  end,  entoconid.    (After  Osborn.)  remain  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,  pi).  Finally,  in  the  bunodont  series,  the  addition  of  a  postero- 
internal  cusp  (Fig.  5,  A?/),  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  MOLARS. 

Antero-internal  cusp  =  protocone     =pr. 

Postero      „        or  6th  cusp  =  hypocone      =  hy. 

Antero-external  cusp  =paracone       =pa. 

Postero      ,,          „  =metacone      =me. 

Anterior  intermediate  cusp  =  protoconule  =  ml. 

Posterior         „  „  =metaconule  =pl. 

LOWER  MOLARS. 

Antero-external  cusp  =  protoconicl  =prd. 

Postero       „  „  =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. 


III.    THE   SKELETON. 

Definition. — The  skeleton  is  a  system  of  hard  parts,  forming  a 
framework  Avhich  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  lacunae, 
connected  together  by  delicate  channels  or  canaliculi,  which  radiate 
in  all  directions  from  the  sides  of  the  lacunae.  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.1 

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  Osttographie  of  De  Blainville,  1835-54  ;  the  section 
devoted  to  the  subject  in  Bronn's  Klassen  und  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 


fa       IP 


CE 


ACE 


No- 


FIG.  6. — Longitudinal  and  vertical  section  of  the  skull  of  a  Dog  (Canis  familiaris),  with 
mandible  and  hyoid  arch,  an,  Anterior  narial  aperture ;  M T,  maxillo-turbinal  bone  ;  ET,  ethmo- 
turbinal ;  Na,  nasal ;  ME,  ossified  portion  of  the  mesethmoid ;  CE,  cribriform  plate  of  the 
ethmo-turbinal :  Fr,  frontal ;  Pa,  parietal ;  IP,  interparietal ;  SO,  supraoccipital ;  ExO,  ex- 
occipital  ;  BO,  bastoccipital ;  Per,  periotic ;  BS,  basisphenoid ;  Pt,  pterygoid ;  AS,  all- 
sphenoid  ;  OS,  orbitosphenoid  ;  PS,  presphenoid  ;  PI,  palatine  ;  VO,  vomer ;  MX,  maxilla  ; 
PMx,  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  fossae,  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  premaxillae  (Figs.  6,  7,  PMx),  which  carry  the 
incisors ;  behind  them  are  the  maxillae,  in  which  all  the  remaining 
upper  teeth  are  implanted.  Both  the  premaxillae  and  maxillae  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 


Per 


between  the  premaxilla  and  maxilla  of  either  side.  Behind  the  nasals 
and  maxillae,  the  anterior  part  of  the  brain-case  is  formed  by  the 
large  paired  frontals  (Figs.  6,  7,  Fr),  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- 

1  /TT'      *7    <y  \  (  •  '•~Side  view  of  skull  of  Cape  Jumping  Hare  (Pedetes 

OSal(rig.  <,0</;  lorms     ca/er).      x?.    PMx,  Premaxilla;  MX,  maxilla;   Ma,  jugal  or 

the     lateral     Wall     of     malar ;  Fr,  frontal ;   L,  lachrymal ;   Pa,  parietal ;  Na,  nasal ; 

the     hinder    part     Of     S9>  S(luamosal  J  r2/>  tympanic  ;  ExO,  exoccipital ;  AS,  alisphen- 

oid  ;  OS,  orbitosphenoid  ;  Per,  mastoid  bulla. 

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  affinity. 
The  relative  position  occupied  by  the  orbito-  and  alisphenoid  is 
sufficiently  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 


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  Mam- 
malia already  men- 
tioned. Attention 

FIG.  8.— The  right  half  of  the  hinder  part  of  the  base  of  the  r  i 

cranium  of  the  Wolf  (Canis  lupus),    c,  Condyloid  foramen  ;  I,  fora-  mav>  >  -QOWCVer,     D6 

men  lacerum  posticum  ;  car,  carotid  canal;  e,  eustachian  canal;  particularly     dl- 

o,  foramen  ovale ;  a,  posterior,  and  a',  anterior  aperture  of  ali-  rected  to  the  SO- 
sphenoid  canal ;  P,  paroccipital  process  of  exoccipital ;  m,  mastoid  ,,  ,  ,.  ., 

process  of  periotic  ;  am,  external  auditory  meatus ;  g,  glenoid  for-  Called      alispnenoid 

amen,  below  which  is  the  glenoid  cavity  for  the  condyle  of  the  man-  canal,    the   position 

dible.    (Flower,  Proc.  Zool.  Soc.,  1869,  p.  25.)  Qf   which  jg    gh()wn 

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  vertebrae,  arranged  in  close  connection  with 
each  other  along  the  dorsal  side  of  the  neck  and  trunk,  and  in  the 
median  line.1  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  Cholcepus).  The  different  vertebrae,  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. 


GENERAL  ANATOMICAL   CHARACTERS 


of  the  trunk,  in  nearly  all  mammals  which  possess  completely 
developed  hinder  limbs,  two  or  more  vertebrae  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  vertebrae  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,  c),  of  the  form  of  a  disk  or  short  cylinder.  The  bodies  of  con- 


FIG.  9.— Anterior  surface  of  Human 
thoracic  vertebra  (fourth),  c,  Body  or 
centrum ;  «c,  neural  canal ;  p,  pedicle, 
and  /,  lamina  of  the  arch ;  t,  transverse 
process  ;  az,  anterior  zygapophysis. 


FIG.  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 ;  t,  trans- 
verse process. 


tiguous  vertebrae  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  vertebrae  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  (p)  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- 
brae, called  the  haemal  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  "  (az,pz),  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 
(f),  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  "  (m) 
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 
vertebras  of  one  are  apt  to  blend  into 
those  of  the  next  region,  either  normally 
or  as  peculiarities  of  individual  skeletons. 

Cervical  Vertebrae. — The  cervical  region 
constitutes  the  most  anterior  portion  of 
the  column,  or  that  which  joins  the 
cranium.  The  vertebras  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 
(the  vertebrarterial  canal,  Fig.  11,  v)  ;  or,  .  J°-  »-fne™  ™™  < 

\  .  1-1111  sixth  cervical    vertebra  of   Dog. 

as  it  is  sometimes  described,  they  have    s>  gpinous  process;  az,  anterior 

tWO  transverse  processes,  superior  and  zygapophysis  ;  t>,  vertebrarterial 
.  /.  i  .  i  ,1  •  •,•  canal ;  t,  transverse  process ;  t ,  its 

inferior,  which  meet  at  their  extremities    inferioylameiia. 

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  vertebrae  (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  vertebrae  is  the  same,  viz.  seven,  with  the  exception  of 
the  Manatee  and  Hoffman's  Two-toed  Sloth  (Cholozpus  hoffmanni), 
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  Vertebrce. — The  dorsal  (or,  as  it  would  be  more  correctly 
termed,  thoracic)  region  consists  of  the  vertebrae  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  Vertebra. — The  lumbar  region  consists  of  those  vertebrae 
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.  As  a  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  vertebrae; 
but  these  may  consist  of  1 2  dorsal  and  7  lumbar  vertebrae,  or  1 3 
dorsal  and  6  lumbar,  or  14  dorsal  and  5  lumbar.  The  smallest 
number  of  dorso-lumbar  vertebrae  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  (Cholcepus)  27,  and  the  Hyrax 
30.  The  prevailing  number  of  rib-bearing  vertebrae  is  12  or  13, 
any  variation  being  generally  in  excess  of  these  numbers. 

Sacral  Vertebra. — The  sacral  region  offers  more  difficulties  o* 
definition.  Taking  the  human  "  os  sacrum "  as  a  guide  for 
comparison,  it  is  generally  defined  as  consisting  of  those  vertebras 
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  vertebrae  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  vertebrae  in  the  following  pages.  The  Cetacea,  having  no  iliac 
bones,  have  no  part  of  the  vertebral  column  modified  into  a 
sacrum. 

Caudal  Fertebrce. — The  caudal  vertebrae  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 
vertebrae  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 
V.  These  are  always  situated 
nearly  opposite  to  an  interverte- 
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. 


FIG.  12.  —  Anterior  surface  of  fourth 
caudal  vertebrae  of  Porpoise  (Phoccena  com- 
munis).  s,  Spinous  process  ;  m,  metapophy- 
sis  ;  (,  transverse  process  ;  Ji,  chevron  bone. 


44 


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. 

Ribs. — The  ribs  form  a  series  of 
long,  narrow,  and  more  or  less  flattened 
bones,  extending  laterally  from  the 
sides  of  the  vertebral  column,  curving 
downwards  towards  the  median  line 
of  the  body  below,  and  mostly  joining 
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,  meeting  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  called  "  floating  " 
ribs.  The  portion  of  each  rib  nearest  the  vertebral  column  and 
that  nearest  the  sternum  differ  in  their  characters,  the  latter  being 
usually  but  imperfectly  ossified,  or  remaining  permanently  cartila- 
ginous. These  are  called  "costal  cartilages,"  or  when  ossified 
"  sternal  ribs." 


JCJt 


Fio.  13. --Human  sternum  and 
sternal  ribs,  ps,  Presternum ;  ms, 
mesosternum  ;  xs, ,  xiphisternum  ;  c, 
point  of  attachment  of  clavicle  ;  1  to 
10,  the  cartilaginous  sternal  ribs. 


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, 
and  the  rib  is  attached 
solely  to  its  corresponding 
vertebra.  The  number  of  pairs  of  ribs  is  of  course  the  same  as  that 
of  the  thoracic  vertebrae. 

The  circumstance  that  in  some  of  the  Anomodont  reptiles  and 


FIG.  14.— Sternum  and  strongly  ossified  sternal  ribs 
of  Great  Armadillo  (Priodon  gigas).  ps,  Presternum ; 
xs,  xiphisternum. 


FIG.  15.— Skeleton  of  Lion  (Felis  leo).  cd,  Caudal  vertebrae ;  cp,  carpus  ;  cr,  coracoid  process 
of  scapula  ;  cv,  cervical  vertebra  ;  d,  dorsal  vertebrae  ;  /&,  fibula ;  fm,  femur ;  h,  humerus  ;  il, 
ilium  ;  isch,  ischium ;  (,  lumbar  vertebrae ;  m,  metatarsus  ;  me,  metacarpus  ;  p,  patella ;  pb,  pubis ; 
ph,  phalanges  ;  pv,  pelvis  ;  r,  radius  ;  s,  sacral  vertebrae  ;  sc,  scapula ;  sk,  skull ;  tb,  tibia  ;  tt, 
tarsus  ;  u,  ulna ;  zy,  zygoinatic  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  girdkin  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 
lastr 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  Ursidce, 
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  Antebrachmm. — 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  5 
foramen,  which  corresponds  with  a  similar 
one  found  in  the  Anomodont  reptiles.  The 
hollow  in  the  head  of  the  ulna  for  the  recep- 
tion of  the  head  of  the  humerus  is  known 
as  the  greater  sigmoid  cavity,  and  that  for 
the  head  of  the  radius  as  the  lesser  sigmoid 
cavity  (Fig.  16).  The  term  olecranon  is 
applied  to  that  process  of  the  ulna  which 
forms  the  prominence  of  the  elbow. 

In  most  mammals  walking  on  four  limbs, 
in  which  the  hand  is  permanently  prone,  the 
ulna  is  much  reduced  in  size,  and  the  radius 
increased,  especially  at  the  upper  end ;  so 
that  the  articular  surface  of  the  latter,  instead  of  being  confined  to 
the  external  side  of  the  trochlea  of  the  humerus,  extends  all  across 


FIG.  16.— Outer  aspect  of 
the  proximal  extremity  of  the 
right  ulna  of  a  Bear  (Ursus). 
a,  Anterior  tubercle  ;  ol, 
olecranon ;  6,  greater  sigmoid 
cavity. ;  c,  lesser  do. 


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. 

Manus. — 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  (u),  the  first  being 
on  the  radial  or  preaxial  side  of  the  limb.1 
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  radiale 
supports  on  its  distal  side  two  bones, 
carpale  1  and  2 ;  the  intermedium  is  in  a 
line  with  the  centrale  and  carpale  3,  which 
together  form  a  median  axis  of  the  hand, 
while  the  ulnare  has  also  two  bones  articu- 
c,  centrale ;  i-5,  the  five  bones  of  lating  with  its  distal  end,  viz.  carpale  4 


Fio.  17.— Dorsal  surface  of  the 
right  manus  of  a  Water  Tortoise 
(Chelydra  serpentina).  After  Ge- 
genbaur. U,  Ulna  ;  R,  radius  ;  u, 
ulnare  ;  i,  intermedium  ;  r,  radiale  ; 


row  supports  a  metacarpal. 

1  The  opinion  has  recently  been  expressed  by  Baur  that  the  bone  termed 
radiale  in  Fig.  17  is  really  a  second  ceutrale,  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 
Intermedium 
Ulnare 
Centrale 
Carpale  1 
Carpale  2 
Carpale  3 
Carpale  4 
Carpale  5 


=  Scaphoid     =  Naviculare. 
=  Lunar         =  Semilunare,  Lunatum. 
=  Cuneiform  =  Triquetrum,  Pyramidale. 
=  Central       =  Intermedium  (Cuvier). 
=  Trapezium  =  Multangulum  majus. 
=  Trapezoid  =  Multangulum  minus. 
—  Magnum     =  Capitatum. 

=  Unciform  =Hamatum,  Uncinatum. 


The  radial  and  ulnar  sesamoids  are  regarded  by  Bardeleben l  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  coffer,  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  Aiiat. 
Anzeiger,  vol.  iv.  pp.  49-52,  1889.) 

1  On  the  Prapollex  and  Prrehallux,  etc.,  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 


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  vertebrae.  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  "  fabellae."  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.  1 7.  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. 

]•  =  Astragalus 1  =  Talus. 

Intermedium   J 

Fibulare  =  Calcaneum  =  Os  calcis. 

Centrale  =  Navicular  =  Scaphoideum. 

Tarsale  1  =  Internal  cuneiform  =  Entocuneiforme. 

Tarsale  2  =  Middle  cuneiform     =  Mesocuneiforme. 

Tarsale  3  =  External  cuneiform  =  Ectocuneiforme. 

Tarsale  4  )  ~  ,  . , 
rr  7  K  f  =  Cuboid. 
Tarsale  5  } 

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  calcaneum  (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 
(c1),  middle  (c2),  and  external  (c3),  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 
Eodentia ;  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  Cryptoprocta  there 
may  be  a  second  centrale,  which  usually 
coalesces  with  the  ectocuneiform. 

The  metatarsal  bones  never  exceed  five 
in  number,  and  the  phalanges  follow  the 
same  numerical  rule  as  in  the  manus,  never 
exceeding  three  in  each  digit.  Moreover, 
the  first  digit,  counting  from  the  tibial  side, 
or  hallux,  resembles  the  pollex  of  the  hand 
in  always  having  one  segment  less  than 
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  Halitherium. 


FIG.  18.— Bones  of  the  right 
Human  foot  T,  Tarsus ;  M, 
metatarsus;  Ph,  phalanges;  c, 
calcaneum;  a,  astragalus;  c6, 
cuboid ;  n,  navicular ;  c1,  inter- 
nal cuneiform  ;  c2,  middle  cunei- 
form ;  <;3,  external  cuneiform.  The 
digits  are  indicated  by  Roman 
numerals,  counting  from  the 
tibial  to  the  fibular  side. 


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  sufficient 
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  papillae,  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 


sm 


FIG.  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) ;  o,  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 ;  o,  aperture  of  do. ;  z,  zygomatic  arch.  (Mivart,  Proc. 
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  duct  (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  duct 


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  submaxillaries,  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  O3sophagus  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 


U 


FIG.  20. — Stomach  and  pancreas  of  the  Genet.  Posterior  or  dorsal  surface,  ce,  (Esophagus  ; 
s,  pancreas ;  pd,  pancreatic  duct ;  bd,  biliary  duct  from  the  liver.  (From  Mivart,  Proc.  Zool. 
Soc.  1882,  p.  305.) 

and  perhaps  S9ftening  or  otherwise  preparing  the  food  for  digestion. 
Sometimes  there  is  a  great  aggregation  of  glands  forming  distinct 
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 
Eodent  Lophwmys.  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  ccecum  coli,  as  it  was 
first  called,  a  name  generally  abbreviated  into 
"  caecum,"  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- 
lationship between  the  size  and  complexity 
of  this  organ  and  that  of  the  stomach. 
Where  the  latter  is  simple  the  caecum  is 
generally  the  largest,  and  vice  versd.  Both  the 
caecum  and  colon  are  often  sacculated,  a  dis- 
position caused  by  the  arrangement  of  the 
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  "  valvulae  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  duodenum, 
there  are  other  structures  in  the  mucous  membrane,  about  the 
nature  of  which  there  is  still  much  uncertainty,  called  "  solitary  "  and 


FIG.  21.  —  Diagrammatic 
plan  of  the  general  arrange- 
ment of  the  alimentary  canal 
in  a  typical  Mammal,  o, 
Oesophagus  ;  st,  stomach  ;  p, 
pylorus  ;  s,  s,  small  intestine 
(abbreviated)  ;  c,  caecum  ;  I,  I, 
large  intestine  or  colon,  end- 
ing in  r,  the  rectum. 


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. 

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,  u) 
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- 


Fio.  22. — Diagrammatic  plan  of  the  inferior  surface  of  a  multilobed  liver  of  a  Mammal. 
The  posterior  or  attached  border  is  uppermost,  u,  Umbilical  vein  of  the  foetus,  represented  by 
the  round  ligament  in  the  adult,  lying  in  the  umbilical  fissure ;  dv,  the  ductus  venosus ;  we, 
the  inferior  vena  cava  ;  p,  the  vena  portae  entering  the  transverse  fissure  ;  llf,  the  left  lateral 
fissure  ;  rlf,  the  right  lateral  fissure  ;  c/,  the  cystic  fissure  ;  U,  the  left  lateral  lobe  ;  Ic,  the  left 
central  lobe  ;  re,  the  right  central  lobe  ;  rl,  the  right  lateral  lobe  ;  «,  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,  rlf  and  llf).  When  these  are  more 
deeply  cut  than  the  umbilical  fissure  (u),  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  (re  and 
Ic)  separated  by  the  umbilical  fissure,  and  those  beyond  the  lateral 
fissures  on  each  side  being  the  right  and  left  lateral  lobes  (rl  and  II). 


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.1 

1  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  amoeboid  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  a  rule  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  -5-^-$  of 
an  inch,  which  is  exceptionally  large,  and  only  exceeded  by  the 
Elephant  (^TTT),  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  T.  javanicus  and  meminna  not  more  than  y^g^-g-.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  (chordae 
tendinice)  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  (os  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 

1  G.  Gulliver,  Proc.  Zool.  Soc.,  1862,  p.  91. 


64  GENERAL  ANATOMICAL   CHARACTERS 

each  guarded  by  three  semilunar  valves.  The  aorta  is  single,  and 
arches  over  the  left  bronchial  tube.  After  supplying  the  tissues  of 
the  heart  itself  with  blood  by  means  of  the  coronary  arteries,  it 
gives  off  large  vessels  ("carotid")  to  the  head  and  ("brachial")  to  the 
anterior  extremities.  The  mode  in  which  these  vessels  arise  from 
the  aorta  varies  much  in  different  mammals,  and  the  study  of  their 
disposition  affords  some  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  the  "  anterior  aorta  "  of  veterinary  anatomy ;  or 
they  may  be  detached  in  various  degrees,  both  arising  separately 
from  the  aorta,  as  in  Man,  or  the  left  carotid  from  the  innominate 
and  the  left  brachial  from  the  aorta,  a  very  common  arrangement ; 
or  the  last  two  from  a  common  second  or  left  innominate,  as  in 
some  Bats  and  Insectivores.  The  aorta,  after  giving  off  the  inter- 
costal arteries,  passes  through  the  diaphragm  into  the  abdomen,  and, 
after  supplying  the  viscera  of  that  cavity  by  means  of  the  gastric, 
hepatic,  splenic,  mesenteric,  renal,  and  spermatic  vessels,  gives  off 
in  the  lumbar  region  a  large  branch  (iliac)  to  each  of  the  hinder 
extremities,  which  also  supplies  the  pelvic  viscera,  and  is  continued 
onwards  in  the  middle  line,  greatly  diminished  in  size,  along  the 
under  surface  of  the  tail  as  the  caudal  artery.  In  certain  mammals, 
arterial  plexuses,  called  retia  mirabilia,  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  Euminants,  or  a  sponge-like  mass  of 
convoluted  vessels,  as  in  the  intercostals  of  Cetaceans,  are 
peculiarities  of  the  vascular  system  the  meaning  of  which  is 
not  in  all  cases  clearly  understood.  In  the  Cetacea  they  are  ob- 
viously receptacles  for  containing  a  large  quantity  of  oxygenated 
blood  available  during  the  prolonged  immersion,  with  consequent 
absence  of  respiration,  to  which  these  animals  are  subject. 

The  vessels  returning  the  blood  to  the  heart  from  the  head  and 
upper  extremities  usually  unite,  as  in  Man,  to  form  the  single  vena 
cava  superior  or  precaval  vein,  but  in  some  Insectivores,  Chiroptera, 
and  Rodents,  in  the  Elephant,  and  all  Marsupials  and  Monotremes, 
the  two  superior  caval  veins  enter  the  right  auricle  without  uniting, 
as  in  birds.  In  Seals  and  some  other  diving  mammals  there  is  a 
large  venous  sinus  or  dilatation  of  the  inferior  vena  cava  immediately 
below  the  diaphragm.  In  the  Cetacea  the  purpose  of  this  is  supplied 
by  the  immense  abdominal  venous  plexuses.  As  a  rule  the  veins 
of  mammals  are  furnished  with  valves,  but  these  are  said  to  be 
altogether  wanting  in  the  Cetacea,  and  in  the  superior  and  inferior 
cava,  subclavian  and  iliac  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  laminae  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  vertebrae,  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.  In  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. 

Trachea. — 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  bifurcation,  and  enters  the  right 
lung. 

Larynx. — 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. — The  thoracic  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. 

Lungs. — 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  in  connection  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.  Th6y  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  Equidce  there  are  large  postrpharyngeal  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 
Macrorhinus)  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  axillae,  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. 

Madder. — 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- 


70  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 


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-          "MPHW/  "*  '"H ^   ^c 

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  Palae- 
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 
advance  of  time, — indicating 
in  this,  as  in  so  many  other 
respects,  a  gradual  progress 
from  a  lower  to  a  higher  type 
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  vertebrae,  and  gives  off  the  compound  double-rooted 
nerves  of  the  trunk  and  the  extremities,  corresponding  in  number 
to  the  vertebrae,  thflSugh  the  interspaces  between  which  they  pass 


B 


FIG.  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 ; 
c.m.s,  calloso-marginal  sulcus;  c,  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 
lateral  gyrus  of  do. ;  os,  supraorbital  sulcus  of  do.  ; 
*/,  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 
cord.  (Mivart,  Proc.  Zool.  Soc.  1882,  p.  516.) 


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  papillae,  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  "  vibrissse  "  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  papillae.  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  papillae  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 — Edentates,  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  foetal  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  externus),  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.  L  pp. 
371-497,  pi.  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  mammillae  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  Sexual  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  Sex, 
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  Bovidce.  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- 
pogori),  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  Ornithorhynchits 
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  foetus  during  intra-uterine  life  takes 
place  through  the  medium  of  certain  structures,  partly  belonging 
to  the  fcetus  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  fcetus,  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  fcetus,  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  fcetus  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  egg  membranes  of  the  Monotremes  present  many  points  of 
agreement  with  those  of  the  ovum  of  the  Marsupials,1  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  K.  Owen  upon  the  development  of  the  Kangaroo  have  been 
confirmed  by  those  of  Dr.  H.  C.  Chapman,2  while  Dr.  Selenka,3  and 
Professor  H.  F.  Osborn4  have  contributed  important  evidence  as  to  the 
structure  and  relations  of  the  festal  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  foetus  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.  Sci.  Philadelphia,  1881,  p.  468. 

8  "  Studien  ueber  Entwickelungeschichte  der  Thiere,"  pt.  4,  Wiesbaden,  1886. 
4  Journal  of  Morphology,  vol.  i.  p.  373  (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  Eodentia,  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 


So  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  fcetal  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  Avith  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  fcetal  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  placentae  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  foetal  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 

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). 


CHAPTEE    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,  Eeptilia,  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 l  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  Society  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 l  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,2 
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.  93-104  (1887). 


ORIGIN  AND   CLASSIFICATION 


The  pes  of  the  Amphibia  and  Anomodontia  agree  in  having  a 
distinct  intermedium,  tibiale,  fibulare,  and  centrale,  whereas  in 
other  Keptiles  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  palseontological  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  Eutheria.  It  would,  however,  be  merely 
encumbering  zoological  science  with  new  names  to  give  them  any 
other  than  the  ordinarily  known  family  designations  of  Ornitho- 
rhynchidce  and  Echidn-idce. 

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 
"  Encyclopaedia." 

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- 
ttis,  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 


CLASSIFICA  TION  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  Archaeoceti.  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,  Ruminantia,  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  Galeopithecidce, 
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  subordirial  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 : — Hapalidce  or  Marmosets ;  Cebidce  or 
American  Monkeys,  with  three  premolar  teeth  on  each  side  of  each 
jaw ;  Cercopithecidce,  containing  the  majority  of  Old-world  Monkeys  ; 
Simiidce,  consisting  of  the  genera  Hylobates,  Simia,  Gorilla,  and 
Anthropopithecus,  the  true  Man -like  Apes;  and,  lastly,  Hominidce, 
containing  the  genus  Homo  alone. 

Subclass  I.  PKOTOTHERIA. 
Order  i.  MONOTREMATA — Monotremes. 

Fam.  1.  Ornithorhynchidce — Duck-bill. 
2.  Echidnidce — Spiny  Anteater. 

Group.  MULTITUBEROULATA.1 

Fam.  1.  Plagiaulacidae — Plagiaulax. 

2.  Polymastodontidae — Polymastodon. 

3.  Tritylodontidse — Tritylodon. 

Subclass  II.  METATHERIA. 
Order  ii.  MARSUPIALIA — Marsupials. 

Suborder  1.  POLTPROTODONTIA — Polyprotodonts. 

1  The  names  of  the  groups  containing  only  extinct  forms  are  printed  in  heavier 
type  than  those  which  contain  species  still  existing. 


CLA  SSIFICA  TION  89 

Fam.  1.  Dromatheriidae — Dromatherium. 

2.  Amphitheriidae — Amphitherium,  etc. 

3.  Spalacotheriidae — Spalacotlieriuni. 

4.  Tritylodontidae — Tritylodon. 

5.  Didelphyidce — Opossums. 

6.  Dasyuridce — Thylacine  and  Dasyures. 

7.  Peramelidce — Bandicoots. 

Suborder  2.  DIPROTODONTIA — Diprotodonts. 
Fam.  8.  Phascolomyidce — Wombats. 
9.  Phalangeridce — Phalangers. 

10.  Diprotodontidse — Diprotodon. 

11.  Nototheriidae — Notothere. 

1 2.  Macropodidce — Kangaroos. 


Subclass  III.  EUTHERIA. 
Order  iii.  EDENTATA — Edentates. 

Fam.  1.  Bradypodidce — Sloths. 

2.  Megatheriidae — Ground  Sloths. 

3.  MyrmecophagidcB — Anteaters. 

4.  Dasypodidce — Armadillos. 

5.  Glyptodontidae — Glyptodonts. 
C.  Manidce — Pangolins. 

7.  Orycteropodidce — Aard-varks. 

Order  iv.  SIRENIA — Sirenians. 

Fam.  1.  Manatidce — Manatees. 

2.  Rhytinidae — Rhytina. 

3.  Halicoridce — Dugongs. 

4.  Halitheriidae — Halithere. 

Order  v.  CETACEA — Cetaceans. 

Suborder  1.  MYSTACOCETI — Baleen  Whales. 
Fam.  1.  Balcenidce — Greenland  Whale,  etc. 

Suborder  2.  ARCKEOCETI. 

Fam.  2.  Zeuglodontidse — Zeuglodonts. 

Suborder  3.  ODONTOCETI — Toothed  Whales. 
Fam.  3.  Physeteridce — Sperm  Whale. 

4.  PlatanistidcK — Freshwater  Dolphins. 

5.  Delphinidw — Dolphins,  Porpoises,  etc. 

Order  vi.  UNGULATA — Hoofed  Mammals. 
Suborder  1.  ARTIODACTYLA — Artiodactyles. 
Section  A.  SUINA — Pig-like  Artiodactyles. 
Fam.  1.  Hippopotamidce — Hippopotamus. 
2.  Suidce — Pigs  and  Peccaries. 


90  ORIGIN  AND   CLASSIFICATION 

3.  Chceropotamidae — Choeropotamus. 

4.  Anthracotheriidae — Anthracothere. 

5.  Merycopotamidae — Merycopotamus. 

6.  Cotylopidae — Oreodonts. 

7.  Anoplotheriidae — Anoplothere. 

8.  Dichodontidae — Dichodon. 

Section  B.  TRAGULINA — Chevrotains. 

9.  Tragulidce — Chevrotains. 

Section  C.  TYLOPODA — Camels. 

10.  Camelidce — Camels  and  Llamas. 

11.  Poebrotheriidae — Poebrotherium. 

Section  D.  PECORA — True  Ruminants. 

12.  Cervidce — Deer. 

13.  Giraffid(e — Giraffe. 

14.  Antilocapridce — Prong-buck. 

15.  Bovidw — Sheep,  Cattle,  etc. 

Suborder  2.  PERISSODACTYLA — Perissodactyles. 

Fam.  16.  Tapiridce — Tapirs. 

1 7.  Lophiodontidae — Lophiodonts. 

1 8.  Palaeotlieriidae — Palaeotheres. 

19.  Equidce — Horses. 

20.  Rhinocerotidce — Rhinoceroses. 

2 1 .  Lambdotheriidae — Palaeosyops. 
2  2.  Chalicotheriidae — Chalicothere. 

23.  Titanotheriidae — Titanothere. 

24.  Macraucheniidae — Macrauchenia. 

Suborder  3.  TOXODONTIA— Toxodonts. 

Fam.  25.  Toxodontidse — Toxodon. 
26.  Typotheriidae — Typothere. 

Suborde'r  4.  CONDYLARTHRA. 

Fam.  27.  Periptychidae — Periptychus. 

28.  Phenacodontidae — Phenacodus. 

29.  Meniscotheriidae — Meniscothere. 

Suborder  5.  HYRACOIDEA — Hyraces. 
Fam.  30.  Hyracidce — Hyrax. 

Suborder  6.  AMBLYPODA. 

Fam.  31.  Pantolambdidae — Pantolambda. 

32.  Coryphodontidae— Coryphodon. 

33.  Uintatheriidae — Uintathere. 

Suborder  7.  PROBOSCIDEA — Proboscideans. 
Fam.  34.  Dinotheriidae — Dinothere. 
35.  ElephantidcK — Elephants. 


CLA  SSI  PICA  TION  9 1 

Group.  TILLODONTIA— Tillodonts. 

Fam.  Anchippodontidae — Anchippodus. 
Calamodontidae — Calamodon. 

Order  vii.  KODENTIA — Rodents. 
Suborder  1.  SIMPLICIDENTATA. 

Fam.  1.  Anomalurida — Anoraalurus. 

2.  Sciuridce — Squirrels  and  Marmots. 

3.  Haplodontidce — Haplodon. 

4.  Ischyromyidae — Ischyromys. 

5.  Castoridee — Beavers. 

6.  Myoxidce — Dormice. 

7.  Lophiomyidce — Lophiomys. 

8.  Muridce — Rats,  Mice,  and  Voles. 

9.  Spaladdce — Mole-rats. 

10.  Geomyidce — Pouched  Rats. 

11.  Dipodidce — Jerboas. 

1 2.  Theridomyidae — Theridomys. 

13.  Octodontidce — Spiny  Mice. 

14.  Castoroididae — Castoroides. 

1 5.  Hystricidce — Porcupines. 

1 6.  ChinchillidoR — Chinchillas. 

17.  Dinomyidce — Dinomys. 

18.  Caviidce — Cavies. 

19.  Dasyproctidce — Agouties. 
Suborder  2.  DUPLICIDENTATA. 

Fam.  20.  Lagomyidce — Picas. 

21.  LeporidtK — Hares  and  Rabbits. 

Order  viii.  CARNIVORA — Carnivores. 

Suborder  1.  CARNIVORA  VERA — Fissipedes. 
Fam.  1.  Felidce — Cats. 

2.  Hycenidce — Hyaenas. 

3.  Proteleidce — Earth-wolf. 

4.  Viverrida — Civets  and  Ichneumons. 

5.  Canidte — Wolves  and  Foxes. 

6.  Ursid(B — Bears. 

7.  Mustelidce — Weasels  and  Otters. 

8.  Procyonidce — Racoons  and  Cat-bear. 
Suborder  2.  PINNIPEDIA — Pinnipedes. 

Fam.  9.  Otariidce — Eared  Seals. 

10.  Trichechidce — Walrus. 

11.  Phocidce — Seals. 

Suborder  3.  CEEODONTA — Creodonts.       . 
Fam.  12.  Hyaenodontidae — Hyaenodon. 

1 3.  Proviverridae — Proviverra. 

14.  Arctocyonidae — Arctocyon. 

1 5.  Mesonychidae — Mesonyx. 


92  ORIGIN  AND   CLASSIFICATION 

Order  ix.  INSECTIVORA — Insectivores. 

Suborder  1.  INSECTIVORA  VERA. 

Fam.  1.  Tupaiidce — Tupaias. 

2.  Macroscelididce — Elephant-Shrews. 

3.  Erinaceidce — Hedgehogs. 

4.  Soricidce — Shrews. 

5.  Talpidce — Moles. 

6.  Potamogalidce — Potamogale. 

7.  SolenodontidcB — Solenodon. 

8.  Centetidce — Centetes. 

9.  Chrysochloridce — Golden  Moles. 

Suborder  2.  DERMOPTERA. 

Fam.  10.  Galeopithecidce — Galeopithecus. 

Order  x.  CHIROPTERA — Bats. 

Suborder  1.  MEGACHIROPTERA — Frugivorous  Bats. 

Fam.  1.  Pteropodidce — Flying  Foxes. 
Suborder  2.  MICROCHIROPTERA — Insectivorous  Bats. 

Fam.  2.   Vespertilionidce — Common  Bats. 

3.  Nycteridce — Nycteris. 

4.  Rhinolophidce — Leaf-nosed  Bats. 

5.  Emballonuridce — Emballonura. 

6.  Phyllostomatidce — Vampyres. 

Order  xi.  PRIMATES. 

Suborder  1.  LEMUROIDEA — Lemuroids. 
Fam.  1.  Hyopsodontidae — Hyopsodus. 

2.  Chiromyidce — Aye-Aye. 

3.  Tarsiidce — Tarsier. 

4.  Lemuridce — Lemurs. 

Suborder  2.  ANTHROPOIDEA — Anthropoids. 
Fam.  5.  Hapalidce — Marmosets. 

6.  Cebidce — American  Monkeys. 

7.  Cercopithecidce — Old  World  Monkeys. 

8.  Simiidce — Gibbons  and  Man-like  Apes. 

9.  Hominidce. — 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. 


CHAPTEE    IV 

GEOGRAPHICAL  AND  GEOLOGICAL  DISTRIBUTION 
I.    GEOGRAPHICAL  DISTRIBUTION.1 

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,  palaeontology  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  Eeindeer  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  ^Eluropus  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  Dasyuridce,  to 
which  that  genus  belongs,  ranges  all  over  Australia,  while  the  order 
Marsupialia,  which  includes  the  Dasyuridce,  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  Muridce  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 
fades  of  their  fauna  as  a  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 
Hyaenas,  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  Palsearctic 


ZOOLOGICAL   REGIONS— PALAEARCTIC  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  Palaearctic  and  Nearctic  regions  have  a  large  number 
of  common  types,  more  especially  among  the  mammals,  and  Dr.  A. 
Heilprin l  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 
Palaearctic,  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  Palaearctic, 
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. 

Pal&arctic  Region. — The  Palaearctic  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  JBovidce,  nearly  all  the  species  of  which 
are  Palsearctic,  although  we  meet  with  one  Goat  (Copra)  in  the 
Nilgherries  of  Southern  India,  and  a  Sheep  (Ovis)  in  the  Nearctic 
region.  The  Musk  Ox  (Ovibos)  is  characteristic  of  the  Palsearctic 
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  (fiupicapra),  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  (Moschus),  the  sole  representative  of  the  subfamily 
Moschince,  is  exclusively  Palaearctic.  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  (Talpa)  are  mainly  Palaearctic, 
although  one  species  enters  Northern  India,  while  the  Desmans 
(Myogale)  of  the  Pyrenees  and  Southern  Kussia  are  unknown 
beyond  the  limits  of  this  region.  The  Water-shrew  (Nedogale)  is 
likewise  a  peculiar  eastern  Palsearctic  type.  Among  the  Rodents, 
the  Picas  or  Tailless  Hares  (Lagomys)  and  the  Dormice  (Myoxus) 
are  essentially  Palsearctic  forms,  only  one  species  of  each  being  found 
beyond  the  limits  of  the  region,  and  the  one  extra-Palaearctic  species 
of  Lagomys  occurring  in  the  cognate  Nearctic  region.  The  Mice  and 
Rats  are  represented  by  the  typical  genus  Mus  and  other  types, 
and  Hares  (Lepus)  and  one  species  of  Squirrel  (Sciurus)  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  (Giro/fa)  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  (Pliacochcerus)  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  Oryx  ranges  into  Arabia 
and  Persia.  In  contrast  to  this  abundance  of  Antelopes  is  the  total 
absence  of  the  Deer  family,  or  Cervidce,  which  are  so  characteristic 
of  the  Paleearctic  and  Oriental  regions.  The  Chevrotains  or 
Tragulidce  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  (Equidce)  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  Hysenas,  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  Otocyon.  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  Simiidce,  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- 
cidce.  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  ffyomoschus,  but  first  described  as  an  extinct  form 
under  the  above  name. 


ioo  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  (Manis),  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 
Khinoceroses,  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  Palaearctic  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 
Palaearctic  Red  Deer  belongs.  True  Chevrotains  (Tragidus)  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  101 

only  by  a  single  species  in  the  desert  area  of  the  Indian  sub-region, 
while  the  two  species  of  Rhinoceros  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  Palsearctic. 
Civets  are  abundant,  comprising  some  peculiar  genera,  of  which  it 
will  suffice  to  mention  the  well  known  Paradoxurus.  Wolves  closely 
allied  to  the  Palaearctic  species  occur  in  Northern  India,  and  there 
are  also  Foxes  related  to  the  typical  species.  The  Dog-like  animals 
which  hunt  in  packs,  and  are  separated  by  some  writers  from  Canis 
under  the  name  of  Cyon,  occur  in  the  present  and  the  Palsearctic 
region.  The  striped  Hyaena  is  the  Indian  representative  of  its  genus. 
Ratels  are  common  to  this  and  the  Ethiopian  region,  and  constitute 
the  genus  Mellivora.  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  Melursus.  Among  the  Rodents  we  may  especially  notice 
the  abundance  of  the  Muridce  and  Sciuridce.  In  the  former  family 
we  have  numbers  of  true  Mice  (Mus),  and  also  the  peculiar  genus 
Nesocia  (Bandicoot-Rat),  while  in  the  latter  both  the  true  Squirrels 
(Sciurus)  and  the  Flying-Squirrels  (Pteromys)  attain  great  develop- 
ment. The  genus  (Pteromys)  is,  indeed,  mainly  characteristic  of  this 
region,  although  in  Kashmir  and  Japan  it  enters  the  Palaearctic. 
The  Bats  are  very  numerous,  being  represented  by  all  the  families, 
with  the  exception  of  the  Phyllostomatidce,  or  Vampyres,  of  South 
America.  Among  the  Insectivora  the  genera  Tupaia  and  Galeo- 
pithecus  (Flying  Lemur)  are  peculiar  to  this  region,  although  not 
found  in  India.  Finally,  in  the  Primates  we  have  the  genera 
Macacus  and  Semnopithecus  very  abundantly  represented,  although 
both  also  enter  the  Palaearctic  region ;  but  the  Anthropoid  types 
are  confined  to  the  south-eastern  half  of  the  region,  and  include  the 
Orangs  (Simia)  of  Borneo,  and  the  smaller  long-armed  Gibbons 
(Hylobates),  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  (Manis),  which,  as  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  (Muridce),  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  (Didelphyidce), 
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  Balrirusa.  Several  other  types  of 
placental  mammals  are  found  in  this  transitional  area,  while  the 
Marsupials  are  represented  by  Phalanger  and  Petaurus. 

Neardic  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  Palsearctic  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  Palsearctic  genus 
Ovis,  namely  the  Big-horn,  and  the  Prong-buck  (Antilocaprd),  which 
is  quite  peculiar.  There  are,  however,  no  Perissodactyla.  The 
Racoons  and  Coatis  (Procyonidce)  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  Eats  (Geomyidce),  and  the  Beaver-like 
rodents  known  as  the  Haplodontidce.  True  Rats  and  Mice  (Mus), 
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  Cricetus,  although  often  separated 
as  Hesperomys.  This  feature  alone  would  seem  to  justify  the  dis- 
tinction of  the  Nearctic  from  the  Palsearctic  region.  The  Musquash 
(Fiber}  is  a  genus  of  Nearctic  rodents  unknown  in  the  Old  World. 
Among  other  characteristic  genera  we  may  mention,  in  the  Carnivora, 
the  Skunk  (Mephitis)  and  the  American  Badger  (Taxidea).  Primates 
are  absent  from  the  entire  region. 

Neotropical  Region. — The  last  of  the  six  main  regions  is  the 
Neotropical,  including  Mexico,  South  America,  and  the  West  Indies. 
A  very  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  regions  as  llanos,  savannas, 
and  pampas.  The  back -bone  of  the  region  is  formed  by  the  great 
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  Ungulates,  we  find  a  total 
absence  of  Antelopes,  Sheep,  and  Oxen,  and  also  of  all  Perissodac- 
tyles  except  Tapirs.  Deer  are,  however,  represented,  although  by 
peculiar  forms  (Cariacus)  unknown  beyond  the  New  World.  The 
Peccaries  (Dicotyles),  which  are  often  made  the  type  of  a  distinct 
family,  take  the  place  of  the  Old  World  Pigs,  while  the  Llamas  and 
Alpacas  (AucJienia)  are  the  substitutes  for  the  Palaearctic  Camels. 
The  Carnivora  include  several  Cats  (Felis),  among  which  the  Puma 
and  the  Jaguar  are  the  most  noticeable ;  and  there  are  also  Racoons, 
Coatis,  Foxes,  and  one  species  of  Bear.  Insectivora  are  totally 
wanting ;  but  the  Bats  are  characterised  by  the  presence  of  the 
Vampyres  (Phyllostomatidce),  which  are  almost  restricted  to  this 
region.  The  Rodents  likewise  include  three  families  unknown 
elsewhere,  namely  the  Chinchillas  and  Viscacha  (Chinchillidce),  the 
Agouties  (Dasyproctidce),  and  the  Cavies  (Caviidce) ;  while  a  large 
number  of  the  Ododontidce  are  Neotropical,  all  the  other  forms 
being  Ethiopian.  In  the  Primates,  again,  we  have  all  the  forms 
quite  peculiar  to  this  region,  and  constituting  two  families,  viz.  the 
Cebidce  or  Prehensile -tailed  Monkeys,  and  the  Hapalidce,  or  Mar- 
mosets, both  of  which  differ  decidedly  in  their  dentition,  as  well 
as  in  other  features,  from  the  Old  World  Monkeys.  Lemuroids 
are  unknown.  Perhaps,  however,  the  mammals  which  may  be 
considered  as  most  characteristic  of  the  Nearctic  region  are  the 
numerous  Edentates,  which  form  three  families,  mostly  confined  to 
it.  These  comprise  the  Bradypodidce  or  Sloths,  which  solely 
inhabit  the  forest  region ;  the  Myrmecopliagidce  or  Anteaters ;  and 


104  GEOGRAPHICAL  DISTRIBUTION 

the  Dasypodidce  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  (Baltmoptera}  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.  forsteri  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  (Balcena)  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.  Uscayensis), 
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  (Balcena  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  Hyperoodon  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  (Neobalcena 
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. 


io6  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  P.  gangetica  have  been 
shown  by  Anderson  to  be  of  no  constant  value.  Orcella  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  (Manatus)  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 
(Halicore)  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.  CAINOZOIC  OR  TERTIARY — 

1.  Pleistocene — River  alluvia,  etc. 

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,  etc. 

III.  PALAEOZOIC  OR  PRIMARY — 

1.  Permian — Beds  overlying  the  Coal. 

2.  Carboniferous — Coal-measures,  etc. 

3.  Devonian — Old  Red  Sandstone. 

4.  Silurian — Wenlock  Limestone,  etc. 

5.  Cambrian — Llanberis  Slate,  etc. 

6.  Archaean — Gneiss  and  other  schists. 

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 


io8  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  Palaeozoic. 

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.1  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  Tritylodon. 


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.1 

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 


FIG.  24. — Frontal  and  oral  aspects  of  the  cranium  of  Tritylodon  longtevus;  from  the  Karoo 
system  of  Basuto-land,  South  Africa.    §  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," Palccontographical  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  by  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. 


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.  24).  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  (Fig.  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 
those  forms  in  which 
these  teeth  are  known, 
while  there  are,  at  least 
usually,  only  two  in 
those  of  the  lower  jaw.  In  other  cases  the  premolars  are  of  a 
secant  type,  with  a  highly  convex  cutting-edge,  and  usually  either 
serrated  or  obliquely  grooved  (Figs.  25,  26).  From  a  certain 
resemblance  between  these  secant  premolars  and  those  of  some  of 
the  smaller  Macropodidce  it  was  at  one  time  considered  that  we  had 
in  these  mammals  representatives  of  Diprotodont  Marsupials.  The 
great  difference  in  the  structure  of  the  molar  teeth  of  these  forms, 


Fio.  25. — The  right  ramus  of  the  mandible  of  Plagiaulax 
beklesl ;  from  the  Purbeck  of  Swanage.  Twice  natural  size. 
i,  Incisor ;  m,  molar ;  6,  coronoid  process ;  c,  condyle.  (After 
Owen.) 


Fio.  26.— The  imperfect  right  ramus  of  the 
mandible  of  Plagiaulax  minor ;  from  Swanage. 
Four  times  natural  size,  p,  Premolars ;  m, 
molars.  (After  Lyall.) 


FIG.  27. — Stereognathus  oolithicus.  Frag- 
ment of  jaw  with  three  teeth  (a,  6,  c),  in 
matrix  ;  from  the  Stonesfleld  Slate.  Natu- 
ral size.  (After  Owen.) 


coupled  with  the  circumstance  that  when  the  number  of  upper 
incisors  is  reduced  below  three  it  is  the  second  in  place  of  the  first 
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  suggested  that 
the  Multituberculata  may  really  represent  an  order  of  Prototheria. 
Some  siapport  is  afforded  to  this  suggestion  by  certain  fragmentary 
bones  from  the  Cretaceous  of  the  United  States,  which  are  regarded 


MESOZOIC  MAMMALS  in 

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  Khsetian  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  Microlestes  antiquus.  Subsequently  Mr.  C.  Moore  discovered  in  a 
bone  bed  of  Ehsetic  (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  Microlestes,  and 
described  specifically  as  M.  moorei.  Under  the  name  of  Hypsi- 
prymnopsis  rhceticus,  Professor  Boyd  Dawkins  described  a  single  tooth 
with  two  roots  discovered  in  the  Rhsetic  Marlstone  at  Watchet  in 
Somersetshire.  Sir  R.  Owen  referred  the  latter  tooth  to  Microlestes, 
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  Microlestes  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  Macropodidce,  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  (Potormis  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  Plagiaulax  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  (b)  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)  Plagiaulax,  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)  R.  Owen, 
art.  "  Palaeontology,"  Encyclopaedia  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,"  Palceontographical  Society,  1871. 


MESOZOIC  MAMMALS  113 

Jurassic  of  North  America ;  and  the  Plagiaidacidce  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  (Meniscoessus  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 
(Triacanthodori),  in  which  the  formula  of  the  lower  teeth  is  i  3,  c  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  an tero- posterior  line,  those  of  the 

i    i  .         ,     .  i .,  n    .  Fio.  28.— Reversed  view  of  the 

upper  and  lower  jaw  being  alike.     Pna-    left  rainus  of  ^  mandible  of 

COdon,  of  the  Jurassic  Of  the  United  States,      Triconodmt,    mordax ;    from    the 

is  probably   inseparable  from  Tricmod&n.    Purbeck  of  8wanag'.    Natural 

T     \,  *        r>7          j  j?       •  /T-L-       o«\       f     Slze-    (After  Owen.) 

In  the  genus  Phascolot/ienum  (iig.  29)  of 

the  Lower  Jurassic  Stonesfield  Slate,  the  lower  teeth  may  be 
classified  as  i  4,  c  1,  p  3,  m  4,  the  premolars  and  molars  being 

8 


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 


Fio.  29. — Inner  view  of  the  right  ramus  of  the  mandible  of  Phascolotherium  bucklandi ; 
from  the  Stonesfield  Slate.  The  outline  shows  the  natural  size,  i,  Incisors  (one  missing)  ;  c, 
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  Amblotherium,  of  the 
Dorsetshire  Purbeck,  to  which  Amphitherium  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 
hypocone.  A  similar 
form  of  molar  occurs 
in  the  existing  Opos- 
sums and  Bandicoots. 
The  number  of  lower 

Fio.  30.— Reversed  inner  view  of  the  left  ramus  of  the  .  . 

mandible  of  Amphilestes  broderipi;    from  the  Stonesfield  teeth    m    AmOiOtfienum 

Slate.    Twice  natural  size.    The  restoration  of  the  anterior  is    i     4,    C    1,     _p     4,    WJ 

teeth  is  conjectural,  and  the  condyle  is  placed  too  high.  ^,_g  ^'umerOUS  allied 
(After  Owen.) 

types,  such  as  Achyro- 

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  case  so-called  genera  like  Stylodon  and  Stylacodon  have  been 
formed  upon  specimens  showing  the  opposite  side  to  that  which 
is  exposed  in  the  types  of  Amblotherium  and  Amphitlwrium.  The 


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  Myrmecobius,  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  Myrmecobius  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  ^  u  _part  of  ^  ^  ramus  of  the  ^ 

apparently  referable  to  the  Same  dible,  viewed  from  the  outer  side,  of  Spala- 
animal.  Upper  and  lower  teeth  cotherium  tncuspidens ;  from  the  Purbeck  of 

of  this  form  are  represented  in  Swanage'  Twic.  Mto»i  *».  (After  Owen.) 
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,  i  3,  c  1, 
p  +  m  10,  by  which  it  is  widely  distinguished  from  all  the  Insect- 
ivora.  Menacodon,  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 


Ii6  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- 
liarity 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. 


CHAPTEE   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  Ornithorhynchidce  and  the  Echidnidce,  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 


1 1 8  MONO  TREMA  TA 


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  vertebrae  are  nineteen  in  number,  and  have 
no  terminal  epiphyses  to  their  bodies.  The  tranverse  processes  of 
the  cervical  vertebras  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, 


ORNITHORH  YNCHID^E  1 1 9 

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  vertebrae 
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 
fossae.  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 
supports  in  the  adult  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-caecal  valve, 
or  marked  distinction  between  large  and  small  intestine,  but  has  a 
small,  slender  vermiform  caecum  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  representatives  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  ORNITHORHYNCHID^E. 

Ornithorhynchus.1 — Cerebral    hemispheres    smooth.     Premaxillae 
and  mandible  expanded  anteriorly  and  supporting  a  horny  beak 
1  Blumenbach,  Voigts  Magazin,  vol.  ii.  p.  205  (1800). 


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.  Vertebrae:  C  7,  D  17,  L  2,  S  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  Naturalist's  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 


ORNITHORHYNCHID& 


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, 


Fio.  32. — Platypus  or  Duck-bill  (Ornithorhynchus  anatlnus).    From  Gould's  Mammals  of 

Australia. 

flattened,  and  glossy  toward*  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, 


1 22  MONO  TREMA  TA 


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 l  announced  the  discovery 
in  an  embryo  of  teeth  which  were  regarded  as  quite  functionless.  In 
the  following  year,  however,  Mr.  0.  Thomas  2  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  among 
other  known  mammals  Mr.  Thomas  considers,  as  was  first  suggested 
by  Professor  Cope,  that  those  of  the  Mesozoic  Multituberculata  (p.  1 09) 
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.  353  (1888). 
-  Ibid.  vol.  xlvi.  p.  126  (1889). 


ORNITHORHYNCHID&  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." 

Eeverting  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.  1 62),  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, 


1 24  MONO  TREMA  TA 


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.  A.  B. 
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  ECHIDNID.E. 

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.  Kami  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- 
rhynchidce.  According  to  Mr.  0.  Thomas,  all  the  living  forms  may 


ECHIDNID&  i25 

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

Echidna.1 — 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  vertebrae  is  C  7,  D  16,  L  3,  S  3,  Ca  12. 
The  one  existing  representative  of  the  genus  (E.  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  EUmentaire  d'ffist.  Nat.  p.  143  (1798). 


1 26  MONO  TREMA  TA 


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. 

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


FIG.  33.— The  Three-toed  Echidna  (Proechidna  Iruijnii).    From  Gervais. 

the  largest  race  of  Echidna  aculeata.  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  vertebrae  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. 

1  Gervais,  OsUographie  des  Monotremes,  p.  43  (1877). 


ECHIDNID&  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  E.  ramsayi  by  Sir  R.  Owen. 
In  referring  this  species  to  the  genus  Echidna,  that  term  must  be 
regarded  as  including  Proechidna. 


CHAPTEE  VI 

THE   SUBCLASS  METATHEKIA  OK  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 
MARSUPIALIA,1  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- 
rous)  this  tooth  re- 
tains its  place  and 

f         t°          until     fhp  Fl0'  34-— Teeth  of  upper  jaw  of  Opossum  (Didelphys  mar- 

J  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  f°rm  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 

9 


1 30  ME  TA  THERIA 

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  vetebrae  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  venae  cavae,1  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  faeces, 
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 
mammae  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   Petaurus  (Belideus)  breviceps  (Forbes,  Proc.   Zool.  Soc.    1881, 
p.  188). 


DISTRIBUTION 


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 


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

exception  of  one  family  (the  Didelphyidce),  limited  in  geographical 
distribution  to  the  Australasian  region,1  forming  the  chief 
mammalian  fauna  of  Australia, 
New  Guinea,  and  some  of  the 
adjacent  islands.  The  Didel- 
phyidce 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 
them  is  derived  from  the  form 


FIG.  36. — Front  view  of  skull  of  Koala  (Phas- 
colarctus  cinereus),  showing  diprotodont  and 
herbivorous  dentition  (Quart.  Journ.  Geol.  Soc. 
vol.  xxiv.  p.  313). 

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  Peramelidce  in  the  former,  as  well  as  in  the  latter ;  but  if 
Mr.  0.  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.  POLTPBOTODONTIA. — Incisors  numerous,  small,  subequaL     Canines 

larger  than  the  incisors.     Molars  with  sharp  cusps. 

a.  Incisors  y.  Hind  feet  with  the  four  outer  toes  subequal, 
distinct,  and  a  well-developed  opposable  hallux.  Didel- 
phyidce. 

/3.  Incisors  ^.   Hind  feet  with  four  outer  toes  distinct.    Hallux 

small  or  rudimentary,  rarely  opposable.     Dasyuridce. 

/4 g\ 

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).  Perainelid(e. 

B.  DIPROTODONTIA. — Incisors  not  exceeding  f ,  usually  y,  but  occasion- 

ally -p  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  T,  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- 
myidce. 


DIDELPHYIDsE 


133 


(3.  Teeth  rooted.  Three  upper  incisors  and  a  canine.  Hind 
limbs  not  disproportionately  large.  Feet  syndactylous, 
broad,  with  four  subequal  outer  toes,  and  a  large 
opposable  hallux.  Phalangeridce. 

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


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^E. 


Dentition :  i  -£-,  c  ^,  p  ^ ,  m  % ;  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.  Caecum  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.  Vertebrae : 
C  7,  D  13,  L  6,  S  2,  C  19-35. 

The  Didelphyidce,  or  true  Opos- 
sums, differ  from  all  other  existing 
Marsupials  in  their  habitat,  being 
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 


FIG.  37. — Skeleton  of  the  right  hind 
foot  of  the  Virginian  Opossum  (Didelphys 
marsupialis). 


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.1 — 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  Didelphijs  is  a  very  large  one,  con- 
taining, according  to  Mr.  0.  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  (D.  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 

1  Illiger,  Prod.  Syst.  Afamm.  et  Aves,  p.  76  (1811). 

2  Linn.  Syst.  Nat.  Ed.  12,  vol.  i.  p.  71  (1766). 


DIDELPHYID&  135 

fourth  or  Micoureine  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 


FIG.  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  based  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 


Family  DASYURID^E 

Dentition  :  i  ^,  c  ^,  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 
csecum.  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  Myrmecobius, 
though  clearly  a  member  of  this  family,  is  so  sharply  distinguished 


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

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

Subfamily  Dasyurinse. — This  comprises  the  more  typical  Dasy- 
uridce,  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.1 — Dentition  :  i  $,  c  \,p  f,  m  %  =  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  re- 
1  Temminck,  Monographies  de  Mammalogie,  vol.  i.  p.  60  (1827). 


DASYURID^E 


137 


sembling  those  of  Dasyurus,  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.  Vertebrae  : 
07,  D13,  L6,  S  2,  C  23.  Marsupial  bones  represented  only  by 
small  unossified  fibro-cartilages. 

The  only  known  existing  species  of  this  genus,  T.  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 


Fio.  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,  "  Hyaena."  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 
mammae.  The  figure  of  the  skull  exhibits  the  peculiar  Dog-like 
form  so  characteristic  of  the  genus. 

Sarcophilus.1 — Dentition :  i  £,  c  ^,  p  f ,  m  £.  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  Mammiftres,  iv.  (1837). 


138  MARSUPIALI A 


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. 

Dasyurus.1 — Dentition  :  i  -£,  c  ^,  p  $ ,  m  % ;  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. 
Vertebra  :  C  7,  D  13,  L  6,  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 ;  D.  viver- 
1  Geoffrey,  Bull.  Soc.  Philom.  vol.  i.  p.  106  (1796). 


DASYURID&  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. 

Phascologale.1 — 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  *$>C^,p$,m£;  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. 
Mammae  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  Chcetocercus  of  Krefft,  but  is  included  by  Mr.  0.  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.3 — The  last  genus  of  the  Dasyurince  is  Antechinomys, 
represented  only  by  A.  laniger  of  Queensland  and  New  South  Wales. 
This  elegant  little  mouse-like  creature,  which  has  large  oval  ears  and 

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

2  Thomas,  Ann.  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  bullee  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  Myrmeeobiinse. — Molars  and  premolars  exceeding 
the  normal  number  of  seven  on  each  side.  Tongue,  long  cylindrical, 
and  extensile. 

Myrmecobius.1 — Dentition  :  i  £,  c  ^,  p  f ,  m  •£  or  $• ;  total  52  or  56, 


FIG.  41. — Myrmecdbius  fasciatus.    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.  Soc.  1836,  p.  69. 


PERAMELID^  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. 
Vertebrae:  C  7,  D  13,  L  6,  S  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  PERAMELID^E. 

M 5)       134 

Dentition  :  i  — 3— ,  c  j,  p  3,  in  -  •  total  46  or  48.    Upper  incisors 

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.  Caecum  of  moderate  size.  Pouch  complete, 
opening  backwards.  Alone  among  Marsupials  they  have  no  clavicles. 

The  Peramdidce  form  a  very  distinct  family,  in  some  respects 
intermediate  between  the  sarcophagous  Dasyuridce  and  the 
phytophagous  Macropodidce.  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 


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  Dasyuridce.  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 


FIG.  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.  Vertebrae  ;  C  7,  D  13,  L  6,  S  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 
1  Geoffrey,  Bull.  Soc.  Philvm.  vol.  iii.  p.  249  (1803). 


PERAMELID&  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,  raffmyana,  and  longicaudata 
from  New  Guinea. 

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

Peragale.1 — 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.  Ears  very  large,  long,  and  pointed. 
Tail  long,  its  apical  half  clothed  on  the  dorsal  surface  with  long 
hairs  which  form  a  crest.  Vertebrae:  C  7,  D  13,  L  6,  S  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. 

Chceropus.2 — 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.  Tail  not  quite  so  long  as 
the  body,  and  covered  with  short  hairs  forming  a  slight  crest.  Ears 
large  and  pointed,  and  folded  down  when  the  animal 
is  at  rest.  Fur  soft  and  loose.  Vertebrae :  C  7,  D 
13,  L  6,  S  1,  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  Peramelidce.  It  was  first  described  as 
C.  ecaiidatus  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. 


PIG.  43.— Skele- 
ton of  right  hind 
foot  of  Chccropus 
castanotis.  c,  Cal- 
caneum  ;  a,  astra- 
galus ;  c6,  cuboid ; 
n,  navicular  ;  c3, 
ectocuneiform ;  II 
and  III,  the  con- 
joined second  end 
third  digits  ;  IV, 
the  large  and  only 
functional  digit  ; 
V,  the  rudiment- 
ary fifth  digit. 


Family  PHASCOLOMYID^E 


Dentition  :  c  •*-,  i  ft,  p  \,  m  %  =  24.  All  the  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 
equal,  stout,  and  short.  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 
bullae  of  the  skull  are  imperfect,  open  behind,  with  their  anterior 


PHA  SCOLOM  YID& 


145 


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


FIG.  44. — Chcaropus  castanotis.    From  Gould. 


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


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

Phascolornys.1 — The  existing  Wombats  (Fig.  45)  comprise  three 
1  Geoffrey,  Ann.  du  Afus6um,  vol.  ii.  p.  365  (1803). 
10 


MARSUPIALIA 


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

In  the  typical  group  we  find  the  following  characters.  Fur 
rough  and  coarse.  Ears  short  and  rounded.  Muffle  naked.  Post- 
orbital  process  of  the  frontal  bone  obsolete.  Ribs  fifteen  pairs. 
Vertebra:  C  7,  D  15,  L  4,  S  4,  C  10-12.  The  Wombat  of  Tas- 
mania and  the  islands  of  Bass's  Straits  (P.  ursinus)  and  the  closely 
similar  but  larger  animal  of  the  southern  portion  of  the  mainland  of 
Australia  (P.  mitchelli)  belong  to  this  group. 

In  the  second  group  the  characters  are  as  follows.  Fur  smooth 
and  silky.  Ears  large  and  more  pointed.  Muffle  hairy.  Frontal 
region  of  skull  broader  than  in  the  other  group,  with  well- 
marked  postorbital  processes.  Ribs  thirteen.  Vertebrae :  C  7,  D 
13,  L  6,  S  4,  C  15-16.  One  species,  P.  laiifrons,  the  Hairy-nosed 
Wombat  of  Southern  Australia. 

In  their  general  form  and  actions  the  Wombats  resemble  small 
bears,  having  a  somewhat  similar  shuffling  manner  of  walking,  but 
they  are  still  shorter  in  the  legs,  and  have  broader,  flatter  backs  than 
bears.  They  live  entirely  on  the  ground,  or  in  burrows  or  holes 
among  rocks,  never  climbing  trees,  and  feed  entirely  on  grass, 
roots,  and  other  vegetable  substances.  They  sleep  during  the  day, 
and  wander  forth  at  night  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  P.  ursinus  of  Tasmania,  is  a  brownish 
gray.  The  large  wombat  of  the  mainland  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  feet. 

It  is  noteworthy  that  P.  mitchelli  was  first  described  from  the 
evidence  of  fossil  remains,  the  living  form  subsequently  described  as 
P.  platyrhinus  being  found  to  be  indistinguishable.  Other  extinct 
species  occur  in  the  Pleistocene  of  Australia. 

Phascolonus.1 — Remains  of  a  large  extinct  Wombat,  which  must 
have  nearly  equalled  the  dimensions  of  a  Tapir,  occur  in  the 
Pleistocene  of  Queensland,  and  have  been  described  as  Phascolonus. 
It  is  probable  that  the  expanded  and  flattened  upper  incisors  from 
the  same  deposits  upon  the  evidence  of  which  the  presumed  genus 
Sceparnodon  was  founded,  are  likewise  referable  to  the  same  form. 
The  characters  of  both  the  upper  and  lower  incisors  distinguish 
Phascolonus  from  Phascolomys. 

1  Owen,  Phil.  Trans.  1872,  p.  257. 


PHALANGERID&  147 


Family  PHALANGERID.E. 

Dentition  extremely  variable,  owing  to  the  presence  of  minute 
rudimental  teeth  not  constant  in  the  same  species,  or  even  in  the 

two  sides  of  the  jaws  of  the  same  individual ;  exclusive,  however,  of 

g        j         /2 3)         (3 4) 

Tarsipes,  the  formula  i  p  c  ^,  p  }Q~^,  ™>  (3114)  represents  fairly  the 

general  condition  of  the  functional  teeth.  First  incisors  long  and 
stout ;  the  lower  pair  very  large  and  pointed,  but  withoiit  the  scissor- 
like  action  found  in  the  existing  Macropodidw ;  second  and  third 
lower  incisors  minute  and  probably  functionless.  Fourth  premolar 
generally  secant ;  milk-molar  generally  minute  and  deciduous  at  an 
early  period.  Molars  either  with  sharp  cutting-crests  or  bluntly 
tuberculate ;  fourth  sometimes  absent.  Mandible  without  pit,  and 
at  most  a  very  minute  perforation  in  the  masseteric  fossa.  Limbs 
subequal.  Fore  feet  with  five  distinct,  subequal  toes,  furnished  with 
claws.  Hind  feet  short  and  broad,  with  five  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.  Caecum  present  (except  in  Tarsipes),  and  usually  large. 
Pouch  complete.  Animals  of  small  or  moderate  size  and  arboreal 
habits,  usually  feeding  on  a  vegetable  or  mixed  diet,  inhabiting 
Australia  and  the  Papuan  Islands. 

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

Phalangers  are  small  woolly-coated  animals,  with  long,  power- 
ful, and  often  prehensile  tails,  large  claws,  and,  as  in  the  American 
opossums,  with  opposable  nailless  great  toes.  Their  expression 
seems  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 
insects,  and  all  relish,  when  in  confinement,  an  occasional  bird 
or  other  small  animal.  Several  of  the  Phalangers  possess  flying 
membranes  stretched  between  their  fore  and  hind  limbs  (Fig.  48), 
by  the  help  of  which  they  can  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  be  any  indication  of  special  affinity,  the  characters  of  the  skull 
and  teeth  sharply  dividing  the  flying  forms,  and  uniting  them  with 
other  species  of  the  non-flying  groups.  Their  skulls  (Fig.  47) 
are  as  a  rule  broad  and  flattened,  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  Dasyuridw   among  the 


Fio.  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  Tarsipedinae. — Cheek-teeth  almost  rudimentary  and 
variable  in  number.  Tongue  long,  slender,  pointed,  and  very  ex- 
tensile. Tail  long.  Caecum  absent. 

Tarsipes.1 — So  named  from  some  supposed  resemblance  of  its 
foot  to  that  of  the  Lemurine  genus  Tar  sins;  but  it  must  be  remarked 
that  it  has  none  of  the  peculiar  elongation  of  the  calcaneum  and 
navicular  so  characteristic  of  that  genus.  Head  with  elongated 

1  Gervais  and  Verraux,  Proc.  Zodl.  Soc.  1842,  p.  1. 


PHALANGERID^E  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  i  y,  c  ^,  p  and  m  %  on 
one  side,  and  -£  on  the  other;  total  20.  Kami  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  Phalangerince,  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.  Vertebrae :  C  7, 
D13,  L  5,  S  3,  C  24. 

Of  this  singular  genus  but  one  species,  T.  rostmtus  (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  Melaleucce,  etc.  One  kept  in  confinement  by  Mr. 
Gould  was  also  observed  to  eat  flies. 

Subfamily  Phalangerinse.  —  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.  Caecum  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.1 — The  typical  genus  Plialanger  (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  mammae.  Skull  (Fig.  47) 

1  Storr,  Prodromus  Meth.  Mamm.  p.  33  (1780).     Syn.  Phalangista,  Geoffrey, 
Huff.  Soc.  Philom.  vol.  i.  p.  106  (1796). 


MARSUPIALIA 


FIG  47. — Left  lateral  view  of  skull  of  Gray  Cuscus  (Phal- 
anger  orientalis).    After  Peters. 


stout  and  strong,  with  large  vacuities  in  the  hinder  half  of  the 
palate,  and  the  auditory  bullae  thick  and  inflated.    Dentition  usually 

*  f  >  c  ^>  P  f  >  m  $•     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. 
Molars  four-cusped, 
with  the  inner  cusps 
of  the  upper  ones 
crescentoid,  and  imperfect  transverse  ridges  connecting  each  pair 
of  cusps. 

The  Cuscuses  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. 

Trichoswus.1 — The  members  of  the  genus  Trichosurus  are  of 
relatively  large  size,  and  are  distinguished  from  PJialanger  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 

*  f  >  c  ir>  P  $ j  m  T-     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  Phalanger. 

The  true  Phalangers  comprise  two  species,  of  which  the  best 
known   is   the  Vulpine  Phalanger  (T.  vulpecula),   so    common    in 
1  Lesson,  Diet.  Class.  d'Hist.  Nat.  vol.  xiii.  p.  333  (1828). 


PHALANGERID&  \  5 1 


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  (car.  fuliginosa).  Its  habits  are  very  similar  to  those  of 
the  Yellow-bellied  Fly  ing  -Phalanger  (Petaurus  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. 

Pseudochirus.1  —  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  mammae.  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  bulla?  generally 

somewhat  inflated.     Dentition  (at  most)  i  —  ~,  c  —Q-^,p  Q>  m  z- 

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  King-tailed  Phalanger  (Pseudochirus  peregiinus), 
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  Phascolardus  ;  according,  however,  to  Mr. 
0.  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 


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


152  MARSUPIALIA 


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.1 — With  the  genus  Petauroides,  containing  only  the 
single  species  P.  volans,  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  fore-arm  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,  arid  not  extending  nearly  as 
far  forwards  as  the  premaxillse.  Large  vacuities  in  hinder  part  of 
palate.  Auditory  bullae  inflated  and  smooth.  Dentition  usually 
*  f >  c  iy>  P  f>  m  f •  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* — The  genus  Dadylopsila  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  f ,  c  ^,  p  f ,  m  £. 
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.  trivirgata,  or  Striped  Phalanger,  inhabits  the 

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


PHALANGERIDjE  153 


Papuan  and  North  Australian  sub -region ;  a  second  species  (D. 
palpator),  characterised  by  the  still  greater  elongation  of  the  fourth 
finger,  occurring  in  South  New  Guinea.  These  animals  are  said 
to  be  of  insectivorous  habits,  the  elongated  fourth  finger,  as  in  the 
analogous  instance  of  the  Lemuroid  genus  Chiramys,  being  appar- 
ently specially  adapted  for  extracting  insects  and  larvae  from  their 
hiding  places. 

Petaurus.1 — 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 
regularly  in  length  from  the  first  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 
vacuities  near  the  second  molars.  Auditory  bullse  inflated,  and 
variable  in  size.  Dentition  :  i  f ,  c  £,  p  f ,  m  £.  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  Mr.  Gould  as  follows.  "This  animal  is 
common  in  all  the  brushes  of  New  South  Wales,  particularly  those 
which  stretch  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-failing  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  night  while  running  over  the 
small  leafy  branches,  frequently  even  to  their  very  extremities,  in 
search  of  insects  and  the  honey  of  the  newly-opened  blossoms.  Its 
structure  being  ill  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  forced  to  flight  it 
ascends  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." 

1  Shaw,  Naturalist's  Miscellany,  vol.  ii.  pi.  Ix.  (1791). 


154 


MARSUPIALIA 


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

Gymnobelideus.1 — Like  Petaums  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.  leadbeateri  of  Victoria,  and  according  to  Mr.  Thomas, 
may  be  regarded  as  the  primitive  form  from  which  the  specialised 
Petaums  has  been  developed. 


FIG.  48. — Squirrel  Flying-Phalanger  (Petaunis  sciureus). 

Dromicia.- — 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  scaly, 
with  fine  hairs,  except  at  the  tip,  which  is  naked  and  prehensile. 

1  M'Coy,  Ann.  Mag.  N.  H.  (3)  xx.  p.  287  (1867). 
2  Gray,  in  Grey's  Australia,  appendix,  vol.  ii.  p.  407  (1841). 


PHALANGERID&  155 

Skull  short  and  broad,  with  the  hinder  part  of  the  palate  in- 
complete, and  the  auditory  bullae  large,  much  inflated,  and  trans- 

010  /o A\ 

parent.    Dentition  :  i  -^  c  -,  p  g,  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  Petaurus  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. 

Distcechunis.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  Acrolates, 
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.  Fjars  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  mammae.  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  maxillae.  Auditory  bullae  low,  rounded,  and 
but  slightly  prominent.  Dentition :  i  f ,  c  ^,p  f ,  m  f .  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  (A.  pygmceus),  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  Phaseolaretinae. — Teeth  large,  normal;  no  rudi- 
mentary premolars  before  the  last  upper  premolar,  or  any  teeth 

1  Peters,  Ann.  Mus.  Genov.  vol.  vi.  p.  303  (1874). 
2  Destnarest,  Nouv.  Diet.  d'Hist.  Nat.  ser.  2,  vol.  xxv.  p.  405  (1817). 


1 56 


MARSUPIALJA 


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  Phascolomyulce.1 

Pluiscolardus? — Dentition:  if,  c%,p  ^,  ra£ ;  total  30.  Upper 
incisors  crowded  together,  cylindroidal,  the  first  much  larger  than 
the  others,  with  a  bevelled  cutting  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- 
brae :  C  7,  D  11,  L8,  S2,  C8. 
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 
the  Australian  colonists  (P.  cin- 
ereus),  an  animal  of  compar- 
atively large  size  and  heavy 
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  Of.  W.  A.  Forbes,  "Anatomy  of  the  Koala,"  Proc.  Zool.  Soc.  1881,  p.  180. 
2  Blainville,  Bull.  Soc.  Philom.  1816,  p.  116. 


Fio.  49.— Skeleton  of  right  hind  foot  of  Koala 
(Phascolarctus  cinereiis),  showing  the  stout  op- 
posable hallux,  followed  by  two  slender  toes, 
which  in  the  living  animal  are  enclosed  as  far 
as  the  nails  in  a  common  integument. 


PHALA  NGERID.'E 


'57 


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  Phalangeridce,  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  Arcliizonurus  which  appears  to  be 


FIG.  50.— The  Koala  (I'hascolarctus  cinereus).    From  Sclater,  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:  i%,  c£,  p%,  m^;  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,  T.  carnifex.  This  animal  presents  a  most  anomalous 

1  Owen,  in  Gervais's  Zool.  et  Pal.  franqaiscs,  1st  ed.  pt.  i.  p.  192  (1849-52). 


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 
Phalangeridce  and 
Macropodidce,  and 
its  dentition  com- 
pletely unlike  that 
of  any  known  pre- 
daceous  animal,  this 
view  has  been  called 
in  question. 

The  dentition  is 
nearer  to  that  of  the 


PIG.  51. — Front  view  of  skull  of  Thylacoleo  carnifex,  restored. 
J  natural  size.     From  Quart.  Journ.  Geol.  Soc.  vol.  xxiv.  p.  312. 


existing  Phalangeridce  than  to  that  of  the  Macropodidce,  and  the 
genus  may  be  provisionally  regarded  as  the  type  of  a  distinct 
subfamily  of 'the  former. 


Family  MACROPODID^E. 

Dentition  i  -,   c  ^  ~~   ,  p  ~,  m  -•      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,  and 
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, 


MACROPODIDsE 


159 


occasionally    prehensile ;   stomach    sacculated.      Pouch    large    and 
opening  forwards. 

The  Macropodidas  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  by  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 
noticed  in  some  detail.  It  is  extremely  long  and 
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, 


FIG.  52.— Skeleton 
of  right  hind  foot  of 
Kangaroo. 


i6o 


MARSUPIALIA 


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  being  no  tusks  or  canines  for  offensive 
or  defensive  purposes. 

The  number  of  vertebrae  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  of  motion  of  the  hand.  The  pelvis  has  large  epipubic  or 
"  marsupial "  bones.  The  femur  is  short,  and  the  tibia  and  fibula 


FIG.  53.— The  Great  Gray  Kangaroo  (Macropus  giganteus). 

are  of  great  length,  as  is  the  foot,  the  whole  of  which  is  applied  to 
the  ground  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  of 
sacculi,  like  those  of  the  human  colon.  The  alimentary  canal  is 
long,  and  the  caecum  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  sojourn  in  the  pouch, 


MA  CROPODID^E  \  6 1 


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  Macropodidce,  a 
few  words  are  necessary  in  respect  of  the  tooth-change,  and  we  may 
here  quote  the  observations  of  Mr.  0.  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  Macropodidce,  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 


1 62  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  be,  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  m  3  and  not  m  4.  The  first  plan  is, 
as  a  rule,  the  better,  since  p  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  Macropodidce  are  divided  into  three  well-marked  sections : 
(1)  the  true  Kangaroos  (Macropodince) ;  (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  Hypsiprymnodontinse. — 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  Potoroince. 

This  subfamily  is  now  represented  only  by  the  genus  Hypsi- 
prymnodon,1  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  nudicaudalus.  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  Macropodidce. 

Triclis.2 — 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  Macropodidce,  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 
Phalangeridce  and  Macropodidce  than  any  other. 

Subfamily  Potoroinse. — The  second  section  or  subfamily,  the 
Potoroince,  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.  JV.  S.  Wales,  vol.  i.  p.  33  (1876). 
2  De  Vis,  Proc.  Roy.  Soc.  Queensland,  ser.  2,  vol.  iii.  p.  8  (1888). 


MACROPODIDJL 


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 


pin- 


FIG.  54.— Skull  and  Teeth  of  Rat  Kangaroo  (Bettongui  lesueuiri).   c,  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.1 — Head  long  and  slender.  Auditory  bullae  some- 
what inflated.  Ridges  on  premolars  few  and  perpendicular. 
Large  palatine  foramina.  Tarsus  short.  Muffle  naked.  Three 
species,  viz.  P.  tridadylus,  P.  gilberti,  and  P.  platyops ;  the  last  two 
being  confined  to  West  Australia. 

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

1  Desmarest,  Nouv.  Diet.  d'Hist.  Nat.  ser.  1,  vol.  xxiv.    Table  Meth.  p.  20 
(1804).     Syn.  Hypsiprymnus,  Illiger,  Prodromus  Syst.  Mamm.  p.  79  (1811). 
-  Gray,  CharlcswortK  s  Mag.  Nat.  Hist.  vol.  i.  p.  584  (1837). 


1 64 


MARSUP1ALIA 


and  obliqiie.  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.1 — Muffle  naked,  as  in  Bettongia,  but  the  edge  of  the 
hairy  part  less  emarginate  backwards  in  the  middle  line.  Ears 
short,  rounded,  and  hairy.  Auditory  bullae  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.  campestiis  of  South  Australia,  originally  referred  to  Bettongia. 


FIG.  55.— Skull  and  Teeth  of  the  Red-necked  Wallaby  (Macropus  ruficollis).  i\  &,  (3,  First, 
second,  and  third  upper  incisors  ;  pm,  fourth  or  posterior  premolar  (the  penultimate  or  third 
having  been  already  shed) ;  ml,  m?,  m?,  m*,  the  four  true  molars.  The  last,  not  fully  de- 
veloped, is  nearly  concealed  by  the  ascending  ramus  of  the  jaw. 

jtEpyprymnus? — Head  short  and  broad.  Auditory  bullae  not 
inflated.  No  palatine  foramina.  Tarsus  long.  Muffle  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  j£.  rufescens. 

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

Subfamily  Maeropodinse. — 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). 
2  Garrod,  Proc.  Zool.  Soc.  1875,  p.  59. 


MACROPODWsE  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  : 

Lagostrophiis.1 — Represented  only  by  the  Banded  Wallaby 
(L.  fasciatus)  of  Western  Australia,  which  presents  the  following 
distinctive  features.  Size  small.  Muffle  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  bullae  ;  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  bullse  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.  lumholtzi  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. 
2  Schlegel  and  Miiller,  Verh.  Nat.  Ges.  Nederland,  p.  138  (1839-44). 


1 66  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.1 — Hind  limbs  relatively  less  large  than  in  Macropus. 
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  bullse  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  madeayi 
connects  the  other  species  with  Macropus. 

Lagorchestes.2 — Muffle  entirely  or  partially  covered  with  hair. 
Fourth  hind  digit  with  a  long  claw,  not  concealed  by  hair.  Tail 
rather  shqrt,  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 — M.  leporoides,  M. 
hirsutus,  M.  conspicillatus. 

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

Onydwgale? — Muffle  hairy.  Fourth  hind  claAv  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  bullse  more  or  less  inflated.  Canine 

1  Schlegel  and  Miiller,  Verh.  Nat.  Ges.  Ncdcrland,  p.  130  (1839-44). 

2  Gould,  Monograph  of  Macropodidce,  pi.  xiii.  (1841). 
3  Gray,  in  Grey's  Australia,  vol.  ii.  appendix,  p.  402  (1841). 


MACROPODID^E  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  Lagarchestes.  Mr.  0.  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.1 — 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  bullse  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  Macropus.  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? — Muffle  generally  completely  naked.  Ears  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  mammae.  Skull  (Fig.  55)  long, 
smooth,  and  rounded  ;  the  nasals  expanded  behind  ;  generally  large 
palatal  vacuities ;  and  the  auditory  bullae  not  inflated.  Canine 
minute,  and  shed  at  an  early  period.  Incisor  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,  CharleswortK s  Mag.  Nat.  Hist.  vol.  i.  p.  583  (1837). 
2  Shaw,  Naturalist's  Miscellany,  vol.  i.  pi.  xxxiii.  (1790). 


168  MARSUPIALIA 


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  Eabbit 
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  same  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  broiight 
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  gigantea  for  a  specific  designation, — the  latter 
having  been  bestowed  upon  it  by  Zimmermann  under  the  impression 


MACROPODID^E  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  Macropus,  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  M.  robustus.  The  fourth  premolar  is 
of  large  or  medium  size  in  these  forms.  Remains  of  M.  giganteus 
occur  fossil  in  the  Pleistocene  of  Australia,  where  we  also  find  the 
allied  extinct  M.  titan,  which  attains  somewhat  larger  dimensions, 
M.  robustus  also  dates  from  the  same  geological  epoch,  where  it  was 
accompanied  by  two  allied  types  known  as  M.  altus  and  M.  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  M.  rujicollis,  M.  ualabatus,  M.  parryi,  and  M.  agilis 
are  the  best  known. 

M.  ualabatus  and  M.  parryi  are  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  M.  rcechus,  and  M.  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.1 

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  Mammals  of  Australia  (1877),  and  Lydekker's  Catalogue  of 
Fossil  Mammalia  in  the  British  Museum,  pt.  v.  (1887). 


170  MARSUPIALIA 


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.  M.  brachyurus  is  remarkable  for 
its  comparatively  short  and  slender  tail  and  small  ears.  The  earliest 
known  species  of  Kangaroo,  referred  to  before,  M.  bmni,  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  (Beizen  <>rer 
Moskome,  etc.)  under  the  name  of  "Filarider."  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- 
idce,  to  which  brief  reference  may  now  be  made.  The  first  of  these 
is  Sthenurus,1  represented  by  a  single  large  species  (S.  atlas),  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  2 
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  Palorchestes  3  (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.  3  Owen,  op.  cit.  p.  797. 


EXTINCT  FAMILIES 


171 


EXTINCT  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  Macropodidce  and  the  Phalangeridce  in  having 
J  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.1 — Dentition  :  i  f ,  c  $,  p  ^,  m  £ ;  total  28.  The  first 
upper  incisor  very  large  and  scalpriform  (Fig.  56).  True  molars 
Avith  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 


FIG.  56. — Left  lateral  aspect  of  the  skull  of  Diprotodon  australis;  from  the  Pleistocene  of 
Australia.     ^  natural  size,    i,  Incisors;  p,  premolar;  TO,  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 — 
Diprotodontidce — having  affinity  on  the  one  hand  with  the  Phalangers 
and  on  the  other  with  the  Kangaroos. 

Nototherium.* — 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  MitclielVs  Eastern  Australia,  2d  ed.  vol.  ii.  p.  362  (1838). 
-  Owen,  Cat,  Mamin.  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  NototJieriidce  may  apparently  be  regarded  as  a 
distinct  family  connecting  the  Diprotodontida  with  the  Phasco- 
lomyidce  and  Phalangeridce. 

Bibliography  of  Marsupialia. — G.  R.  Water-house,  Nat.  Hist,  of  tJic  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  ;  0.  Thomas,  "On  the  Homologies  and  Succession 
of  the  Teeth  in  the  Dasyurids,"  Phil.  Trans.  1887  ;  and  "Catalogue  of  Mar- 
supialia and  Monotremata  in  the  British  Museum,"  1888. 


CHAPTER  VII 

THE  SUBCLASS  EUTHERIA  AND  THE  ORDER  EDENTATA 

THE  whole  of  the  remaining  groups  of  mammals  are  included  in  a 
single  subclass,  known  by  the  names  Eutheria,  Monodelphia,  or 
Placentalia.1  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  fcetus  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 
habite  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 
palaeontology  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  palaeontology  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  Hyracidce,  and  in  their  case  palaeon- 
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  our  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  Toxodon,  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 
Ta3niodontia)  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  palseontological 
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 l  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 
Linnaean  name  Bruta  is  occasionally  substituted.  But  that  term  is 
quite  as  objectionable,  especially  since  the  group  to  which  Linnaeus 
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  Natures.  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  been  suggested  for  this  proposed  subclass. 


GENERAL   CHARACTERS 


177 


remarkable  exceptions  of  Tatusia  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  Onjcteropus 
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  to  this  order 
that,  beyond  the  negative 
characters  just  mentioned, 
there  seems  little  to  connect 
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. 
Palaeontology  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 


FIG.  57. — Upper  surface  of  the  brain  of  the  Broad- 
banded  Armadillo  (Xenurus  ttnicinctus).  The  large 
olfactory  lobes  are  seen  at  the  anterior  extremity 
(left  of  figure) ;  the  hemispheres  have  only  three 
sulci.  (From  Garrod,  Proc.  Zool.  Soc.  1878,  p.  230.) 


1 78  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)  Bradypodidce,  or  Sloths;  (2)  Myrmecophagidce,  or  Ant- 
eaters  ;  (3)  Dasypodidce,  or  Armadillos ;  (4)  Manidce,  Pangolins  or 
Scaly  Anteaters ;  and  (5)  Orycteropodidce,  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 
Bradypodidce  alone;  and  (2)  the  Entomophaga,  insect-eaters,  or 
Vermilingua,  containing  all  the  other  families,  from  which  some- 
times the  Orycteropodidce  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  Manidce  to  the  Myrmecophagidce  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  Chamseleon  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  animals1  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 
Bradypodidce,  Megatlienidce,  and  Myrmecophagidce  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. 


BRA  D  YPODIDA:  \  79 


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  (Dasypodidce) 
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  Manidce  and  Orycteropodidce  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  Orycteropodidce.1 


Family  BRADYPODID^E. 

Externally  clothed  with  long,  coarse,  crisp  hair.  Head  short 
and  rounded.  External  ears  inconspicuous.  Teeth  f  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.  Tail  rudimentary.  Stomach  complex.  No 
caecum.  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  PILOSA. 

Brady 'podidce. 

Megatfieriidce, 

MyrmecopJiagidoe. 
Suborder  LORICATA. 

Dasypodidoe. 
Suborder  SQUAMATA. 

Manidce. 
Suborder  TUBULIDEMTATA. 

Orycteropodidce. 

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,"  Proe.  Zool.  Soc.  1883,  p.  178. 


i8o  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 


Fio.  58.— Two-toed  Sloth  (Cholaepus  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 


BRA  D  YPODID^E  \  8 1 


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  Bradypodidce,,  as  well  as  in  the  Myrmecophagidce,  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  Ckofaptu  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  Linnsean  genus  Bradypus, 
but  Illiger  very  properly  separated  the  species  with  but  two  claws 
on  the  fore  feet,  under  the  name  of  Cholcepus,  leaving  Bradypus 
for  those  with  three. 

Bradypus}- — Three-toed  Sloths.  Teeth  usually  |-  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.  Vertebrae :  C  9,  D  and  L  20  (of  which  15  to  17  bear  ribs), 
S  6,  C  1 1 .  All  the  known  species  present  the  remarkable  pecu- 
liarity of  possessing  nine  cervical  vertebrae,  i.e.  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). 


1 82  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  mammse  are  two,  and  pectoral  in 
position. 

"  Ai "  is  the  common  name  given  in  books  to  the  Three-toed 
Sloths.  They  were  all  comprised  by  Linnaeus  under  the  species 
Bradypus  tndadylus.  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.  torquatus) ;  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.  infuscalus  and  B.  castaneiceps)  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. 

Cholcepus.1 — 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), 

1  Illiger,  Prodromus  Syst.  Mamm.  et  Avium,  p.  108  (1811). 


MEGA  THER1ID& 


unlike  the  true  canines  of  heterodont  mammals.  Vertebrae  :  C  6 
or  7,  D  2 3-24,  L  3,  S  7-8,  C  4-6.  One  species  (C.  didactylus)  has 
the  ordinary  number  of  vertebrae  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.  didadylus,vf}iich 
has  been  longest 
known,  is  com- 
monly called  by 
the  native  name 
of  Unau.  It  in- 
habits the  forests 
of  Brazil.  C.  hoff- 
manni  (Fig.  58) 
has  a  more  north- 
ern geographical 
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  Choloepus  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^E. 

The  members  of  this  family  are  all  extinct.  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 
Bradypodidce  and  the  Myrmecophagidce,  combining  the  head  and 
1  Burmeister,  Sitzb.  Ak.  Berlin,  vol.  xxviii.  p.  613  (1882). 


FIG.  59. — Skull  of  Two-toed  Sloth  (Cholcepus  didactylus).    From 
Proc.  Zool.  Soc.  1871,  p.  432. 


1 84 


EDENTA  TA 


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,  Nothrotherium  escrivanense, 
exceeding  the  largest  existing  Anteater,  and  the  Megatherium 
being  larger  than  a  Ehinoceros.  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  $  in  Nothrotherium.1  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 


FIG.  60. — Section  of  upper  molar  teeth  of  Megatherium  ameriwnum.     x  J. 
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  grinding 
surface  the  prominent  ridge  of  hard  dentine  follows  the  external 
contour,  and  is  surrounded  only  by  a  thin  layer  of  cement,  as 
in  the  existing  Sloths.  The  Ground  Sloths,  as  the  members 

1  Lydekker,  in  Nicholson  and  Lydekker's  Manual  of  Palaeontology,  vol.  ii. 
p.  1299  (1889).     Originally  described  under  the  preoccupied  name  Ccelodon. 


MEGA  THERIID^E 


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.1 — The  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 


Fio.  61. — Oral  surface  of  mandible  of  Megatherium,  americanum. 
a,  Condyle  ;  b,  massetcric  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  (M.  cuvieri  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,  Tableau  EUm.  d'Sist.  Nat.  des  Animaux,  p.  146  (1798). 

2  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. 


i86 


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  M.  mirabile. 
Three  other  South  American  species  have  been  described ;  but  M. 
laurillardi,  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 


FIG.  62. — Skeleton  of  Megatherium,  from  the  specimen  in  the  Museum  of  the  Royal  College 
of  Surgeons,     x  "A- 

superior.  The  full  length  of  a  mounted  skeleton  (Fig.  62),  from 
the  fore  part  of  the  head  to  the  end  of  the  tail,  is  1 8  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  premaxillse  exhibited  in  Fig.  62,  there  was 
a  prenasal  bone,  towards  which  a  process  extended  upwards  and 


MEGATHERI1D&  187 

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

The  vertebral  column  consists  of  seven  cervical,  sixteen  dorsal, 
three  lumbar,  five  sacral,  and  eighteen  caudal  vertebrae.  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  calcaneum  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 


1 88  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  wrhole  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  Mylodon.  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  Myrmecophagidce.  The  humerus  generally  has  an  ent- 
epicondylar  foramen ;  and  the  form  and  relations  of  the  bones  of 


MEGA  THERIID& 


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  Subtrian0!!-  Fl°'  63-~skeleton  of  Mylodon  robvstus  (Pleistocene,  South 

America).    From  Owen. 

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  M.  (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  Megatherium. 
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.  harlani,  from  beds  of  corre- 
sponding age  in  Kentucky.  The  Patagonian  M.  (Grypotherium) 
darwini  is  a  remarkable  form,  characterised  by  the  presence  of  a 
bony  arch  connecting  the  premaxillse  with  the  nasals,  of  which,  as 
already  mentioned,  there  is  an  incomplete  development  in 
Megatherium.  Megalonyx,  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.  Nothrotherium  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 


190  EDENTATA 

of  Argentina,  described  as  Promegatherium  and  Promylodon,  are 
respectively  distinguished  from  Megatherium  and  Mylodon  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  Moropus  and 
Morotherium,1  and  originally  regarded  as  Edentates,  would  appear  to 
be  aberrant  Ungulates. 


Family  MYRMECOPHAGID^E. 

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 
vertebrae,  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  excellence.  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  Bradypodidce. 

Myrmecvphaga.2 — Skull  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  maxillae,  and  the  long 
and  narrow  nasal  bones,  the  premaxillae  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,  or  any 
other  demarcation  between  the  orbits  and  the  temporal  fossae.  Palate 
extremely  elongated,  and  produced  backwards  as  far  as  the  level  of 

1  See  E.  D.  Cape,  Amer.  Naturalist,  vol.  xxiii.  p.  \52  (1889).   . 
2  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  51  (1766). 


MYRMECOPHAGID^E  191 

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.  Vertebrae  :  C  7,  D  15-16,  L  3-2,  S  6,  C  31. 
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  caecum.  Mammas  two, 
pectoral. 

There  is  one  species,1  M.  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  iu  support  of  this  view. 


1 92 


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  (Tatusia),  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 


FIG.  64.— The  Great  Anteater  (Myrmecophagajubata).    (From  Sclater,  List  of  Animals  in 
Zoological  Society's  Gardens,  1883,  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  Myrmecophaga,  but  Avith  the  muscular 
pyloric  gizzard  not  quite  so  strongly  developed.  There  is  a  distinct 
ileo-colic  valve  and  a  short  globular  caecum.  The  fore  foot  has  a  very 
large  claw  on  the  third  toe,  moderate-sized  claws  on  the  second  and 

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


MYRMECOPHA  GID^E 


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.  Vertebrae :  C  7,  D  17,  L  2,  S  5,  C  37.  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 


FIG.  65.— Tamandua  Anteater  (Tamandua  tetradactyla).    From  Prof.  Zool.  Soc.  1871,  pi.  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.  Vertebrae:  C  7,  D  16,  L  2,  S  4,  C  40.  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 

foregoing  genera.  Commence- 
ment of  the  colon  provided  with 
two  small  caeca  (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 
and  silky. 

This  genus  has  also  but  one 
species  certainly  known,  the  Little  or  Two-toed  Anteater  ((7.  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. 


Fio.  66.—  C#ca  of  the  Two-toed  Anteater 
(Cycloturus  didaetylus).    i,  Ileum  ;  c,  colon. 


Family  DASYPODID^E. 

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  inner 
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  ossified 
dermal  scutes  are  everywhere  covered  by  a  layer  of  horny  epi- 
dermis. Teeth  numerous,  simple,  of  persistent  growth,  and  usually 


DASYPODID&  195 

monophyodont,  but  in  one  genus  (Tatusiii)  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  vertebras 
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  Tatusia  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  Dasypodidce  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  s,till  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  Chlamydophorinse. — In  most  anatomical  characters, 
especially  the  structure  of  the  fore  foot,  this  little  group  resembles 
the  Dasypodina ;  but  it  differs  remarkably  from  all  other  known 
Armadillos,  living  or  extinct,  in  the  peculiar  modification  of  the 
dermal  armour. 

8 
Chlamydophorus}- — Teeth     — ,    subcylindrical,    somewhat    com- 

o — y 

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.  Vertebrae :  C  7, 
D  11,  L  3,  S  10,  C  15.  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.  Eyes  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  Piiodon.  Nipples 
two,  pectoral.  Visceral  anatomy  closely  resembling  that  of  Dasypus, 
the  caecum  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 

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


DASYPODID&  i97 

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 
skiu  of  the  back  as  far  as  its  edge,  instead  of  only  along  the  median 
line. 

Subfamily  Dasypodinse. — 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.  Mammae  one  pair, 
pectoral. 

Dasypus.1 — Teeth  ^  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  Dasypodince  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.  Vertebras  :  C  7,  D  11-12,  L  3,  S  8,  C  17-19.  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  (D.  sexcindus)  of 
Brazil  and  Paraguay.  A  very  similar  species,  D.  villvsus,  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. 

Xenurus.1 — Teeth  -|  or  |,  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.  Vertebra:  C  7,  D  12-13,  L  3,  S  10,  C  18. 
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,  A". 
unicinctus,  is,  after  Piiodon  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.z — 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.  Amphibian,  etc.,  p.  36  (1830). 
2  F.  Cuvier,  Hist.  Nat.  des  Mammifires  (1822). — Priodontes. 


DASYPODIDsE  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. 
Vertebrae:  07,  D  12,  L  3,  S10,  C  23.  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.1 — Teeth  -|  or  -|,  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.  Vertebrae:  C  7,  D  11,  L  3,  S  12, 
C  13.  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.  conunis,  the 
1  Illiger,  Prodromus  Syst.  Mamm.  et  Avium,  p.  Ill  (1811). 


200  EDENTA  TA 

Matico ;  and  T.  muriei.     Eemains  apparently  referable  to  T.  conums 
are  of  not  uncommon  occurrence  in  the  Brazilian  cavern-deposits. 

Subfamily  Tatusiinae. — This  group  contains  but  one  genus, 
Tatusia.1  Teeth  -|  or  f ,  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  Oryderopus)  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.  Vertebrae  :  C  7,  D  9-11,  L  5,  S  8,  C  20-27.  Head  narrow, 
with  a  long,  narrow,  subcylindrical,  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. 


FIG.  67. — The  Peba  Armadillo  (Tatusia  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  of 
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,  obtusely-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  mammae,  in  addition  to  the  usual  pectoral  pair,  and  in 

1  Lesson,  Man.  de  Mammalogie,  p.  309  (1827) ;  ex.  F.  Cuvier,  Tatusie. 


DASYPODID&  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,  T,  novemcinda  (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- 
phradus  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 
T.  novemcinda,  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.1 

In  the  Pleistocene  cavern -deposits  of  Brazil  have  been  found 
remains  of  T.  novemcincta,  and  also  of  T.  pundata,  which  appears  to 
be  an  extinct  form  nearly  allied  to  T.  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.  Sc.  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 


E  DENT  A  TA 


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


Family  GLYPTODONTID.E. 

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 
Dasypodidce  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  £  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 
Texas  and  Mexico.  Many  species  of  the  family 
have  been  described  and  figured,  especially  by 
Burmeister  (in  the  Annales  del  Museo  publico  de 
Buenos  Aires),  among  which  the  following  may 
be  noticed.  Hoploplwr^us  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  long 
subcylindrical  terminal  tube  similar  to  the  one  represented  with  the 
carapace  of  Glyptodon  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 


FIG.  68.—  Tooth  of  Glyp. 
todon  from  the  side,  and 
from  the  grinding  surface. 
(After  Owen.) 


GL  YPTODONTID& 


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.  Doedicurus,  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, 


FIG.  69.—  Glyptodon  davipes  (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  Panochthus,  probably  carried  horny  spines  in  the  living 
condition.  The  typical  genus  Glyptodon  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  TJwracophorus, 


204  EDENTA  TA 

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 

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  vertebne,  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  vertebrae  is  a  character  in  which  this  and  the  following 
family  differ  from  all  the  preceding  forms. 

Manis.1 — Skull  somewhat  of  the  form  of  an  elongated  cone,  with 
the  small  end  turned  forwards  ;  very  smooth  and  free  from  crests 
and  ridges.  No  distinction  between  the  orbits  and  temporal  fossae. 
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.  Kami  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  vertebrae  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). 


MANID&  205 


secreting  follicles,  the  ducts  of  which  terminate  in  a  common 
orifice.  No  caecum.  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  Manis  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  foatal 
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.  0.  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  betAveen  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 


2o6 


EDENTA  TA 


as  an  organ  of  touch.  There  are  three  species,  viz.  Manis  javanica, 
ranging  from  Burma,  through  Malacca  and  Java,  to  Borneo ;  M. 
awita,  found  in  China,  Formosa,  and  Nipal ;  and  the  common  Indian 
Pangolin,  M.  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 

IV  J  group  :  —  the 

Long-tailed  Pan- 
golin (M.  mac- 
rura),  which  has 
a  tail  nearly  twice 
as  long  as  its 
body,  and  con- 
taining as  many 
as  forty-nine 
caudal  vertebras, 
being  the  largest 
number  known 
among  mammals ; 
the  White-bellied 
Pangolin  (M.  tri- 
cuspis),  Fig.  70, 
closely  allied  to 
the  last,  but  with 
longer  and  tri- 
cuspid  scales,  and 
white  belly  hairs. 
These  two,  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  (M. 
temmincki  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,  M.  temmincki,  extending  also  into  south 
and  eastern  equatorial  Africa. 

According  to  Professor  W.  K.  Parker,1  who  remarks  upon  the 

peculiarly  aberrant  nature  of  the  group,  the  horny  scales  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 

1  Mammalian  Descent,  p.  95. 


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


MANIDJE  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  observes1  that  there  are  occasional  in- 
stances of  the  presence  of  eight  cervical  vertebrae  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  Manis  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. 


2o8  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  M. 
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. 

Palceamanis.1 — 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  M.  gigantea,  upon  the 
evidence  of  which  the  genus  Palceomanis  has  been  established. 


Family  ORYCTEROPODID.E 

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  vertebrae  with  no  accessory  zygapophyses. 
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. 

Oryderopus.2 — 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.  cvii.  p.  1180  (1888). 
2  Geoffroy,  Decade  Philosophique,  1795  (teste  Agassiz). 


ORYCTEROPODIDM  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.  Vertebrae:  C  7,  D 13,  L  8,  S6,  027.  The  large 
number  of  lumbar  vertebrae  is  peculiar  among  Edentates.  Tongue 
less  vermiform  than  in  Myrmecophaga,  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-csecal  valve,  and  a  considerable-sized  csecum ; 
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  Oryderopus  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  Manis,  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  Manis  by  the  abortion  of  the  foetal  villi  at  the  two  poles 
of  the  ovum. 

The  Orycteropodidce  have  long  been  regarded  as  widely  different 

from  other  Edentates,  their   presumed  affinity  with  the  Manidce 

being  more  or  less  problematical ;  but  the  discovery  recently  made 

by  Mr.  0.  Thomas l  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  Tatusia),  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  palaeontological 
knowledge  of  the  extinct  allies  of  either.  Macrotherium  even, 
usually  supposed  from  the  structure  of  its  phalangeal  bones,  to  be 
related  to  Manis,  has  lately  proved  to  have  the  teeth  and  vertebrae 
of  a  perissodactyle  Ungulate,  and  one  could  not  dare  to  suggest 
that  ancestors  of  Manis,  or  Orycteropus  were  to  be  sought  in  that 
direction.  Lastly,  as  the  numerous  fossil  American  Edentates  do 


OR  YCTEROPODW^E  2 1 1 

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.  cethiopicus)  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,  differing  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  iibcr  die  Edentaten,  2d  ed.  1852). 
Among  numerous  memoirs  on  special  groups  the  following  may  be  cited : — 
Myrmecophagidce : — R.  Owen,  "Anatomy  of  Great  Anteater,"  Trans.  Zool.  Soc. 
vol.  iv.  ;  G.  Pouchet,  Mem.  sur  le  Grand  Fourmilier,  1874 ;  W.  A.  Forbes, 
"Anat.  of  Great  Anteater,"  Proc.  Zool.  Soc.  1882,  p.  287.  Megatlieriidce : — R. 
Owen,  Extinct  Gfigantic  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  Rtpublique  Argentine,  t.  iii.  Mammiferes,  1879, — which  contains 
full  references  to  various  memoirs  by  Owen,  Gervais,  Reinhardt,  and  others. 
Glyptodontidce : — 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  Ripublique 
Argentine,  1879  ;  H.  Gervais  and  F.  Ameghino,  Les  Mammiferes  Fossiles  de 
I'Amtrique  M6ridionale,  Paris,  1880, — which  also  contains  a  list  of  all  the 
S.  American  Edentates  described  at  that  date.  Dasypodidce : — 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  cerebrales  des  Edeutes,"  Nouv.  Arch,  du  Museum,  torn.  v.  ;  W.  Turner, 
Jour.  Anatomy,  i.  313  (1867).  For  the  dentition  of  Orycteropus  see  0.  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  VIII 

THE    ORDERS    SIREXIA    AND    CETACEA 

Order  SIRENIA. 

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  very 
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, 


SIRENIA  21.3 

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  Manatus,  and  they  were  fully  developed  in 
Halitherium  and  other  fossil  forms.  The  anterior  caudal  vertebrae 
have  well-developed  chevron  bones.  In  one  genus  (Manatus)  there 
are  only  six  cervical  vertebrae.  There  are  no  clavicles.  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  (Rhytina)  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,  caecal  sacs  or  diverticula.  The  intestinal  canal  is  long,  and 
has  very  muscular  walls.  There  is  a  caecum,  either  simple,  conical, 
and  with  extremely  thick  walls,  as  in  Halicore,  or  bifid,  as  in  Maimtus. 
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, 


214  SIRENIA 

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  kidneys  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  mammae  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  algae  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  species  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  stelleri,  of  the 
North  Pacific,  was  totally  exterminated  through  the  agency  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  Eed  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. 


MANATID&  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  MANATID^E. 

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.1 — Incisors  -f ,  rudimentary,  concealed  beneath  the  horny 
oral  plates,  and  disappearing  before  maturity.  Molars  \\,  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 vertebrae  present  the  remarkable  anomaly  of  being  reduced  to 
six  in  number,  the  usual  vertebral  formula  being  C  6,  D  1 7,  L  2, 
and  C  23-25.  Rostrum  of  the  skull,  formed  by  the  union  of  the 
premaxillae  in  front  of  the  anterior  narial  aperture,  shorter  than  the 
length  of  the  aperture  and  scarcely  deflected  from  the  basi-cranial 
axis  ;  premaxillae  and  mandibular  symphysis  not  markedly  deflected 
(Fig.  72).  Tail  entire,  rounded,  or  shovel- shaped.  Rudimentary 
nails  on  the  fore  limbs.  Caecum  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  Manati  was  apparently  first  applied  to  this  animal  by 
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  Linnaeus  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  Avill  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  Mcth.  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 


FIG.  71. — American  Manatee  (Manatus  americanus),  from  life.    Proc,  Zool.  Soc.  1881,  p.  457. 

valve-like  slits,  at  the  apex  of  the  muzzle.  The  eyes  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 


MAN  ATI D^;  217 


backwards  considerably  behind  the  orbits.  Their  sides  are  formed 
by  the  ascending  processes  of  the  premaxillse  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  premaxillse,  supported  below  and  at  the  sides  by  the  maxillae. 
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 


FIG.  72. — Skull  of  African  Manatee  (Manatus  senegalensis).    \  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  SIRENIA 

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,  .2  ^  0  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.  As  a  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,  A  and  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 


MANATID&  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 

B 


FIG.  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  Murie, 
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 l  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). 


220  SIRENIA 

bones.  Of  these  the  American  Manatee  may  be  known  as  M. 
americanus,  although  it  has  been  described  under  the  names  of 
M.  latirostris,  and  M.  australis.  The  African  Manatee  (M.  senegalensis) 
differs  from  the  American  species  in  the  following  cranial  characters  : 
the  anterior  part  of  the  rostrum  is  shorter,  shallower,  and  altogether 
smaller ;  the  orbit  is  smaller ;  the  zygomatic  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,  edge ;  the  upper  surface 
of  the  frontal  is  flat,  instead  of  concave ;  the  foramen  magnum  and 
occipital  condyles  are  narrower  from  side  to  side,  and  the  symphysis 
of  the  mandible  is  smaller  and  shallower. 

Finally,  M.  inunguis  is  a  fluviatile  species  confined  to  the 
Amazon  and  Orinoco,  which  has  been  but  recently  fully  brought 
under  the  notice  of  zoologists. 


Family  HALICORID^E. 

Halicore.1 — In  the  upper  jaw  a  pair  of  large,  nearly  straight,  tusk- 
like  incisors,  directed .  downwards  and  forwards,  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  age  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  f ,  sometimes  f,  altogether,  but  not  all  in  place  at  once, 
as  the  first  falls  before  the  -last  rises  above  the  gum ;  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  no  enamel.  The  summits  of  the  crowns  are  tuberculated  before 
wearing,  afterwards  flattened  or  slightly  concave.  Skull  with  rostrum 
formed  by  the  union  of  the  premaxillae  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  the  lower  jaw  bent 
down  in  a  corresponding  manner.  Vertebrae  :  C  7,  D  18-19,  L  and 
C  30.  Tail  broadly  notched  in  the  middle  line,  and  with  two 
pointed  lateral  lobes.  No  nails  on  the  fore  limbs.  Caecum  single. 

1  Illiger,  Prodromus  Syst.  Mamm.  et  Avium,  p.  140  (1811). 


RHYTINID&  221 

The  Dugongs  are  more  distinctly  marine  in  their  habits  than  the 
Manatees,  feeding  chiefly  on  sea -water  algae.  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 : — H.  tabernaculi 
from  the  Red  Sea,  H.  dugong  from  the  Indian  seas,  and  H.  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  RHYTINID^E. 

Ehytina.1 — No  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.  Vertebrae  :  C  7,  D  19,  L  and  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 
csecal  appendages  to  the  pyloric  cavity.  Caecum  simple. 

Only  one  species  of  this  genus  is  known,  E.  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 
ill-fated  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,2  and  its 

1  Illiger.  Prodromits  Syst.  Mamm.  et  Avium,  p.  141  (1811). — Amended  from 
Rytina. 

2  Nordenskibld,  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. 


222  SI  REN  I  A 

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  laminarise  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  SIRENIANS. 

Halitherium.1 — The  Miocene  and  early  Pliocene  seas  of  Europe 
abounded  in  Sirenians,  to  which  the  generic  name  of  Halitherium 
Avas  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  -f-  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 
— Halitheriidce — were  thus,  in  dental,  cranial,  and  other  osteological 
characters,  less  specialised  than  are  either  of  the  existing  species, 

1  Kaup,  Ncues  Jahrbuch,  1838,  pp.  319  and  536. 


HALITHERIID&  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. Halitherium  remains  have  been  found  in  many 
parts  of  Germany,  especially  near  Darmstadt,  also  in  France,  Italy 
Belgium,  Malta,  etc. 
Until  a  few  years  ago 
none  were  known  from 
England,  probably  owing 
to  the  absence  of  beds 
of  an  age  corresponding 
to  those  in  Avhich  they 
are  found  on  the  Eu- 
ropean continent  ;  but  Fio.  74.— The  penultimate  and  last  right  lower  molars 
a  skull  and  several  °^  ^a^t^er^um  fossile ;  from  the  Miocene  of  the  Continent. 
,  .  (After  De  Blainville.) 

teeth  have  been  detected 

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  •§-  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  HalitJierium. 
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  fwms. — 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 
Halicoridce.  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  Rytiodus  of  E.  Lartet.  Of  this 


224  SIRENIA 

genus,  which  may  be  identical  with  Trachytherium  of  the  French 
Miocene,  better  preserved  remains  have  subsequently  been  described 
by  Delfortrie.  These  show  that  the  rostrum  is  more  elongated 
than  in  Halitherium,  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  f,  c  ^, 

/ 1  8\ 

p  +  m  (  jg  j  =  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  E.  Owen  under  the  name  of  Prorastomus  sirenoides.  Some 
writers  regard  this  genus  as  the  type  of  a  distinct  family — the 
Prorastomatidce.  Unfortunately  we  have  no  knowledge  of  the  geo- 
logical antiquity  of  the  formation  in  which  it  was  embedded.  Lastly 
must  be  mentioned  the  Eotherium  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  Manatus,  and  resembles 
that  of  Halitherium.  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  Manatus  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,  Symbolce  Sirenologicae,  St.  Petersburg, 
3  fasciculi,  1846-61-68— an  exhaustive  account  of  the  anatomy,  affinities,  and 
literature  of  the  group,  with  copious  illustrations  of  the  osteology  of  Rhytina. 


CETACEA  22$ 

Anatomy  of  Duyong : — 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).  Manatee: — W.  Vrolik,  Bijdragen  tot  dc  Dicrkundc,  1851; 
J.  Murie,  "On  the  Form  and  Structure  of  the  Manatee,"  Trans.  Zool.  Soc.  Lond. 
vol.  viii.  p.  127,  1872,  and  "Further  Observations  on  the  Manatee,"  Ibid.  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,  torn.  i.  p.  332,  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  sonie  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  vertebra?,  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  vertebrae  are  united  together  to  form  a  sacrum.  The  lumbar 
and  caudal  vertebras  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.1 
There  are  largely  developed  chevron  bones,  the  presence  of  which 
indicates  the  distinction  between  the  caudal  and  lumbar  vertebrae. 

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  maxilla?, 
premaxillae,  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  laminae  or  "baleen." 
The  salivary  glands  are  rudimentary  or  absent.  The  stomach  is 
multilocular,  its  structure  being  fully  noticed  under  the  genus 


Fr 


Pa 

JSO 


FIG.  "5.  —  A  section  of  the  skull  of  a  young  Dolphin  (Globicephalus  melas).  xj.  PMx,  Pre- 
maxilla  ;  MX,  maxilla  ;  ME,  ossified  portion  of  the  mesethmoid  ;  an,  anterior  nares  ;  Na, 
nasal  ;  IP,  inter-parietal  ;  Fr,  frontal  ;  Pa,  parietal  ;  50,  supraoccipital  ;  ExO,  exoccipital  . 
HO,  basioccipital  ;  Sq,  squamosal  ;  Per,  periotic  ;  AS,  alisphenoid  ;  PS,  presphenoid  ;  Pt, 
pterygoid  ;  pn,  posterior  nares  ;  PI,  palatine  ;  Vo,  vomer  ;  s,  symphysis  of  mandible  ;  id, 
inferior  dental  canal  ;  cp,  coronoid  process  of  mandible  ;  cd,  condyle  ;  a,  angle  ;  sh,  stylo-hyal  ; 
T)h,  basi-hyal  ;  th,  thyro-hyal.  (From  Flower's  Osteology  of  Mammalia.) 


The  intestinal  canal  is  simple,  and  only  in  some  species 
provided  with  a  small  caecum.  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  mirabilia.  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  vesiculae  seminales,  nor  os 
penis.  The  uterus  is  bicornuate,  and  the  placenta  nondeciduate 
and  diffuse.  The  mammse  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  passages 
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  (Orca)  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  medusae,  while  the  principal  staple  of  the  food  of 
many  is  constituted  by  the  various  species  of  cephalopods,  Loligo 
and  other  Teuthida,  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  at  a  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  outAvard  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  Zeuglodon,  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  csecum  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,1  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  ami  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,1 
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  Squalodon,  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  foatal 
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  vertebrae,  on  which  the  genus  Palceocetus  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  Oturiidw  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,1  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  "  2  to  the  Porpoise 
contains  a  larger  element  of  truth  than  the  speculations  of  many 
accomplished  zoologists  of  modern  times.  The  fact  that  Platanista, 
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  Mar  sou  in.     "Porpoise"  is  said 
to  be  derived  from  "  Parc-j)oisson." 


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  BAL^NOIDEA,  Whalebone,  or  True  WJides.1 

Family  BAI^ENID^. 

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  vertebrae ;  their  capitular 
processes,  when  present,  not  articulating  directly  with  the  bodies  of 
the  vertebrae.  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  caecum. 

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.2  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 

1  Icel.   hvalr ;    Dan.    and   Swed.    hval ;   Anglo-Saxon  hwcel ;    Germ,    wal, 
walfisch.    The  meaning  apparently  is  "roller,"  the  word  being  closely  allied  to 
"wheel"  (Skeat). 

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


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.  Each  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  maxillae,  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  papillae,  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  papillae,  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  papillae  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.1 

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  (Balcena)  and  Rorquals  or  Fin-Whales 
(Balcenoptera),  the  latter  distinguished  by  their  smaller  heads, 
elongated  and  slender  form,  free  cervical  vertebrae,  tetradactylous 
maims,  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. 

Balcena.2 — Skin  of  throat  smooth,  not  furrowed.  No  dorsal  fin. 
Cervical  vertebras  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. 


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

The  Greenland,  or  more  properly  Arctic,  Right  Whale  (Balcena 
mysticetus)  attains,  when  full  grown,  a  length  of  from  45  to  50 
feet.  Its  usual  vertebral  formula  is  C  7,  D  12,  L  14,  C  22. 
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. 

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


BAL^ENID^E  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  circum- 
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 
Eschricht  and  Reinhardt  in  the  case  of  the  Greenland  seas,  "  every- 
thing tends  to  prove,"  Scammon  says,  "  that  the  Balcena  mysticetus 
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  a  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. 


Fio.  77. — Southern  Right  Whale  (Balcena  australis). 


The  southern  Right  Whale  (B.  australis,  Fig.  77)  resembles  the 
last  in  the  absence  of  dorsal  fin  and  of  longitudinal  furrows  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 


BAL^ENID^K  239 


(fifteen)  of  ribs  and  dorsal  vertebrae.  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  Atlantic,  B.  japonica  of 
the  North  Pacific,  B.  australis  of  the  South  Atlantic,  and  B.  anti- 
podarum  and  B.  novce-zealandice  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  Imnted  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. 


FIG.  78. — The  right  tympanic  bone  of  an  immature  individual  of  the  Greenland  Whale 
(Balcsna  mysticetus),  from  the  inner  (A)  and  outer  (£)  aspects.  J  natural  size.  (From  the 
Proc.  Zool.  Soc.) 

Remains  of  Right  Whales  are  of  not  uncommon  occurrence  in  the 
Pliocene  Crag  deposits  of  England  and  Belgium.  The  tympanics 
of  B.  affinis  from  these  deposits  appear  to  indicate  a  species  closely 
allied  to  B.  mysticetus,  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  Balcenula  balcenopsis,  the  generic  distinction  being 
made  on  account  of  the  free  condition  of  the  atlas  and  seventh 
cervical  vertebrae ;  but  it  seems  scarcely  advisable  to  regard  such  a 
feature  as  indicating  more  than  a  less  specialised  species.  Balcena 
(Balcenotus)  insignis  is  a  whale  of  somewhat  larger  dimensions,  in 
which  the  atlas  is  generally,  and  the  seventh  cervical  vertebra 


BAL^ENID^E  241 

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

Neobalcena.1 — Head  about  one-fourth  the  total  length.  Skin  of 
the  throat  not  plicated.  A  small  falcate  dorsal  fin.  Vertebrae, 
C  7,  D  17,  L  3,  C  16  =  43  The  cervical  vertebras  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  Balcena,  but  with  certain 
very  characteristic  peculiarities  of  shape.  Baleen  very  long,  slender, 
elastic,  and  white.  A  single  species,  at  present  very  rare,  N.  mar- 
ginata,  from  the  Australian  and  New  Zealand  seas  is  the  smallest 
of  the  Whalebone  Whales,  being  not  more  than  20  feet  in  length. 

Ehachianedes.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  Balcena.  The  baleen  is  the 
shortest  and  coarsest  of  any  of  the  group.  Its  osteology  is  im- 
perfectly known.  One  species,  R.  glaucus,  the  Gray  Whale  of  the 
North  Pacific. 

Megaptera.3 — Head  of  moderate  size.  Baleen  plates  short  and 
broad.  Vertebrae,  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  Balcenoptera,  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  loops)  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.  Sci.  PMlad.  1869,  p.  15. 

3  Gray,  Zoology  of  Erebus  and  Terror,  p.  16  (1846). 

16 


242 


CETACEA 


the  winter  of  1883-84.1     The  usual  length  of  the  adult  ranges  from 
45  to  50  feet,  the  female  being  larger  than  the  male.     Whales  of 


Fio.  79. — Humpbacked  Whale  (Megaptera  boops). 

the  genus  Megaptera  are  found  in  the  South  Atlantic  and  in 
both  the  North  and  the  South  Pacific.  They 
resemble  those  of  British  seas  so  closely  that  it 
is  doubtful  whether  the  differences  which  have 
been  observed,  and  upon  which  several  species 
have  been  founded,  may  not  be  individual  peculi- 
arities; but  zoologists  have  not  yet  had  the 
opportunity  of  examining  and  comparing  such 
a  series  of  specimens  of  different  ages  and  sexes 
from  different  localities  as  would  be  necessary 
to  determine  these  points  satisfactorily. 

Tympanic  bones  of  Megaptera  occur  in  the 
English  and  Belgian  Crags,  although  somewhat 
less  commonly  than  those  of  Balcena  and  Balcen- 

'  optera;  they  have  been  described  under  the 
names  of  Megapteropsis  and  Burtinopsis. 

Balcenoptera.2 — Head  small  and  flat,  and 
pointed  in  front.  Body  long  and  slender.  Skin 
of  throat  plicated.  A  small  falcate  dorsal  fin. 
Baleen  short  and  coarse.  Cervical  vertebrae  free. 
Scapula  low  and  broad,  with  a  large  acromion 

i    and  coracoid  process.     Pectoral  limb  tetradacty- 

|  lous,  small,  narrow,  and  pointed.  Tympanic 
(Fig.  81)  long,  much  inflated,  and  rounded,  with 
the  involuted  portion  thickened  and  pyriform, 

;  and  the  notch  for  the  Eustachian  canal  sharply 
defined ;  inner  surface  flattened,  without  the 
vertical  groove  found  in  Megaptera. 

The  Korquals,  Fin- Whales,  Fin-backs,  Fin- 
ners,  or  Eazor-backs,  as  they  are  variously  called, 

1  See  J.   Struthers,   ' '  On  the  Anatomy  of  Megaptera 
longimana,"  Journ.  Anatomy  and  Physiology,  1887-89. 

2  Lacepede,  "Table  ties  Ordres,"  Hist.  Nat.  dcs  Cttacts, 
p.  xxxvi.  (1804). 


243 


have  the  plicated  skin  of  the  throat  like  that  of  Megaptera,  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- 
turing them  by  the 
old  methods,  these 
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)  Balcenoptera  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 
vertebrse,  of  which  1 6  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  (Euphausia  inermis)  which 


FIG.  81. — The  right  tympanic  of  Balcenoptera  musculus  from 
the  inner  (A)  and  outer  (B)  aspects.  J  natural  size.  (From  the 
Proc.  Zool.  Soc.) 


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)  Balanoptera  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  vertebras, 
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) 
Balcenoptera  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  vertebras,  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)  Balcenoptera  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  vertebrae,  1 1  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 


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.1 

Tympanics,  vertebrae,  and  other  bones  of  Korquals  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  Balcenoptera,  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  Balsenopterine  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 
Balcenoptera.  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 
vertebras  is  also  greater.  In  the  typical  forms  (e.g.  C.  brialmonti 
and  C.  dubium)  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 
Megaptera,  but  has  the  greater  part  of  the  cavity  filled  by  bone. 
There  is  a  talon  to  the  condyle  of  the  mandible. 

Palceocetus,  as  already  mentioned  (p.  232),  is  founded  upon  the 
ankylosed  cervical  vertebrae  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  Balcenidce  it  indi- 
cates a  Right  Whale. 

1  See  P.  J.  Van  Beneden,  "  Histoire  Naturelles  des  Balenopteres, "  Mem.  Acad. 
Bclyiquc,  xli.  1887. 


246  CETACEA 

Suborder  ARCILEOCETT. 
Family  ZEUGLODONTID^E. 

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  i  f ,  c  \,  p  and  m  £  =  36,  resembling  that 
of  some  Seals.2  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  fossae  very  large.  A 
strong  sagittal  crest.  Rostrum  long  and  narrow,  differing  from 
that  of  other  Cetaceans  in  the  large  extent  to  which  the  premaxillae 
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 
vertebrae  free.  The  characters  of  the  dorsal  vertebrae  and  mode  of 
articulation  of  the  ribs  appear  to  have  resembled  those  of  Platanista 
rather  than  JBalcena,  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.3 

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 
Garni  vora.     ' '  On  the  Systematic  position  of  Zeuglodon, "  Studies  from  tJie  Museum 
of  Zoology,  Dundee,  vol.  i.  No.  9,  1890. 

2  An  appearance  in  one  specimen  has  been  described  by  C.  G.  Cams  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. 

3  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. 


PHYSETERID&  247 

yoked  together."  This  name  was  substituted  by  Owen  for  the 
earlier  one  Basilosawns  of  Harlan,  with  the  consent  of  that  author, 
on  the  mammalian  nature  of  the  animal  being  demonstrated.1  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. 

Subwder  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.  Eami  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  Avell-developed  capitular  processes,  articulating 
with  the  bodies  of  the  vertebrae.  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  PHYSETERID^E. 

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  GET  ACE  A 

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  vertebrae,  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  vertebrae.  (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  Physeterinse. — 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.1 — Upper  teeth  apparently  of  uncertain  number,  rudi- 
mentary, and  functionless,  being  embedded  in  the  gum.  Lower  jaw 


FIG.  82.— 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  edges 
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 

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


PHYSETERID.-E 


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.  Vertebrae : 
C  7,  D  11,  L  8,  C  24;  total  50.  Atlas  free;  all  the  other  cervical 
vertebrae  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 


FIG.  83.— The  Sperm  Whale  (Physeter  macrocephaius'). 


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  GET  ACE  A 

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  1 2  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.  Vertebrae :  C  7,  D  13  or  14,  L  and  C  30  ;  total  50  or  51. 
All  the  cervical  vertebrae  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  breiiceps. 
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  potsii),  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.  breviceps.  This 
animal  appears  to  attain  the  length  of  10  feet  when  adult,  and  has 
1  Gray,  Zoology  of  Erebus  and  Terror,  p.  22  (1846).  Usually  spelt  Kogia. 


PHYSETERID&  251 

been  met  with  at  various  distant  localities  in  the  Southern  Ocean, 
and  also  oft'  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  Ziphiince. — 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  Delphinidce.  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. 

Hyperoodon.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  premaxillse  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  premaxillse, 
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 
maxillae  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. 
Vertebras:  C  7,  D  9,  L  10,  C  19 ;  total  45.  All  the  cervical 
vertebras  united.  Upper  surface  of  the  head  in  front  of  the  blow- 


FIG.  S4.—Hyperoodon  rostmtiis.    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.2  So  different,  indeed,  is  the  appearance  of 
the  skull  of  an  old  male  from  that  of  a  female  individual  that 

1  Lacepecle,  "Table  des  Ordres,"  Hist.  Nat.  des  Cttacts,  p.  xliv.  (1804). 
2  See  the  figures  in  the  Proc.  ZooL  Soc.  1882,  pp.  728,  729. 


PHYSETERID&  253 

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

The  Hyperoodon,  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  Society'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 
Da  vis'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 


254  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  yields  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  Hyperoodon  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.1 — 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  premaxillae  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  premaxillae.  Anteorbital  notch  not  distinct.  Rostrum  (seen 
from  above)  triangular,  gradually  tapering  from  the  base  to  the 
apex ;  upper  and  outer  edges  of  maxillae '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.  Vertebrae :  C  7,  D  10,  L  10,  C  22  ;  total  49.  The 
three  anterior  cervical  vertebrae  united,  the  rest  free. 

The  type  of  this  genus  is  Z.  cavirostris  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  Epiodon 
being  probably  referable  to  the  type  species. 

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

Mesoplodon.2 — A  much  compressed  and  pointed  tooth  in  each 

1  Cuvier,  Ossemens  Fossiles,  2d  ed.  vol.  v.  p.  352  (1823). 

2  Gervais,  Ann.  Sci.  Nat.  ser.  3,  vol.  xiv.   p.  16  (1850).     For  the  very  com- 
plicated synonymy  of  this  genus,  see  Traits.  Zool.  Soc.  vol.  viii.  p.  208. 


PHYSETERID^E 


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. 


FIG.  85. — Mesoplodon  bidens.    From  Reinhardt. 

Skull  with  the  region  around  the  nares  as  in  Hyperoodon,  except 
that  the  nasals  are  narrow  and  more  sunk  between  the  upper  ends 
of  the  premaxillse ;  like  those  of  Hyperoodon,  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 


FIG.  86. — 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.  Vertebrae : 
C  7,  D  10,  L  10  or  11,  C  19  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  in  the  case  of  M.  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  (M.  grayi] 
a  row  of  minute,  conical,  pointed  teeth,  like  those  of  ordinary 
Dolphins,  17  to  19  in  number,  are  present  even  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  M.  bidens,  M.  europceus,  M.  densirostris,  M.  layardi, 
M.  grayi,  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 


. 

comer  of  the  figure  forms  the  anterior  ar-  folk.  Several  Species  have  been 
ticulation  with  the  tympanic.  (From  the  foun(Jed  UDOn  the  evidence  of 
Cat.  Foss.  Mamm.  Brit.  Miis.  pt.  v.  p.  70.)  ,,  -•->  .  ,.  •,  ,. 

these  rostra,      renotic  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. 

Bemrdius.1  —  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  premaxillae  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 
maxillae  at  base  of  rostrum.  Vertebrate  :  C  7,  D  10,  L  12,  C  19  ; 
total  48.  The  three  anterior  cervicals  ankylosed,  the  rest  free  and 
well  developed. 

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

1  Duvernoy,  Ann.  Sci.  Nat.-Zoologie,  ser.  3,  vol.  xv.  p.  41  (1851). 


SQUALODONTID^E  257 

Choneziphius.1 — The  rostral  portions  of  crania  from  the  Antwerp 
and  Suffolk  Crags,  on  the  evidence  of  which  this  genus  has  been 
established,  agree  with  those  of  Mesoplodon  in  having  the  premaxillse 
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  premaxill*  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. 


Family  SQUALODONTID^. 

Numerous  extinct  forms,  chiefly  known  by  teeth  and  fragments 
of  crania,  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  Archseoceti,  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. 

Squalodon.2 — All  the  forms  may  be  included  in  this  genus,  the 
so-called  Rhizoprion  not  being  distinct.  Dentition  :  i  f ,  c  -J-,  simple 
teeth  of  the  molar  series  (premolars  ?)  -£,  two-rooted  molars  -f  =  ^f ; 
total  60.  The  double-rooted  molars  differ  from  those  of  Zeuglodon 
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  contemporaneous  with  the  beds 
in  which  they  occur,  indicate  the  survival  of  the  genus  into  the 
Pleistocene  period. 

Family  PLATANISTID^E. 

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

1  Duvernoy,  op.  cit.  p.  61. 

2  Grateloup,  Act,  Ac.  E.  Sci,  Bordeaux,  1840,  p.  208. 
17 


258  CETACEA 

points  in  common,  and  do  not  altogether  come  under  the  definition 
either  of  the  Physeteridce  or  the  DelpMxidce,  especially  in  the 
important  -character  of  the  mode  of  articulation  of  the  ribs  with 
the  dorsal  vertebrae,  the  tubercular  and  capitular  articulations, 
distinct  at  the  commencement  of  the  series,  gradually  blending 
together,  as  they  do  in  most  ordinary  mammals.  The  cervical 
vertebras  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.1 — Teeth  about  %%  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.  Maxillse  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. 
Vertebras:  C  7,  D  10,  L  9,  C  26  ;  total  52.  A  small  cascum.  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 


FIG.  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.  AmpUb.  etc.,  p.  35  (1830). 

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


PLA  TANISTIDsE 


259 


iii..1 — 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.  Vertebrae  :  C  7,  D  13,  L  3,  C"18  ;  total 
41.  Transverse  processes  of  lumbar  vertebrae  very  broad.  Sternum 
short  and  broad,  and  consisting  of  a  single  segment  only.  Dorsal 
fin  a  mere  ridge.  The  long  cylindrical  rostrum  externally  furnished 
with  scattered,  stout,  and  crisp  hairs.  One  species  only  is  known, 
/.  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. 
Vertebrae:  C  7,  D  10,  L  5,  C  19 ;  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 


Fio.  89. — Pontoporia  blainvillei.    (From  Bunneister.) 

transverse,  crescentic.  This  genus  connects  the  last  two  forms  with 
the  true  Delphinidce.  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  Pontoporui,  but  they 
have  been  regarded  by  other  writers  as  indicating  a  distinct  genus, 
for  which  the  names  Palceopontoporia  and  Poiitistes  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  vertebras,  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  Stem,  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, 
Ixacanthus,  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). 


26o  CETACEA 

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


Family  DELPHINID^E. 

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  vertebrae  gradually  trans- 
ferred from  the  arches  to  the  bodies  of  the  vertebrae  without  any 
sudden  break,  and  becoming  posteriorly  continuous  serially  with  the 
transverse  processes  of  the  lumbar  vertebrae.  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.1 — 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). 


DELPHINID^E 


261 


small,  not  meeting  in  the  middle  line,  but  approxi- 
mating posteriorly.  Vertebrae:  C  7,  D  11,  L  6, 
C  26 ;  total  50.  Cervical  region  comparatively 
long,  and  all  the  vertebrae  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  (M.  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  circumpolar   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  Wernerian  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 


262  CRT  ACE  A 

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.1 — 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  -f-  to  ^$, 
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. 
Premaxillae  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.  Vertebra  :C7,  D  11,  L  9,  C  23 ;  total  50. 
Cervical  vertebrae  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. 


FIG.  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.  leucas  (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  Lacepede,  Hist.  Nat.  dcs  Cetarts,  p.  xli.  (1804). 


DELPHINID&  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. 

Eemains  of  a  Cetacean  from  the  Lower  Pliocene  of  Tuscany  have 
been  referred  by  Brandt  to  this  genus  under  the  name  D.  brocchii. 

In  all  the  remaining  genera  of  Delphinidce  the  cervical  region  of 
the  vertebral  column  is  very  short,  and  the  first  two,  and  usually 
more,  of  the  vertebrae  are  firmly  united. 

Phoccena.1 — Teeth  f f,  small,  occupying  nearly  the  whole  length 
of  the  rostrum,  with  compressed,  spade -shaped  crowns,  separated 
from  the  root  by  a  constricted 
neck  (Fig.  92).  Kostrum  rather 
shorter  than  the  cranium 
proper,  broad  at  the  base  and 
tapering  towards  the  apex. 
Premaxillae  raised  into  tuber- 
osities  in  front  of  the  nares. 
The  frontal  bones  forming  a  , 

FIG.  92.— Teeth  of  Porpoise.    Twice  natural  size. 

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.  Vertebrae  :  C  7,  D  13,  L  14,  C  31 ;  total  65  (subject 
to  slight  individual  variations).  First  to  sixth  cervical  vertebrae, 
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,  marsouin.  "  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  Phocana  of  Cuvier,  of  which  the 
Porpoise  of  the  British  seas,  Phoccena  communis,  Cuvier  (Delphimts 
phoccena,  Linnaeus),  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). 


264 


CETACEA 


from  nose  to  median  notch  between  the  flukes  of  the  tail,  62i 
inches ;  from  the  nose  to  the  anterior  edge  of  the  dorsal  fin,  29 
inches  ;  height  of  dorsal  fin,  4|  inches  ;  length  of  base  of  dorsal  fin, 
8  inches ;  length  of  pectoral  fin,  9^  inches  ;  breadth  of  pectoral  fin, 
3|  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 


FIG.  93. — The  Common  Porpoise  (Phocfena  communis). 

moderate  size,  and  slightly  falcate.  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 


DELPHINID& 


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 
Phoccena  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  (b) 
may  be  regarded 
as  a  kind  of  crop, 
or  dilatation  of 
the  large  oeso- 
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 
Ungulates  and 
certain  Rodents ; 
but,  although  its 
walls  do  not  appear  to  contain  peptic  glands,  its  contents  undergo 
partial  digestion — probably  caused  by  the  regurgitation  into  it 


FIG.  94. — Diagrammatic  section  of  the  stomach  of  the  Porpoise, 
a,  (Esophagus  ;  6,  left,  or  cardiac,  compartment ;  c,  middle  compart- 
ment ;  d  and  e,  the  two  divisions  of  the  right,  or  pyloric,  compart- 
ment ;  /,  pylorus  ;  g,  duodenum,  dilated  at  its  commencement ;  h, 
biliary  duct. 


266  CRT  ACE  A 

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  /)  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  (/).  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,  GloU- 
cepJialus,  and  Grampus)  there  are  two  such  small  sacs  of  very  similar 
size  and  form,  communicating  by  circular  pylorus -like  apertures ; 
and  in  Hyperoodon  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  pJwccenoides  by  Cuvier, 
D.  melas  by  Schlegel,  forms  the  type  of  Gray's  genus  Neomeris.1 
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  J.apan. 
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),  f  4  to  f£.  Vertebrae :  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). 


DELPHINID&  267 


small,  narrow,  and  ovate.  Typified  by  C.  hearisulei,  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.1 — Teeth  -J-f  to  ^,  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 — 0.  hrevirostris,  from  the  Bay  of 
Bengal,  and  0.  fluminalis,  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.2 

Orca? — Teeth  about  ^-|,  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  arid  posterior  surfaces.  Rostrum  about  equal  in 
length  to  the  cranial  part  of  the  skull,  broad  and  flattened  above, 
rounded  in  front ;  premaxillae  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.  Vertebrae:  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  vertebras  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, 
falcate  dorsal  fin,  whence  their  common  German  name  of  Scliwert- 
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  JVTiales  in  Brit.  Mus.  p.  285  (1866). 

2  Anatomical  and  Zoological  Researches,  comjmsing  an  Account  of  the  Zoological 
Results  of  the  two  Expeditions  to  Western  Yunnan,  in  1868  and  1875  (1878). 

3  Gray,  Zoology  of  Erebus  and  Terror,  p.  33  (1846). 


268  CRT  ACE  A 

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. 


FIG,  95. — The  Killer  Whale,  or  Grampus  (Orca  gladiator).    From  Hunter. 

Orca  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. 

Psevdorca.1 — Teeth  about  ^$.  Cranial  and  dental  characters 
generally  like  those  of  Orca,  except  that  the  roots  of  the  teeth  are 
cylindrical.  Vertebrae:  C  7,  D  10,  L  9,  C  24;  total  50.  First 
to  sixth  or  seventh  cervical  vertebrae  united.  Bodies  of  the  lumbar 
vertebrae  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,  falcate.  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 
Phoccena  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. 

Q TO 

Globicephalus? — Teeth  8_12>  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. 

1  Reinhardt,  Overs.  Dan.  Sezsk.  Fork.  1862,  p.  151. 
2  Lesson,  N.  Tab.  d.  Etgne  Animal— Mamm.  p.  200  (1842). 


DELPHINID&  269 

Premaxillae  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  maxillae  in  the  anterior  half  of  this 
region.  Pterygoids  of  normal  form,  meeting,  or  very  nearly  so, 
in  the  middle  line.  Vertebra:  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 
vertebrae  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,  Ca'ing  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 
themselves  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  Flo.  W.-TI*  left  periotic  bone 
indistinguishable,  either  by  external  or  of  Giobicepiiaius  unddens;  from  the 
osteological  characters,  from  those  of  the  Suffolk  Cn*  Xatu[al  ?«• .  The 

AT        V.    A    1  grooved  surface  on  the  right  is  the 

JNortn  Atlantic.  anterior  facet  for  articulation  with 

Teeth,  periotic  (Fig.   96)  and   tym-    the  tympanic ;  the  posterior  tym- 
panic    bones     from     the     Suffolk    Crag,     Panic  articulation  being  on  the  op- 
_  .  7  .     T  *  posite  side  of  the  figure.    (Prom  the 

described  as  G.  unaaens,  indicate  a  form  Cat  Foss-  Mamm.  Brit.  M™.  pt  v.) 
apparently  closely  allied  to  the  existing 

species.  The  periotic  is  figured  in  order  to  illustrate  the  dis- 
tinctive characters  of  that  bone  in  the  Delphinida1. 


270  GET  ACE  A 

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. 
Vertebra:  C  7,  D12,  L19,  C  30 ;  total  68.  General  external 
characters  much  as  in  GlobicepJialus,  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  GlobicepJialus,  Grampus,  and  Lagenarhynclms.  From  the 
latter  they  differ  chiefly  in  the  smaller  number  (about  -J-g-)  and  much 
larger  size  (6-7  mm.  in  diameter  at  base  of  crown)  of  the  teeth. 

Lagenorhynchus.3 — 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),  f  f  to  f  f .  Vertebrae 
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 
vertebrae  free. 

If  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). 

2  Gray,  Proc.  Zool.  Soc.  1870,  p.  77. 
3  Gray,  Zoology  of  Erebus  and  Terror,  p.  35  (1846). 


DELPHINID^E  271 

Delphinus.1 — Teeth  very  numerous  in  both  jaws,  £$•  to  £$, 
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  middle  line  throughout  their  length. 
Palate  with  deep  lateral  grooves.  Vertebrae  73  to  75.  Pectoral  fin 
of  moderate  size,  narrow,  pointed,  somewhat  falcate.  Second  and 
third  digits  well  developed ;  the  rest  rudimental. 

The  type  of  the  genus  is  the  Common  Dolphin  of  the  Mediter- 
ranean (D.  delphis,  Fig.  97),  also  found  in  the  Atlantic,  and  of 


FIG.  97. — The  Common  Dolphin  (Delphinus  cUlphis).     From  Reinhardt. 

which  a  closely  allied  if  not  identical  form  is  met  with  in  the 
Australian  seas  (D.  forsteri)  arid  in  the  North  Pacific  (D.  bairdi). 
Other  species  are  D.  janira,  D.  major,  etc. 

Tursiops.z — Rostrum  tapering  moderately  from  base  to  apex ; 
palate  not  grooved ;  symphysis  of  mandible  short ;  other  cranial 
characters  as  in  Delphinus.  Teeth  f-^  to  f  •§-,  stout  (6  to  7  mm.  in 
antero-posterior  diameter).  Vertebrae :  C  7,  D  13,  L  17,  C  27;  total 
64.  Limbs  as  in  Delphinus.  Represented  by  the  widely  distributed 
T.  tursio  ;  T.  catalania  being  a  second  form.  Fossil  remains  of  this 
genus  from  the  Italian  Pliocene  have  been  recently  described. 

Prodelphinus.3 — Rostrum  somewhat  variable ;  mandibular  sym- 
physis short  (less  than  one -fifth  the  length  of  the  ramus);  other 
cranial  characters  as  in  the  preceding  genus.  Teeth  f$  to  -£$, 
small,  not  exceeding  3  mm.  in  diameter.  Vertebrae  73  to  78. 
Limbs  as  in  Delphinus.  Four  leading  types  of  this  genus  are 
recognised  (all  of  which  have  numerous  synonyms)  viz.  P.  obscurus, 
P.  euphrosyne,  P.  doris,  and  P.  longirostris. 

Peron's  Dolphin  (Delphinus  leucorhamphus,  Pe"ron,  or  Leuco- 
rhamphus  peroni,  Lilljeborg)  resembles  some  forms  of  Prodelphinus  in 
its  cranial  characters ;  but  having  no  dorsal  fin,  it  has  been  separated 
generically  by  some  writers.  It  is  not  improbable  that  Delphinus 
borealis,  Peale,  from  the  North  Pacific,  in  which  there  is  likewise  no 
dorsal  fin,  may  be  an  allied  form. 

Steno.*- — Rostrum  long,  narrow,  and  compressed,  very  distinct 
from  the  cranium ;  mandibular  symphysis  as  long  as,  or  longer  than 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  108  (1766). 

2  Gervais,  Hist.  Nat.  des  Mammifercs,  vol.  ii.  p.  323  (1855). 

3  Gervais,  Osteographie  des  Cttacds,  p.  604  (1880). 

4  Gray,  Zoology  of  Erebus  and  Terror,  p.  43  (1846). 


272  .     GET  ACE  A 

one-fourth  the  length  of  the  ramus ;  other  cranial  characters  as  in 
the  preceding  genus.  Teeth  £ \  to  f  4,  of  comparatively  large  size 
(5-6  mm.  in  diameter) ;  surface  of  their  crowns  finely  grooved. 
Vertebrae:  07,  D 12,  L 15,  C  32  ;  total  66.  Kepresented  by 
S.  rostratus,  from  which  the  forms  which  have  received  other  names 
are  probably  not  specifically  separable. 

Sotalia.1 — 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  Steiio.  Teeth  tolerably  large  (4-5  mm.  in  diameter),  f  $  to  f f, 
with  smooth  enamelled  surface.  Vertebrae  :  C  7,  D  12,  L  10-14, 
C  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  (S.  sinensis)  and  S.  pallidus 
from  the  river  Amazon. 

Bibliography  of  Cetacea. — D.  F.  Eschricht,  Untersuchungen  ilber  die  Nordischcn 
Wallthiere,  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,  Osteographie  des 
Cetaces,  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  iiber  die  Fossilen  und  Subfossilen  Cetaceen  Europa's,"  in 
Mem.  de  I'Acad.  Imp.  de  St.  Petersbourg,  7ifeme  ser.  vol.  xx.  1873  ;  C.  M.  Scammon, 
Marine  Mammals  of  the  N.  W.  Coast  of  North  America,  1874  ;  W.  H.  Flower, 
"  On  the  characters  and  Divisions  of  the  Families  of  the  Delphinidae,"  Proc.  Zool. 
Soc.  1883,  p.  466,  and  List  of  the  Spccimeiis  of  Cetacea  in  the  British  Museum, 
1885  ;  F.  W.True,  "Review  of  the  Family  Delphinidaj, "  Bull.  U.S.  Nat.  Museum, 
No.  36,  1889  ;  P.  J.  Van  Beneden,  Histoire  Naturellc  des  Cetaces  des  Mers 
d' 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.  £.  Belgiqiie  and  Ann.  Mus.  E.  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). 

1  Gray,  Cat.  Seals  and  Whales  Brit.  Mus.  2d  ed.  p.  393  (1866). 


CHAPTEK    IX 

THE    ORDEK    UNGULATA 

UNDER  this  term  may  be  included  provisionally  a  large  and  rather 
heterogeneous  group  of  mammals,  the  existing  members  of  which 
form  the  Pecora  and  Belluse  of  Linnaeus,  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.1  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  Wincza  (Morpholoq.  Jahrb.  xvi. 
p.  647). 

18 


274 


UNGULATA 


The  more  generalised  of  the  fossil  forms  do  not  conform  in  all 
respects  to  the  above-mentioned  characters ;  clavicles  being  present 
in  Ti/potherium,  and  perhaps  in  some  of  the  Condylarthra,  while  in 
the  latter  group  the  humerus  may  have  an  entepicondylar  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^structure  is  exhibited  by  the  extinct 
Macrotherium,  provisionally  included  in  the  Perissodactyla,  where 
the  digits  terminated  in  long  and  curved  claws. 

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 
grinding  surface,  are  termed  brachydont  (Fig.  122) ;  while  those  with 
higher  crowns,  in  which  the  bases  of  the  infoldings  of  enamel  are 
invisible  from  the  grinding  surface,  are  known  as  hypsodont  (Fig.  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  bunodont ;  but  when 
they  are  expanded  in  an  antero- 
posterior  direction  and  curved  into 
a  crescent  shape  the  tooth  is 
described  as  selenodont. 

The  whole  order  may  be 
divided  into  the  Ungulata  Yera, 
containing  the  suborders  Perisso- 
dactyla and  Artiodactyla,  and  a 
somewhat  heterogeneous  assem- 
blage of  animals  which  may  be 
called  Subungulata  or  Ungulata 
Polydactyla.  Cope  has  pointed 
out  a  character  in  the  structure 
of  the  carpus  by  which  the  latter 
are  differentiated  from  the  former. 
Thus  in  all  the  Subungulata  the 
bones  of  the  proximal  and  distal 
row  retain  the  primitive  or  more 
typical  relation  to  each  other  (see 
Fig.  98) ;  the  os  magnum  of  the 
second  row  articulating  mainly 
with  the  lunar  of  the  first,  or 
with  the  cuneiform,  but  not  with  the  scaphoid.  But  in  the  group  to 


B 


FIG.  98. — Right  fore  foot  of  Indian  Ele- 
phant. X  J.  U,  ulna  ;  R,  radius  ;  c,  cunei- 
form ;  I,  lunar ;  sc,  scaphoid  ;  u,  unciform ; 
m,  magnum  ;  td,  trapezoid  ;  tm,  trapezium  ; 
I  to  V,  first  to  fifth  digit. 


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.1 

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  mammae  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  Biwidce  (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  nianus  of  the  extinct  Cotylops. 


276 


UNGULATA 


respects  a  great  contrast  to  such  decadent  types  as  those  represented 
by  the  Tapirs  and  Ehinoceroses. 

The  principal  anatomical  characters  by  which  the  Artiodactyles 
are  distinguished  from  the  Perissodactyles  are  as  folloAvs.  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. 


FIG.  99.— Bones  of  right  fore  foot  of  existing  Artiodactyles.  A,  Pig  (Sus  scrofa),  xj;  B, 
Red  Deer  (Cervus  elaphus),  x }  ;  C,  Camel  (Camelus  bactrianus),  x|.  U,  Ulna;  R,  radius ;  c, 
cuneiform  ;  I,  lunar ;  s,  scaphoid  ;  M,  unciform  ;  ra,  magnum  ;  td,  trapezoid  ;  tm,  trapezium. 
From  Flower's  Osteology  of  Mammalia. 

Dorsal  and  lumbar  vertebrae  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  277 

of  the  astragalus  divided  into  two  nearly  equal  facets,  one  for  the 
navicular  and  the  other  for  the  cuboid  bone.  The  calcaneum  with 
an  articular  facet  for  the  lower  end  of  the  fibula.  Stomach  almost 
always  more  or  less  complex.  Colon  convoluted.  Caecum  small. 
Placenta  diffused  or  cotyledonary.  Mammae  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. 
Modification  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.    Eeduction  of  the  ulna  from  a  complete 
and  distinct  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   navicular  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 -cylinder. 

4.  Development  of  horns  or  antlers  on  the  frontal  bones,  and 
gradual  complication  of  form  of  antlers. 

5.  By  inference  only,  increasing  complication  of  stomach^with 
ruminating  function  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  proba- 
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  Arctic  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.  Hippopotamidce ;  II.  Suidce,  or  true  Pigs;  III. 
Dicotylidce,  or  Peccaries.1 

Family  HIPPOPOTAMIDCE. 

Muzzle  very  broad  and  rounded.  Feet  short  and  broad,  having 
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  large, 
curved,  continuously  growing ;  those  of  the  upper  jaw  directed 
downwards.  Stomach  complex.  No  caecum. 

Hippopotamus.2 — This  genus  may  be  taken  to  include  all  the 
known  members  of  the  family ;  it  appears  to  have  been  always 

1  In  the  table  on  p.  89  the  Peccaries  are  included  in  the  Suidce. 
-  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  101  (1766). 


HIPPO  PO  TAMIDsE 


279 


m 


The  dentition  may  be  expressed  by  the 
The  crowns  of  the  molars  (Fig.  100) 


confined  to  the  Old  World, 
formula  i  .     „,  c  T,  p  - 

1 — 8         I         4 

when  worn  present  trefoil-shaped  surfaces  of  dentine ;  and  those  of 
the  premolars  are  sharp.  The 
facial  portion  of  the  skull  is  much 
elongated,  the  orbits  are  tubular 
and  very  prominent,  and  the  man- 
dible has  a  large  rounded  descend- 
ing flange  at  its  angle.  The  ears 
are  small,  the  tail  is  short,  and  the 
legs  are  likewise  so  short  that  the 
belly  is  raised  but  a  little  distance 
above  the  ground.  The  brain  is 
not  richly  convoluted,  and  differs 
very  considerably  from  that  of 

,        -T..  ...  FIG.  100.— Gnnding  surface  of  a  worn  molar 

the    FlgS,  approximating   in  SOme        Of  Hippopotamus  amphibius.    (FromOwen.) 

respects  to  that  of  the  Camel  and 

Giraffe,  but  on  the  whole   standing  very  much   by  itself.      The 

stomach  of  the  common  species  is  of  enormous  dimensions,  having 


FIG.  101. — The  Hippopotamus  (Hippopotamus  amphibius). 

an  axial  length  of  1 1  feet,  and  measuring  upwards  of  1 5  feet  along 
the  greater  curvature.  Its  axis  is  longitudinal,  the  pylorus  being 
situated  almost  in  the  pelvis,  and  it  is  divided  into  three  distinct 
compartments,  of  which  the  third  is  cylindrical.  The  liver  of  the 
adult  is  of  extremely  simple  form,  elongated  transversely,  and  narrow 
from  above  downwards.  With  the  exception  of  a  few  tufts  of 


280  UNGULATA 


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.  amphibius),  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  1 2  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 
(H.  liberiensis),  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  Chceropsis  ;  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 


SUID^E  28 1 

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  H.  palceindicus  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.  sivalensis  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  (H.  pentlandi  and  H.  minutus)  have  left  their 
remains  in  enormous  quantities  in  the  caves  and  fissures  of  Sicily 
and  Malta. 

Family  SUIDJE. 

An  elongated  mobile  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  flattened.  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  cardiac  orifice. 
A  caecum.  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.  FIG.  102.— Grinding  surface  of  a  worn 

gusi — Dentition :  i  f ,  c  ±,  p  4-,  m  third  risht  lower  molar  of  the  Wild  Boar 

.,          1^1   jj         TT  •  1'     •        (Susscrofa).    After  Owen. 

f ;  total  44.      Upper  incisors  dimin-  v 

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  slightly 
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.  Syst.  Nat.  12th  ed.  vol.  i.  p.  102  (1766). 


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.1  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  Avith  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  :  i  f ,  c  ^,  m  f  ;  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.  Vertebne  :  C  7,  D  13-14,  L  6,  S  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.  1 03-1 05)  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- 
maxillae  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. 


SUID^E 


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 
Japan.  The  fol- 
lowing    among 
others      have 
been    admitted 
by     many    zo- 
ologists as  dis- 
tinct     species  : 

£>US     SCrOjCl,          pI0.  103.— Left  lateral  view  of  the  dentition  of  the  Boar  (Sits  scrofa), 
the    \Vild    Boar    the  roots  of  the  teeth  being  exposed  by  removing  the  external  lamina 

of  Europe,  Asia  ofbone' 

Minor,  and  North  Africa,  once  common   throughout  the  British 

Isles  ;    S.  sennaaretisis,  North-East  Africa ;   S.  cristahts,   India ;    S. 


FIG.  104. — Left  lateral  view  of  the  skull  of  Sits  longirostris.    }  natural  size.    (From  Nehring.) 

vittatus,    Java,    Borneo,    Amboyna,    Batchian  ;>  S.  papuensis,   New 
Guinea ;   S.  timorensis,  Timor  and  Eotti ;   S.  andamanensis,  Anda- 


UNGULATA 


man  Islands ;  S.  taevanus,  Formosa ;  S.  leucomystax,  Japan ;  S. 
veirucosus,  Java,  Borneo,  Ceram ;  S.  barbatus,  Borneo ;  S.  celebensis, 
Celebes,  Philippines,  and  Moluccas ;  S.  longirostris,  Borneo  and 
Java.  The  last  four  species  form  an  allied  group  in  which  the 
facial  portion  of  the  skull  may  be  greatly 
elongated ;  S.  barbatus  and  S.  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.  S.  vittatus,  S.  leucomystax,  S.  cris- 
tatus,  S.  taevanus,  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 
well  ascertained  that  no  animal  of  the 
kind  ever  lived  upon  the  island  until 
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  historic  evidence  upon  the  subject, 
and  the  researches  of  naturalists,  as  Nathusius,  Riitimeyer, 
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 


FIG.  105.  —  Frontal  aspect  of 
the  cranium  of  Sits  longirostris. 
I  natural  size.  (From  Nehring.) 


SUID& 


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 


FIG.  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 


UNGULATA 


specimens  in  the  London  Zoological  Gardens  has,  however,  afforded 
opportunities  for  careful  anatomical  observation.1 

C. — Two  well-marked  species  of  African  Swine  have  been  with 
more  reason  separated  under  the  name  of  Potamochcerus.  The  denti- 
tion differs  from  that  of  the  true  Sus,  inasmuch  as  the  anterior 
premolars  have  a  tendency  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 


FIG.  107. — The  Red  River-Hog  (Sus  porous).    From  Sclater,  Guide  to  Animals 
in  Zoological  Society's  Gardens,  1883,  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  S.  africanus,  the  South  African  River-Hog, 
or  Bosch-Vark,  of  a  gray  colour,  and  the  other  S.  porcus,  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.  salvanius,  present  the  striped 
character  of  the  true  Sus,  a  strong  indication  of  close  affinities, 
whereas  in  all  the  following  forms  this  is  absent. 

The  genus  Sus,  in  the  above  extended  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,  Proc.  ZooL  Soc.  Lmid.  1883,  p.  413. 


SUID&        .  287 

S.  falconeri  arid  S.  karmdiensis  are  characterised  by  the  extremely 
complex  structure  of  the  molars,  in  which  they  show  decided  signs 
of  approximation  to  the  Wart-Hogs ;  the  same  feature  being 
exhibited  by  S.  phacochwroides  of  the  Algerian  Pliocene.  S.  titan 
and  S.  giganteus,  of  the  Indian  Pliocene,  together  with  S.  antiquus 
and  S.  erymanthius,  of  the  corresponding  European  deposits,  are  very 
large  species  characterised  by  their  comparatively  simple  molars ; 
S.  titan  being  fully  as  large  as  a  Tapir.  S.  hysudricus  of  the  Pliocene 
of  India,  and  S.  palceochcerus  of  that  of  Europe,  are  smaller  allied 
species  not  improbably  related  to  S.  andamanensis,  with  which  they 
agree  in  molar  structure.  S.  arvernensis,  of  the  Upper  Pliocene 
of  France,  appears  to  be  allied  to  S.  africanns ;  while  in  the 
diminutive  S.  punjabiensis  of  the  Pliocene  of  North- Western  India 
we  probably  have  the  direct  ancestor  of  S.  salvanius. 


FIG.  108. — Head  of  Babirusa  (Dabirusa  alfurus). 

Babirusa.1 — Dentition :  i  f ,  c  \,  p  f ,  m  | ;  total  34.  The  total 
number  of  teeth  is  therefore  considerably  reduced,  the  outer  upper 
incisor  and  the  two  anterior  premolars  of  both  jaws  being  absent. 
The  molars,  especially  the  last,  are  smaller  and  simpler  than  in  Sus; 
but  the  great  peculiarity  of  this  genus  is  the  extraordinary  develop- 
ment of  the  canines  of  the  male.  These  teeth  (Fig.  108)  are 
ever-growing,  long,  slender,  and  curved,  and  entirely  without  enamel 
covering.  Those  of  the  upper  jaw  are  directed  upwards  from  their 
base,  so  that  they  never  enter  the  mouth,  but  piercing  the  skin  of 
the  face,  resemble  horns  rather  than  teeth,  and  curve  backwards, 
downwards,  and  finally  often  forwards  again,  almost  or  quite 
touching  the  skin  of  the  forehead.  Vertebrae:  C  7,  D  13,  L  16, 
S  4.  There  is  but  one  species  (B.  alfunis),  found  only  in  the 
islands  of  Celebes  and  Bum.  Its  external  surface  is  almost 
1  Lesson,  Man.  d.  Mamm.,  p.  337  (1827),  "Babirusa." 


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." 

Phacochcerus.1 — 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  i  J,  c  ^,  p  -f ,  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  Phacochoerus ;  and,  as  already  mentioned,  some  of  the  fossil 
Indian  and  African  species  of  Sus  indicate  the  mode  in  which  this 
1  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,  ^Elian'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  DICOTYLID.E. 

Snout  as  in  Suidce.  Dentition :  i  § ,  c  y,  p  ^,  m  f ;  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  caecum. 
Confined  to  the  New  World. 

Dicotyles.1 — 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.  Vertebrae:  C  7,  D  14,  L  5,  S  4,  C  7. 
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,  Regjie  Animal,  vol.  i.  p.  237  (1817). 
19 


2QO  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,1  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  Eed  River  of  Arkansas  through  the  forest  districts  of 


FIG.  109.— The  Collared  Peccary  (Dicotyles  tajam). 

Central  and  South  America  as  far  as  the  Rio  Negro  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 

1  Professor  Cope  considers  that  there  is  a  third  species,  for  which  lie  has  pro- 
posed the  name  D.  angularis. 


DICOTYLID&  291 


inflicting  severe  wounds  with  its  sharp  tusks.  A  hunter  who  en- 
counters a  herd  of  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  of  villages  and  cultivated  lands  they  often 
inflict  great  devastation  upon  the  crops  of  the  inhabitants. 

Remains  of  the  two  existing  species  of  Peccary,  as  well  as  of  one 
much  larger  extinct  form,  are  found  in  the  cavern-deposits  of  Brazil ; 
while  large  Peccaries  also  occur  in  the  Pleistocene  of  the  United 
States,  which,  although  they  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  Dicotyles. 

Allied  Extinct  Genera. — In  the  Tertiary  deposits  of  both 
the  Old  and  New  World  occur  remains  of  Pig -like  animals 
which,  so  far  as  we  can  judge,  appear  to  connect  the  Peccaries 
so  closely  with  the  true  Pigs  as  to  render  the  Dicotylidw 
really  inseparable  from  the  Suidce.  Of  these  the  American 
genus  Chcenohyus  has  the  lower  canine  with  a  triangular  cross 
section  and  received  into  a  notch  in  the  upper  jaw,  as  in  the  Pec- 
caries, but  the  fourth  upper  premolar  is  simpler  than  the  molars,  as 
in  the  under-mentioned  genus  Hyotherium.  The  typical  forms  have 
only  three  premolars,  but  in  others,  which  it  has  been  proposed  to 
separate  generically  as  Bothriolabis,  there  are  four  of  these  teeth. 
Hyotherium,,  of  the  Pliocene 
and  Miocene  of  the  Old 
World,  is  a  generalised 
form  allied  both  to  Sus  and 
Dicotyles  as  well  as  to  certain 
extinct  genera.  The  upper 
molars  (Fig.  110)  are  char- 
acterised by  their  square 

Crowns    the  last    having   no    Fl°-  HO.—  The  three  left  upper  molars  of  Hyotheriwn 
T    ..       '  .  T  .    T  T    T  perimense.  from  the  Pliocene  of  India. 

distinct  third  lobe,  and  com- 
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  infoldings  of  the 
enamel,  producing  a  very  complex  pattern  on  the  worn  crowns. 
The  European  Miocene  genus  Listriodon,  with  the  dental  formula 
i  |}  c  |5  p  3^  m  3s  differs  from  all  the  preceding  in  having  the 
anterior  and  posterior  pairs  of  tubercles  of  the  molars  united  into 
ridges  running  across  their  crowns,  so  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 
palaeontological  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. 

Chceropotamidce. — In  this  family  the  molars  are  intermediate  in 
structure  between  those  of  the  Suidce  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  Cebochozrus  of 
the  Phosphorites  of  Central  France ;  Cfueropotamus  of  the  Upper 
Eocene,  the  type  species  of  which  was  of  the  size  of  a  large  Pig, 
with  the  dental  formula  if,  c  ^,  p  £,  m  f ,  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  Achcenodon  should  perhaps  also  be  placed  here. 

Anthracotheriidce. — The  genera  Anthracotherium  and  Hyopotamus, 
of  the  upper  Eocene  and  Miocene, 
have  the  typical  Eutherian  dental  for- 
mula;  the  upper  molars  (Fig.  Ill) 
carrying  three  columns  on  the  anterior 
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 

Fio.in.-The  imperfect  third  left  tne  columns  or  tubercles  will  be  at 
upper  molar  of  Hyopotamus  giganteus,  once  apparent,  the  broken  antero- 
JJJSiSJ*  (Fr'  "  median  column  representing  the  proto- 

conule.      Some  of   the    species   are    of 
large  size,  while  others  are  comparatively  small. 


EXTINCT  FAMILIES 


293 


Fl°-  112-—  A  ri8ht 

upper  molar  of  Mery- 


Merycopotamus.  —  The  genus  Mwycopotamus  of  the  lower  Pliocene 
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 
quadrituberculate  ;  and  may  thus  be  regarded  as 
typical  examples  of  the  brachy-selenodont  modifica- 
tion of  molar  structure. 

Cotylopidce.  —  The  Miocene  genus  Cotylops  (Ore- 
odon1}  is  the  type  of  a  large  American  family  in 
which  the  upper  molars  are  selenodont  and  usually 

,  ,  i  -i     .L-I       i  •        •  • 

have  tour  columns,  while  the  lower  canine  is  approxi- 

mated  to  the  incisors  and  its  form  and  function  Pliocene,    India. 

assumed   by  the  first   premolar.     The  last  upper  <Fromr  *he  Pal(eont°- 

•       •  i  i  rm  •  logia  Indica.) 

premolar  is  simpler  than  the  molars.  I  here  is  no 
flange  to  the  angle  of  the  mandible  ;  and  the  feet  have  four  digits. 
The  affinities  of  this  peculiar  family  are  probably  widely  spread, 
but  they  may  have  been  derived  from  the  Anthracotheriidce.  The 
type  genus  has  the  full  Eutherian  dentition,  but  in  some  of  the 
more  specialised  forms  (Cyclopidius)  the  upper  incisors  may  be 
wanting,  and  large  vacuities  occur  in  the  lachrymal  region.  The 
generalised  genus  Protoreodon,  of  the  Upper  or  Uinta  Eocene,  has 
five  cusps  on  the  upper  molars,  arranged  as  in  the  Anthracotheriida1. 
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  outer  columns,  outer  wall  of  upper 
molars  concave  and  inclined  inwards.  —  Agriochcerinw 
(Agriochcerus). 

2.  Orbits  closed,  a  lachrymal  fossa,  no  diastema,  the  last  upper 

premolar  with   one  outer  column  ;  outer  wall   of  upper 
molars  flattened.  —  Cotylopince  (Cotylops,  Eporeodon,  Mery- 
cochcerus,  Cyclopidius,  etc.) 
II.  Upper  molars  with  five  columns.  —  Protoreontince  (Protoreodon). 

Aiwplotheriidce. 
—  This  family  in- 
cludes several 
Upper  Eocene 
European  genera, 
with  selenodont 
upper  molars, 
carrying  five 
columns  arranged 
as  in  Anthraco- 
tlierium.  One  of 


FIG.  113. — Restoration  of  Anoplotherium  commune 
(Upper  Eocene).    Cuvier. 


This  name  (Leidy,  1851)  is  preoccupied  by  Orodus  (Agassiz,  1838). 


294  UNGULA  TA 


the  earliest  known,  Anoplotherium,  was  fully  described  by  Cuvier 
from  remains  found  in  the  Paris  gypsum-beds  (Upper  Eocene). 
Its  forty-four  teeth  formed  a  series  unbroken  by  a  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  Xiphodon.  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 
Hyopotamus  picteti,  of  the  Swiss  Eocene,  is  a  species  of  Dacrytherium. 

Ccenotheriidce. — 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  Anthracotheriidce.  The  known 
forms  are  from  the  Upper  Eocene  and  Lower  Miocene  of  Europe. 
In  Ccenotherium  the  molars  are  selenodont,  while  they  are  bunodont 
in  Dichobunus.  Homacodon,  of  the  Bridger  Eocene  of  the  United 
States,  is  closely  allied  to  the  the  latter.  The  first  lower  premolar 
of  Didwbunus  assumes  the  form  and  function  of  a  canine.  Spanio- 
therium  (Metriotherium)  is  a  much  larger  form,  in  which  the  molars 
are  not  unlike  those  of  Anthracotherium,  if  the  arrangement  of  the 
cusps  were  reversed ;  it  occurs  in  the  Eocene  Phosphorites  of 
France.  It  is  suggested  that  the  Tylopoda  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  Hyopotamw, 
tending  to  form  transverse  ridges ;  its  family  position  is  uncertain. 

Dichodontidce. — 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.  Dichodon,  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  Selenodonts.  In  Lophiomeryx,  of  the  same 
beds,  the  somewhat  higher  crowns  of  the  molars  approximate  to 
those  of  the  Cervidce,  but  the  hinder  lobes  of  the  upper  ones  are 
imperfectly  developed ;  the  genus  may  be  allied  to  the  Tragulidce. 
In  the  small  Gelociis,  of  the  Lower  Miocene,  the  molars  are  not 


CAMELID^E  295 

unlike  those  of  Dichodon ;  but  the  navicular  and  cuboid  bones  of 
the  tarsus  were  fused  together,  and  the  metatarsals  had  united  to 
form  a  "cannon-bone,"  although  the  metacarpals  still  remained 
distinct.  It  is  not  improbable  that  upper  incisors  were  wanting ; 
and  it  has  been  suggested  that  we  have  in  this  genus  the  ancestral 
type  of  the  Tragulidce  and  Cervida. 


TYLOPODA. 
Family  CAMELID.-E. 

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  Camelidce.  The  special  characters 
which  the  Llamas  and  Camels  have  in  common,  and  the  combina- 
tion of  which  distinguishes  them  from  the  rest  of  the  Artiodactyles, 
are  as  follows.  The  premaxillae  have  the  full  number  of  incisor 
teeth  in  the  young  state,  and  the  outermost  is  persistent  through 
life  as  an  isolated  laniariform  tooth.  The  canines  are  present  in 
both  jaws,  and  those  of  the  mandible  are  differentiated  from  the 
long,  procumbent,  and  spatulate  incisors,  being  suberect  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  premolars  is  usually  detached  from  the  series,  and  is 
of  simple  pointed  form.  The  auditory  bulla  is  filled  with  cancellous 
tissue.  The  hinder  part  of  the  body  is  much  contracted,  and  the 
femur  long  and  vertically  placed,  so  that  the  knee-joint  is  lower 
in  position,  and  the  thigh  altogether  more  detached  from  the 
abdomen  than  in  most  quadrupedal  mammals.  The  limbs  are 
long,  but  with  only  the  third  and  fourth  digits  developed ;  no 
traces  of  any  of  the  others  being  present.  The  trapezoid  and  mag- 
num of  the  carpus,  and  the  cuboid  and  navicular  of  the  tarsus  are 
distinct.  The  two  metapodial  bones  of  each  limb  are  confluent  for 
the  greater  part  of  their  length,  though  separated  for  a  considerable 
distance  at  the  lower  end.  Their  distal  articular  surfaces,  instead 
of  being  pulley-like,  with  deep  ridges  and  grooves,  as  in  other  recent 
Artiodactyles,  are  simple,  rounded,  and  smooth.  The  proximal 
phalanges  are  expanded  at  their  distal  ends,  and  the  wide,  depressed 
middle  phalanges  are  embedded  in  a  broad  cutaneous  pad,  forming 
the  sole  of  the  foot,  on  which  the  animal  rests  in  walking,  instead 
of  on  the  hoofs.  The  ungual  phalanges  are  very  small  and  nodular, 
not  flattened  on  their  inner  or  opposed  surfaces,  and  not  completely 
encased  in  hoofs,  but  bearing  nails  on  their  upper  surface  only. 
The  cervical  region  is  long  and  flexuous,  and  the  vertebrae  of  which 
it  is  composed  are  remarkable  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,  Macrauchenia  and  Myrmecopliaga).  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.1 
The  placenta  is  diffuse,  as  in  the  Suina  and  Tragulina,  not  coty- 
ledonary,  as  in  the  Pecora.  Finally,  the  Camelidce  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* — Dentition  of  adult :  i  ^,  c  ^,  p  f ,  m  •§• ;  total  34.  First 
upper  premolar  simple,  placed  immediately  behind  the  premaxillae, 
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  premaxillae  not  articulating 
with  the  arched  and  somewhat  elongated  nasals.  Vertebrae :  C  7, 
D  12,  L  7,  S  4,  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.  bactrianus,  Fig.  114).3  The  former 

1  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.  SysL  Nat.  12th  ed.  vol.  i.  p.  90  (1766). 

3  There  is  much  confusion  as  to  the  proper  use  of  the  names  Camel  and 
Dromedary.  It  is  now  generally  accepted  that  the  former  is  the  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 


CAMELWsE  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 


Fio.  114. — The  Bactrian  Camel  (CVmrfus  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.  At  such  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 


"  If  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.  In  a  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  vertebrae  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.1 — Dentition  of  adults  normally  :  i  J,  c  y,  p  f ,  m  f  ; 
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). 


CAMELID^E 


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  premaxillae.  Vertebrae:  C  7,  D  12,  L  7,  S  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 


FIG.  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 


3oo 


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  Natures,  of  Linnaeus.  They  were, 
however,  separated  by  Cuvier  in  1800  under  the  name  of  Laina, 
changed  by  Illiger  in  1811  to  Auchenia  (in  allusion  to  the  great 
length  of  neck,  av^ijv),  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  denning  their  dis- 
tinctive characteristics. 
These  are  (1)  the  Llama, 
Auchenia  glama  (Linn.),  or 
Lama  peruana  (Tiedemann) ; 
(2)  the  Alpaca,  A.  pacos 
Linn.) ;  (3)  the  Guanaco  or 
Huanaco,  A.  huanacus  (Mo- 
lina) ;  and  (4)  the  Vicugna, 

Fio.  H0.-Head  of  Vicugna,  from  an  animal  living      A     uimfjna  /Molina^)      or    A 
in  the  Gardens  of  the  Zoological  Society  of  London.          .      *  na/'    ( 

vicunna,    (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, 


CAMELID^E 


301 


FIG.  117. — Head  of  Guanaco,  from  an  animal  living 
in  the  Gardens  of  the  Zoological  Society  of  London. 


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, 
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 
it  is  as  vigilant,  wild,  and 
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,1  speaking  from  observation  on  wild  animals,  says : — 

"  It  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.  As  a  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  Naturalist'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  of  ten -quo  ted 
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  : — 

"  In  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  •  for 


CAMELID&  3°3 

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  A.  (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.  magiut,  the  premolars  were  reduced  to  one  in  each 
jaw ;  and  the  same  condition  obtains  in  A.  (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  Camelidce,  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 
is  i  f ,  c  ^,  p  %,  m  % ;  and  from  this  type  a  transition  may  be 
traced  to  Poebrotherinm,  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  Leptotraguhis  may  have  been  derived  from  the 


TRAGULID&  305 

Bunodont   genus   Homacodon   of   the    Bridger   Eocene,    mentioned 
among  the  Cienoiheriidce. 


TRAGULINA. 
Family  TRAGULID^E. 

No  teeth  in  premaxillse.  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  f ,  m  %.  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  bullse  of  skull  filled  with 
cancellar  tissue.  No  frontal  appendages.  Euminating,  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  Tragulidce, 
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 
Moschus,  one  of  the  Cervidce ;  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  palaeonto- 
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,1  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  T.  javanicus,  T.  napu,  T.  stanleyanus,  and 

1  Pallas,  Spidlegia  Zoologica,  vol.  xiii.  p.  27  (1779). 
20 


306 


UNGULATA 


T.  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. 
Dorcatherium l  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  i  $,  c  ^,  p  f,  m  f  =  34.  Vertebrae  :  C  7, 
D  13,  L  6,  S  5,  C  12-13.  The  only  existing  species,  D.  aquaticum 
(Fig.  118),  from  the  west  coast  of  Africa,  is  rather  larger  than  any 


Fio.  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.  naui),  only 
differing  in  size,  occur  in  the  lower  Pliocene  and  Miocene  of 
Europe ;  fossil  species  are  also  found  in  the  Indian  Pliocene. 
In  D.  naui  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 

1  Kaup,  Ossemens  Fossiles  de  Darmstadt,  pt.  5,  p.  92  (1836).  This  name, 
which  was  proposed  for  a  fossil  species,  antedates  Hyomoschus,  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  Cervidce,  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. 
Baehitheriwm,  of  the  French  Phosphorites,  apparently  presents 
affinity  with  Gelocus,  Prodremotherium,  and  Dorcatherium.  In  this 
genus  the  first  of  the  four  loAver  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  f ,  m  f,  in  con- 
tinuous series.  Auditory  bull*  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  arising  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  Euminants  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,  etc.)  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.  Zoo?.  Soc.  1877,  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  Cervidce  and  the  Bovidce, — otherwise  the 
antlered  and  horned  Euminants.  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,  Erahmatherium,  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  Palceomeryx,  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 
Sivatheriwm)  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 
skin  dies  and  peels  off,  leaving 
the  bone  bare  and  insensible, 
and  after  a  time,  by  a  process 
of  absorption  near  the  base,  it 
becomes  detached  from  the  skull 

FIG.  119.— A  shed  right  antler  of  the  Red  Deer   and    IS    "  shed  "    (Fig.   119).       A 
(Cervus  elaphus),  found  in  an  Irish  lake,    a,  Brow   more    Or    less    elongated   portion 

line.'  6(AfteVonJenyreS  ""* '  *'  "^  ^  ^    OT  "  ^^^  "  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  Cervidce,  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  tres  tine ;  the  mass  of 
points  at  the  summit  of  the  antler  being  termed  either  the  royal 
and  surroyal  tines,  or  collectively  the  crwon.  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. 


FIG.  120. — Head  of  Deer  (Cervus  schomburgki),  showing  antlers. 
From  Sclater,  Proc.  ZooL  Soc.  1877,  p.  682. 


L:\GULATA 


2.  The  horns  of  the  Bovidce  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 
(Antilocapra),  in 
which  the  horns 
also  differ  from 
those  of  all  others 
in  being  bifurc- 
ated. Horns  are 
not  present  at 
birth,  but  begin 
to  grow  very  soon 
afterwards.  The 
From  males  of  all  exist- 
ing Bomdce  possess 
them,  and  they  are  also  present  (though  usually  not  so  fully 
developed)  in  the  females  of  all  except  the  genera  Hoselaphus, 
Strepsiceros,  Tragelaphus,  Antilope,  sEpyceros,  Saiga,  Cobus,  Cervicapra, 
Pelea,  Nanotragus,  Neotragus,  Cephalophus,  and  Tetraceros ;  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 


FIG.  121.— Head  of  Antelope  (Gazella  granti),  showing  horns. 
Sir  V.  Brooke,  Proc.  Zool  Soc.  1878,  p.  724. 


PECORA 


generally  be  distinguished,  inasmuch  as  the  true  molars  of  the  Cervulce 
are  more  or  less  brachydont,  and  those  of  the  JBovidce  generally 
hypsodont,  i.e.  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.  1  2  3),  the  crown  being  lengthened 
and  the  root  small,  the  neck  does  not  come  up  FIO.  122.—  Crown  sur- 
to  the  alveolar  level  until  a  considerable  part  facf  of  "  ™rn  left  "Pi** 

....  ,.  *  molar  of  Palceomeryx  siva- 

oi  the  surtace  has  worn  away,  and  the  crown  of  unsis,  to  show  brachydont 


the  tooth  thus  appears  for  the  greater  part  of  type.  (From  the 
the  animal's  life  partially  buried  in  the  socket.  WM' 
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  necessarily  nearly  parallel,  whereas 


CT  FIG.  123. — Inner  and  outer  aspects  of  an  almost  unworn  left  upper  molar  of  the  Nilghai 
(Boselaphus  tragocamdus),  to  show  hypsodont  type.  (From  the  Palasontologia  Indica.) 

in  the  first-described  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  are  filled  up  with  cement,  which  gives  stability  to 
the  whole  organ,  and  is  entirely  or  nearly  wanting  in  the  short- 
crowned  teeth.  The  same  modification  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  Anchitherium  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  Cervidce 
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  Antelopes  (as 
Tragelaphus)  retain  much  of  the  brachydont  character,  which  is, 
however,  completely  lost  in  the  more  modern  and  highly  specialised 
Sheep  and  Oxen. 


FIG.  124. — Stomach  of  Ruminant  opened  to  show  internal  structure,  o,  (Esophagus ;  6, 
rumen  or  paunch  ;  c,  reticulum  or  honey-comb  bag ;  d,  psalterium  or  manyplies ;  e,  abomasuin 
or  reed ;  /,  duodenum. 

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  oasophagus  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. 


CERVID/E  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.  Eunning  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  laminse  arranged  very  much  like 
the  leaves  of  a  book,  and  very  close  together.  Their  surfaces 
are  roughened  by  the  presence  of  small  projections  or  papillae. 
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  CERVIDVE. 

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.  Gall-bladder  absent  (except  in 
Moschus).  This  family  contains  numerous  species,  having  a  wide 
geographical  distribution,  ranging  in  the  New  World  from  the  Arctic 


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  Mosehinse. — 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  Bovidce. 

Moschus.1 — The  Musk-Deer  (Fig.  125)  in  many  respects  stands  by 


FIG.  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.  Vertebrae  :  C  7,  D  1 4,  L  5,  S  5,  C  6. 
Among  the  anatomical  peculiarities  in  which  it  differs  from  all 
true  Deer  and  agrees  with  the  Bovidce  is  the  presence  of  a  gall- 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  91  (1766). 


CERVID^E  315 

1  (ladder.  The  hemispheres  of  the  brain  are  but  slightly  convoluted, 
and  the  cotyledons  of  the  placenta  are  arranged  in  a  peculiar  linear 
manner.1 

Although,  owing  to  variations  of  colour  presented  by  different 
individuals  in  different  localities  and  seasons,  several  nominal  species 
have  been  described,  zoologists  are  now  generally  agreed  that  there 
is  but  one,  the  Moschus  moschifems  of  Linnaeus.  In  size  it  is  rather 
less  than  the  European  Eoe  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 
belongs,  in  fact  the  large  majority  of  mammals,  have  some  portion 
of  the  cutaneous  surface  peculiarly  modified  and  provided  with 
glands  secreting  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  auatomy  of  Moschus  see  Flower,  Proc.  Zool.  Soc.   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  Cepvinse. — This  subfamily  includes  all  the  true  Deer. 
According  to  the  arrangement  proposed  by  Sir  V.  Brooke  l  the 
existing  Cervince  may  be  divided  into  the  sections  Plesiometacarpalia 
and  Telemetacarpalia. 

Plesiometaearpalia. — 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  premaxillae  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- 
maxillse  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.  Soc.  1878,  p.  889. 
2  De  Blainville,  Bull  Soc.  Philom.  1816,  p.  74. 


CERVID&  317 

Cervulus  of  Blainville  and  most  zoologists  ; — Styloceros  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 
graze.  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  Cervulus.  Sir  Victor  Brooke,  who 
investigated  this  question  in  1878  (see  Proceedings  of  the  Zoological 
Society  of  London  for  that  year,  p.  898),  came  to  the  conclusion  that 
there  are  certainly  three  which  are  quite  well  marked,  viz. — 

C.  muntjac  (Fig.  126),  found  in  British  India,  Burma,  the  Malay 
Peninsula,  Sumatra,  Java,  Hainan,  Banca,  and  Borneo.  The  general 


31 8  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  3 1  feet,  tail  7  inches,  height  26  to  28  inches.  The  female  is  a 
little  smaller.  The  specimens  from  Java,  Sumatra,  and  Borneo  are 


FIG.  126.— The  Muiitjac  (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  0.  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.  fcce. 

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  Palceomeryx. 

JElaphodus.1 — Antlers  very  small,  unbranched,  supported  on  long, 

1  Milne-Edwards,  Nouv.  Arch,  du  Mustum,  vol.  vii.  Bull.  p.  93  (1872). 


CERVID& 


319 


slender,  converging  pedicles.  Ascending  rami  of  premaxillae  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 


FIG.  127. — Male  of  Elaphodus  michianus.    From  Sclater  Proc.  Zool.  Soc.  1876,  p.  273. 

glands,  nor  is  there  a  tufted  gland  on  the  metatarsus.  The  limbs 
have  the  same  peculiarities  as  in  Cervulus,  but  the  mesocuneiform 
may  also  ankylose  with  the  ectocuneiform,  and  traces  of  the  nieta- 
carpals  may  remain.  The  hair  is  coarse  and  somewhat  quill-like. 

Cermis.1 — The  great  majority  of  the  Deer  of  the.  Old  World  may 
be  included  in  this  large  genus,  which  is  one  not  e&sy  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.  Vertebrae  :  C  7,  D  13,  L  6,  S  4,  C  11-14.  The  size  of  the 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  92  (1766). 


32o  UXGULATA 


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  Koe.  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.  eldi  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  Eusine  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.  davidiamis  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  Axine  group  includes  only  the  well-known  Axis  of  India, 
readily  distinguished  by  the  white  spots  with  which  the  body  is 


CERVID^E 


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  Palsearctic  region,  frequently  known  as  the  Pseudaxine 
group,  which  appears  to  connect  the  Axine  with  the  Elaphine 
type.  Well-known  representatives  of  this  group  are  C.  sika  (Fig. 
128)  of  Japan,  C.  mantchuricus  of  China,  and  C.  taevanus  of  Formosa. 


Fio.  128. — The  Japanese  Deer  (Ceraus  sika).    From  Lord  Powerscourt,  Proc.  Zool.  Soc. 
1884,  p.  200. 

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.  sihi  have  been  acclimatised  in  Ireland 
by  Viscount  Powerscourt,  at  Powerscourt,  County  Wicklow.  A 
number  of  Deer  from  the  Pliocene  of  Europe,  such  as  C.  perri&i 
and  C.  etueriarum,  appear  to  be  allied  both  to  the  Pseudaxine  and 
Axine  groups. 

The  Elaphine  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  Rusirie 
species)  are  spotted,  are  born  at  the  end  of  May  or  the  beginning 


FIG.  129.— Head  of  the  Wapiti  (Cervus  canade)isis). 

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 


CERVID^E  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  (C. 
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  Avhite  spots ;  there  is,  however,  a  uniformly  dark  brown  variety 
found  in  Britain.  The  bucks  and  does  live  apar.t,  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.  browni  and  C.  falconeri  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.  sedgemcki,  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. 

Telemetaearpalia. — 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,  Caprenh/.<. 
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). 


FIG.  130. — Skeleton  of  the  Gigantic  Irish  Deer  (Cervus  giganteus).    After  Owen. 

Rangifer,1 — The  Reindeer,  or  Caribou  as  it  is  termed  in  North 
America,  is  the  sole  representative  of  the  genus  Eangifer,  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 

1  Hamilton-Smith,  in  Griffith's  Animal  Kingdom,  vol.  v.  p.  304  (1827). 


CERVID& 


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 
branched  or 
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- 
ment of  the 
other.  The 

dentition  of  the  Eeindeer  is  frequently  remarkable  for  the  very 
small  size  of  the  posterior  lobe  of  the  last  lower  molar.  Vertebrae  : 
07,  D14,  L5,  S5,  C  11. 

The  Eeindeer  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 
Aveight  of  300  Ibs.,  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 


Fio.  131. — Skull  and  antlers  of  the  Reindeer  (Rangifer  tarandus), 
from  an  English  Pleistocene  deposit,  br,  Brow  tine ;  bz,  bez  tine. 
(After  Owen.) 


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. 

Alces.1— 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.  It  is  the  largest 
existing  member  of  the 

Fio.  132.— Hinder  part  of  the  base  of  the  cranium  of  the  family,  attaining  SOHie- 
Virginian  Deer  (Cariacus  virginianus).  From  Garrod,  Proc.  times  a  height  of  8  feet 
Zool.  Soc.  1877,  p.  13.  •? ,  r™ 

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  Ibs. 
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  great 
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  Avay.  The  antlers  and  hoofs  are  his 
i  Hamilton-Smith,  in  Griffith's  Animal  Kingdom,  vol.  v.  p.  303  (1827). 


CERVID&  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. 

Cervakes.1 — A  remarkable  extinct  Deer  from  the  Pleistocene  of 
North  America,  described  as   Cewalces,   appears  in  some  respects 


FIG.  133.— Head  of  Elk  (Alces  machlis). 

(although  a  true  Telemetacarpalian)  to  connect  Alces  with  Germs. 
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.  Premaxillse  not 
always  articulating  with  nasals.  Auditory  bullae  slightly  inflated, 
rugose  externally.  Vertebrae  :  C  7,  D  13,  L  6,  S  6,  C  8.  Tail  very 
short.  Glands  in  fore  feet  rudimentary ;  large  in  hind  feet. 

The  Roe,  or  Roe  Deer  (Capreolm  caprea),  is  a  small  form  dis- 

1  Scott,  Proc.  Ac.  Nat.  Sci.  Philad.  1885,  p.  181. 
2  Hamilton-Smith,  in  Griffith's  Animal  Kingdom,  vol.  v.  p.  313  (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  Capi-eolus. 

Hydropotes.1 — No  antlers  in  either  sex.     Lachrymal  fossa  deep 
and  short  (Fig.  134);  lachrymal  vacuity  of  moderate  size.     Orbits 


PIG.  134.— The  left  lateral  view  of  the  skull  of  a  male  Chinese  Water  Deer  (Hydropotes 
inermis),  with  the  wall  of  the  maxilla  cut  away  to  show  the  root  of  the  canine.  J  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.  Vertebrae :  C  7,  D  1 2, 

L  6,  S  4,  C  10.  No  tufts 
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 

Fio.  135.-Upper  surface  of  the  brain  of  Hydropotes      different  in  Other  respects. 
inermis.    (From  Garrocl,  Proc.  Zool.  Soc.  1877,  p.  792.)      Thus    the  brain    (Fig.   135) 

has  the  hemispheres  much 

convoluted,  as  in  other  Cerviiue,  and  approximates  to  that  of  Pudua  ; 
1  Swinhoe,  Proc.  Zool.  Soc.  1870,  p.  90. 


CERVID^E  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  Platyp'osopus,  presents  such 
a  marked  remblance  to  Hydropotes  in  the  form  of  the  angle  as  to 
suggest  a  more  or  less  intimate  affinity. 

Cariacus.1 — Skull  (Fig.  132)  with  the  vomer  dividing  the 
posterior  nares  into  two  distinct  chambers ;  premaxillae  not  reach- 
ing nasals.  Antlers  never  greatly  exceeding  the  length  of  the  head. 
Lachrymal  vacuity  very  large,  and  lachrymal  fossa  small.  Auditory 
bullse  slightly  inflated.  Vertebra:  C  7,  D  13,  L  6,  S  4,  C  13.  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.  mexicamis  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. 
palndosus  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  bullae  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-like  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. 

Pudua.1 — Antlers  in  the  form  of  minute  simple  spikes. 
Distinguished  from  the  Coassine  group  of  Cariacus  by  the  articulation 
of  the  premaxillae  with  the  nasals  (as  in  the  Furciferine  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  Cenidce  occur,  of  which  the  more  important 
may  be  briefly  mentioned.  Amphitragulus,  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  Palceomeryx  (Dremotherium  or  Micro- 
meryx)  generally  has  but  three  lower  premolars,  and  the  brachydont 
upper  molars  (Fig.  122),  like  those  of  Amphitragulus,  want  the  small 
accessory  inner  column2  found  in  modern  Deer.  In  P.  feignouxi,  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^E. 

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  ankylosed  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,  Proc.  Zool.  Soc.  1850,  p.  242. 

2  This  accessory  column  is  shown  in  the  figure  of  the  molar  of  Bosetyjhus  on 
p.  311. 


. 


GIRAFFID& 


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.  Vertebrae : 


Fio.  136.— The  Giraffe  (Gira/a  camelopardalls). 

C  7,  D  14,  L  5,  S  3,  C  20.     Gall-bladder   generally  absent,     Male 
reproductive   organs  and  placenta  of  a  Bovine  type.     Dentition : 

»fc«fcl»l>w»f. 

Giraffa.1 — The  Giraffe  (G.  camelopardalis)  is    the    sole    existing 
representative  of  the  genus,  now  confined  to  the  Ethiopian  region. 

1  Zimmermann,  Geograph.  Gcschichte,  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 
vertebrae  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  S.  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  Extinct  Types. — The  Pliocene  deposits  of  many  parts  of  the 
Old  World  yield  remains  of  a  number  of  large  Kuminants  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  Bovidce,  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  Giraffidce  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  Vishnutherium,  of  the  Pliocene  of  India 
and  Burma,  seems  to  make  the  nearest  approach  to  the  Giraffe,  but 


ANTILOCAPRID&  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.  Sivatherium  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  ANTILOCAPRID^E. 

Closely  allied  to  ihefiovidce,  but  the  horns  deciduous  and  branched. 

Antilocapra.1 — 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  BOVID^E. 

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  Cervidce  the  number  is  usually  from  5  to  12,  Capreolus  and 
Hydropotes  having  the  fewest.  In  Giraffa  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  Bovidce,  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.1 — If  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  mammae. 
1  Blainville,  Bull  Soc.  Philom.  1816,  p.  75. 


BOVIDsE 


335 


In  the  typical  forms,  such  as  the  Bubaline  Antelope  (A.  buba- 
linus),  the  Harte-beest  (A,  caama,  Fig.  137),  and  the  Tora  Antelope 
(A.  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  nearly  white  beneath.  In  the  Pliocene  of  the  Siwalik  Hills  in 
Northern  India  there  occur  remains  of  an  Alcelaphus  (A.  palceindicus) 


FIG.  13".— The  Harte-beest  (Alcelaphus  cacuiia). 

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  (A.  albifrons)  and  Bontebok  (A.  pygargus), 
belonging  to  the  genus  Damalis  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  Mr.  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.  A.  (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. 


FIG.  138.— Head  of  Alcelaphus  torn.    From  Sclater,  Proc.  Zool.  Soc.  1873,  p.  762. . 

Connochcetes.1 — Head  short  and  massive,  with  the  muffle  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  premaxillse  much  expanded  laterally, 
and  firmly  ankylosed.  Vertebrae  :  C  7,  D  14,  L  6,  S  4,  C  16. 
Hoofs  very  narrow.  Tail  very  long,  covered  throughout  with  long 
hairs.  Four  mammae.  Two  species,  C.  taurina  and  C.  gnu  (Fig.  139), 

1  Lichtenstein,  Berlin  Ges.  Natuforsch.  Freunde  Magazin,  vol.  vi.  pp.  152, 165 
(1814). 


BOVID/E 


337 


both  from  South  Africa.  The  former,  or  Brindled  Gnu,  is  distin- 
guished by  the  absence  of  long  hair  on  the  face,  the  black  (instead 
of  Avhite)  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 


FIG.  139.— The  White-tailed  Gnu  (Cowudueta  gnu). 

development  of  the  horns  of  the  Gnu,1  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  base 
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. 

1  F.  E.  Blaauw,  Proc.  Zool.  Soc.  1889,  p.  2. 
22 


338  UNGULATA 


CepJuilophus.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  bulk  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  vertebrae  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 
(T.  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 
no.  i40.-Paiatai  and  outer  aspects  of  been  provisionally  referred  to  this 

the  three  right  upper  premolars  and  first    genUS. 

molar  of  the  Four-horned  Antelope  (Tetra-  CervicdWl'ine      Section.  -  Small      Or 

ceros  quadrlcornis).      From  the  Palceon-    -,  A     f  -,  c       i   ±     ±1. 

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.s  —  Distinguished  from  the  next  genus  by  having  the 
crown  of  the  head  tufted,  muzzle  hairy,  premaxillse  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.  saltianus),  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   Griffith's  Animal  Kingdom,  vol.   iv.  p.   258  (1827). 
Taken  to  include  Grimmia,  Terphonc,  etc.,  of  Gray. 
-  Leach,  Trans.  Linn.  Soc.  vol.  xiv.  p.  524  (1823). 
3  Hamilton-Smith,  in  Griffith's  Animal  Kingdom,  vol.  iv.  p.  269  (1827). 


BOVIDsE  339 

Mr.  W.  T.  Blanford 1  observes  that  "  the  JSeni-Israel,  or  Om-dig-dig, 
one  of  the  smallest  Antelopes  known,  abounds  on  the  shores  of 
the  Eed  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  f rentals,  and  rising 
immediately  above  postorbital  process  of  frontals,  in  front  of  the 
fr  onto -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.3 

The  typical  species  is  the  Royal  Antelope  (N.  pygmceus)  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 
N.  scoparia,  N.  montanus,  and  JV.  hastatus,  both  these  appendages 
are  present;  while  in  the  Oreotragine  group  (N.  melanotis  and 
N.  oreotragus)  the  former  are  present  and  the  latter  absent. 

Pelea* — Horns  rather  small,  compressed,  upright,  scarcely 
diverging,  and  placed  immediately  over  the  orbits.  No  suborbital 
gland,  nor  lachrymal  fossa ;  premaxillse  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.5 — 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. 

-  Sundevall,   Krnigl.    Vetensk.  Akad.  Handl.   for  1844,  p.   191.      Taken  to 
include  Calotragus,  Scopophorus,  Nesotragus,  Pediotragus,  and  Orcotragus  of  Gray. 

3  See  V.  Brooke,  Proc.  Zool.  Soc.  1872,  pp.  642  and  875. 

4  Gray,  Cat.  Ungulate  Mamm.  Brit.  Mus.  p.  90  (1852). 

5  Andrew  Smith,  Illustrations  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  premaxillee  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.  Helicoplwra,  from  the 
Lower  Pliocene  of  Attica,  is  regarded  as  allied  to  Cobus,  but  it  has 
no  distinct  supraorbital  pits. 

Cervicapra.1 — An  allied  South  African  genus  in  which  the  tail  is 
short  and  bushy  and  the  premaxillae  do  not  reach  the  nasals.  Three 
species. 

The  Keitbok  (C.  arundineum)  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 
Palsearctic,  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.2 — 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 

1  De  Blainville,  Bull.  Soc.  Philom.  1816,  p.  75.     Syn.  Eleotragus. 
-  Pallas,  Spicilcgia  Zoologica,  vol.  i.  p.  3  (1767). 


BOVID^E  341 

Jumna  which  cannot  be  distinguished  from  those  of  the  existing 
species. 

sEpyceros.1 — 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  (^.  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.  JE.  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.  Vertebrae  :  C  7,  D  13,  L  6,  S  4,  C  10.  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.3 — 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. 
Vertebrae  :  C  7,  D  13,  L  6,  S  4,  C  14.  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.  Velensk.  Akad.  Handl.  for  1845,  p.  271. 

2  Gr&ySList  Mamm.  Brit.  Mus.  p.  160  (1843). 

3  Hodgson,  Proc.  Zool.  Soc.  1834,  p.  81. 

4  De  Blainville,  Bull.  Soc.  Philom.  1816,  p.  75.     Is  taken  to  include  Procapra 
and  Tragops. 


342  .    UNGULATA 


patch,  and  bordered  externally  by  a  diffused  dark  line  (see  Fig. 
121,  p.  310).  The  Gazelles,  of  which  there  are  some  twentj'- 
four  existing  species,  are  typically  Palsearctic  desert  forms,  the 
Springbok  (G.  euclwre)  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 l  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.  rufifrom,  G.  l&vipes,  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.  soemmerringi,  G.  granti  (Fig.  121), 
G.  ihomsoni. 

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.  deperdita  (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. 
1  Proc.  Zool.  Soc.  1873,  p.  537.  Three  species  subsequently  described  are 
here  added  to  the  list. 


BOVID^E  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.1 — 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  4-£  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  (H.  leuco- 
phceus)  is  distinguished  by  the  glaucous  hue  of  the  hair.  The  other 
species  are  the  Equine  Antelope  (H.  equinus)  and  Baker's  Antelope 
(H.  bakeri)  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  redicornis  is  not  generically 
separable. 

Oryx.2 — 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  all  the  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  colour  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.  beisa)  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,  Bull.  Soc.  Philmn.  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 


FIG.  141. — The  Gemsbok  (Oryx  gazeUa). 

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.  beatrix)  is  a  much 
smaller  animal,  with  the  black  markings  confined  to  the  head,  fore 
limbs,  and  flanks.  Finally,  the  Leucoryx  (0.  leucoryx)  of  North 
Africa,  while  agreeing  in  size  with  the  Beatrix,  differs  by  its  curved 
horns  and  uniform  coloration. 

The  extinct  Palceoryx,  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  Hippotragus. 


BOVID^E  345 

Addax.1 — Horns  Avith  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  (A.  nasomaculatus),  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 ; 
premaxillse  reaching  nasals.  Muffle  large  and  moist ;  nostrils 
approximated.  Molars  hypsodont  or  brachydont.  Vertical  white 
stripes  frequently  present  on  the  body. 

a.  Hind  limbs  much  shorter  than  tliefore.     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  in  those  of  the  upper  jaw.  One 
Oriental  genus. 

Boselaphus.z — 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  inner  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,  Anal.  Nat.  1815,  p.  56. 

2  De  Blainville,   Bull.  Soc.  Phil&m.   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  Ethiopian  genera. 
Tragelaphws.1 — Female  hornless.    Horns  of  males  (Fig.  142)  over 


Fio.  142.— Head  of  Tragelaphits  gratus.    From  Sclater,  Proc.  Zool.  Soc.  1883,  p.  30. 

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. 


BO  VI D& 


347 


The  Harnessed  Antelopes  are  among  the  handsomest  of  the 
whole  group.  The  small  Guib  (T.  scriptus)  is  not  larger  than  a 
Goat,  but  T.  angasi  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. 
T.  grains  agrees  with  this  group  in  coloration  (the  mane  being 


FIG.  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,  T.  spekei,  while  having  the  long  hoofs  of  T.  grains,  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 
(T.  sylvaticus)  of  South  Africa  has  no  stripes,  and  short  hoofs. 

Strepsiceros.1 — Females  hornless.     Horns  (Fig.  143)  more  twisted 
than  in  Tragelaphus,  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  Somali-land  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,  Palceoreas,  which  occurs  in  the  Lower  Pliocene  of  Europe 
and  Algeria,  appears  to  present  affinities  both  to  Oreas  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  Antilope  torticornis,  of  the  French  Pliocene, 
1  Desmarest,  Mammalogie,  p.  471  (1822). 


BOVID&  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  Palceotragine  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 ; 
Palceotragoceros  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  Palsearctic  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.1 — Horns  short  and  cylindrical,  rising  perpendicularly 
from  the  forehead  for  some  distance,  then  bending  sharply  back- 
wards and  downwards,  forming  hooks  with  pointed  tips.  Premaxillae 
not  reaching  the  nasals.  One  species,  Palsearctic. 

The  Gemse,  or  Alpine  Chamois  (R.  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. 


35° 


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 


FIG.  144. — Nemorhcedus  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. 

Nemorhcedus.1 — 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  premaxillae 
not  quite  reaching  the  nasals.  Some  nine  species,  ranging  from 
the  Eastern  Himalayas  to  North  China  and  Japan,  and  southwards 

1  Hamilton-Smith,  in  Griffith's  Animal  Kingdom,  vol.  v.  p.  352  (1827). 


BOVIDsE  351 

to  Formosa,  the  Malay  Peninsula,  and  Sumatra.  The  smallest 
species  is  the  Himalayan  Goral  (N.  goral).  Of  the  larger  forms  we 
may  mention  the  Himalayan  Serow  (N.  bubalinus)  the  Cambing- 
Utan  (N.  sumatrensis)  of  Sumatra,  and  the  Japanese  N.  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." 

Haploceros.1 — The  Eocky-Mountain  Goat  (Haploceros  montanus), 
inhabiting  the  northern  parts  of  California,  appears  to  be  very 
closely  allied  to  Nenwrhcedus.  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. 

Budorcas? — The  Takin  (B.  taxicolor)  of  the  Mishmi  Hills  in 
Assam,  and  an  allied  species  from  Eastern  Tibet,  are  larger  forms 
apparently  related  to  Nemorhcedus,  but  with  a  much  greater  develop- 
ment of  the  horns.  The  horns  of  what  is  considered  to  be  the 
male 3  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- 
Avard  flexure  or  angulation.  The  horns  of  the  male  may  be  2  feet  in 
length,  with  a  basal  diameter  of  1 3  inches.  The  muzzle  is  hairy,  with 
a  small  naked  muffle.  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. 

Scarcely  anything  is  known  of  the  habits  of  the  Takin,  which 
never  appears  to  have  been  seen  alive  by  Europeans. 

Caprine  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   Griffith's  Animal  Kingdom,   vol.  v.  p.  354  (1827). 
Amended  from  ' '  Aplocerus. " 

2  Hodgson,  Journ.  As.  Soc.  Bengal,  vol.  xix.  p.  65  (1850). 

3  See  A.  0.  Hume,  Proc.  Zool  Soc.  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  Palaearctic,  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 


FIG.  145. — The  Alpine  Ibex  (Capra  ibex). 

the  latest  developments  of   the  Bovidce,  since  they  are   unknown 
before  the  Pliocene  period ;  and  are  essentially  mountain  forms. 

Capra.1 — Horns  flattened  from  side  to  side,  and  either  curving 
backwards  (Fig.  145)  or  spirally  twisted.  No  suborbital  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.  Vertebrae:  C  7,  D  13,  L  6,  S  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. 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  94  (1766). 


BOVID&  353 

The  European  Ibex  or  Steinbok  (Fig.  145),  which  may  be 
taken  as  a  typical  Goat,  stands  about  2£  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,1  gives  the  distribution 
of  the  various  species  of  Goats,  with  some  remarks  on  their 
peculiarities : — 

(1)  C.  ibex,  confined  to  the  Alps  of  Switzerland,  Savoy,  and 
the  Tyrol,  and  now  nearly  extinct,  except  where  artificially  pre- 
served. (2)  C,  sibirica,  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  Euboea, 
has  been  described  under  this  name,  and  is  apparently  nearly  allied 

1  Proe.  Zool.  Soc.  1886,  p.  314  ;  and  1887,  p.  552. 

2  Specimens  referred  by  Dinnik  to  C.  caucasica  have  been  made  the  types  of 
another  species — C.  severtzovi. 

23 


354  UNGULATA 


to  C.  cegagrus.  (9)  C.  u-alie,  an  apparently  well  -  characterised 
species  from  the  highest  ranges  of  Abyssinia.  (10)  C.  fakoncri ; 
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.  hylocnus, 
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.  sivalensis 
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.1 — 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. 
Vertebrse :  C  7,  D  1 3,  L  6,  S  4,  C  1 0-1 4.  Some  twelve  species, 
mainly  Palsearctic,  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.  nahura)  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 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  97  (1766). 
2  There  may  be  a  beard  on  the  throat,  as  in  0.  cycloceros. 


BOVID^E  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  poli,  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,1  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  Proc.  Zool.  Soc.  1884,  p.  326. 


356 


UNGULA  TA 


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 
0.  vignei.  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 


FIG.  146. — The  Moufflon  (Ovis  musimon).     From  a  living  animal  in  the  London  Zoological 

Gardens. 

India,  Persia,  and  Baluchistan,  0.  gmelini  of  Asia  Minor  and  Persia, 
0.  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  0.  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 
mentioned,  the  skull  and  horns  resemble  those  of  the  Bharal, 
although  the  tail  is  longer,  and  there  is  a  thick  fringe  of  long  hair 
on  the  throat,  chest,  and  fore  legs. 


•BOVID&  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  (Ovis  savigni),  apparently  allied  to  the  Argali,  has,  however, 
been  described  from  the  so-called  Forest-bed  of  the  Norfolk  coast. 

The  Sheep  was  a  domestic  animal  in  Asia  and  Europe  before 
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. 

Ovibos.1 — 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  caprine ;  no  suborbital  gland, 
no  lachrymal  fossa  or  fissure  in  skull ;  orbits  tubular  ;  a  large  narial 
aperture  and  very  short  nasals  ;  premaxillae  not  reaching  nasals. 
1  De  Blainville,  Bull.  Soc.  Philom.  1816,  p.  76. 


358  UNGULATA 


Tail  short,  and  molar  teeth  caprine.      One  existing  and  two  fossil 
species,  Palsearctic  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 


•  Fro.  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 


BOVID&  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  Avell  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 l  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,  Avas  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 
0.  camfrons)  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  Palsearctic, 
Oriental,  and  Ethiopian  regions,  with  one  Nearctic  species  and  an 
outlying  and  aberrant  species  in  Celebes. 

Eos.2 — The  whole  of  the  species  of  Oxen  were  included  by 
Linnaeus  in  the  single  genus  Bos,  and  although  the  species  have 
been  distributed  by  modern  zoologists  in  several  genera — such  as 
Anoa,  Bubalus,  Bison,  Poephagus,  Bihos,  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. 
2  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  98  (1766). 


BOVID&  36i 

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  premaxillae  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,1  namely  the  large  B.  coffer,  occurring  typically  at  the  Cape, 
but  said  by  this  writer  to  range  to  Abyssinia,  and  the  smaller 
B.  pumihis,  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.  1873,  p.  474. 


362  UNGULATA 


B.  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.  cequinoctialis  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.  coffer,  and  in  some  specimens 
they  are  more  widely  separated  on  the  forehead,  and  are  marked  at 
their  bases  by  distinct  rugse.  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  premaxillse  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  Linnaeus, 
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 

1  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. 


BO  VID^E 


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,1  "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. 


FIG.  148. — The  American  Bison  (Bos  americamis).    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  Eocky  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 ;  arid  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 


Fio.  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  premaxillse  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 


BOVID^E  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  vertebrae,  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 
premaxillse  do  not  join  the  nasals,  and  the  occipital  aspect  of  the 
skull  is  characterised  by  the  deep  incisions  made  by  the  temporal 
fossae.  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.  As  a  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 
Mi  than,  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. 


FIG.  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  palceogaurus,  is  believed  to  indicate 
a  species  nearly  allied  to  the  Gaur,  if  indeed  it  be  specifically 
distinct. 


BOVID&  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 
fossae  scarcely  intrude  upon  the  occipital  aspect  of  the  skull ;  and 
the  premaxillse  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  Csesar,  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  primigenius,  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.1  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  premaxillse 
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 
(B.  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  Scotlatid,  "  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.  Palseontological 

ABC 


FIG.  151. — Bones  of  right  fore  foot  of  existing  Perissodactyles.  A,  Tapir  (Tapirus  indicus), 
X I ;  B,  Rhinoceros  (Rhinoceros  sumatrensis),  x  t ;  C,  Horse  (Equus  caballus),  x  J.  U,  ulna  ; 
R,  radius  ;  c,  cuneiform  ;  I,  lunar ;  s,  scaphoid  ;  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 


PER1SSODACTYLA  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 -tAvo,  usually  twenty -three  in  the  existing  species.  Nasal 
bones  expanded  posteriorly.  An  alisphenoid  canal.  Femur  with 
a  third  trochanter.1  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  caecum  is  large  and 
capacious,  the  placenta  diffused,  and  the  mammae  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  Palceothermm.  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  TAPIRHXE. 

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.1  Outer  columns  of  upper  molars  conical.  Four  digits 
in  the  manus,  and  three  in  the  pes. 

Tapirus? — Dentition  i  f ,  c  1,  p  £,  m  § ;  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.  1ST.  Parker,  Proc.  Zool  Soc.  1882,  p.  775. 
-  Cuvier,  Tableau  fiUment.  de  I'Hist.  Nat.  p.  152  (1798) ;  ex  Brisson. 


TAPIRID^E  371 

triangular,  and  pointed  in  front.  Vertebrae:  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  maxillae  (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  (Tapirus  proper).  Three  species,  T.  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  limbs,  being  glossy 
black,  and  the  intermediate  part  of  the  body  white ;  T.  americanus 
(T.  terrestris,  Linn.),  the  common  Tapir  of  the  forests  and  lowlands 
of  Brazil  and  Paraguay  (Fig.  152) ;  and  T.  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  they 
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  T.  priscus,  T.  arvernensis,  etc.,  have  been 
found  in  France,  Germany,  and  in  the  Ked  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, 


Fio.  152. — The  American  Tapir  (Tapirus  americanus). 

as  they  are  known  to  have  extended  eastward  to  China  (T.  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  T.  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  during  such  an 
enormous  period  of  time. 

The  anatomy  of  the  soft  parts  of  the  Tapirs l  conforms  to  the 

1  See  J.  Murie,  Journ.  Anat.  and  Physiol.  vol.  vi.  p.  131, 1871  ;  W.  N.  Parker, 
Proc.  Zool.  Soc.  1882,  p.  768  ;  and  F.  E.  Beddard,  Proc.  Zool.  Soc.  1889,  p.  252. 


LOPHIODONTID&  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.  T.  americanus 
differs  from  T.  indicus  by  the  absence,  or  at  any  rate  the  less 
development,  of  the  intestinal  valvulae  conniventes,  the  presence 
of  a  moderator  band  in  the  heart,  the  shape  of  the  glans  penis, 
and  the  more  elongated  caecum,  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. 

Palceotapims. — 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, 
*  f  >  c  T>  P  f  >  m  f  >  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  vulpi- 
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  Eoliippus.  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 


FIG.  153. — Right  side  of  skull  of  Hyracotherinum  leporinum,  from  the  London  Clay.  J  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  ;  p  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  Lophiodontidce  and  Palccotheriidce  is  really  an  arbitrary  one. 
Epihippus,  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 

1  The  Swiss  P.  sidcrolithicus  has  only  one  cusp  in  the  last  upper  premolar. 


PA  L.-EO  THERIIDjE 


375 


with  the  Tapiridce ;  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  PAL^EOTHERIID.E. 

Molars  (Fig.   155)  brachydont,  with  the  valleys  between  the 
ridges  never  filled  with  cement ;  upper  premolars  either  simpler  than 


FIG.  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. 
Orbit  (at  least  usually)  confluent 
with  temporal  fossa.  Three  digits 
on  each  foot.  This  family  in- 
cludes extinct  genera  ranging  from 
the  Middle  and  Upper  Eocene  to 
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, 
Palceotherium,  was  made  known  to 
science  in  the  early  part  of  the  T 

*   JIT" ,  i  Palceotherium  magnum.    (Alter  uwen.j     /,  /, 

present    century    by    Cuvier,    Who  External  surfaces  of  outer  columns;  a,  postero- 

restored    the    skeleton    (Fig.    154)  external  column  (metacone);   6,  antero-ex- 

*  1 1  i  11  M       ii,      fj_i  temal  column  (paracone)  i  c.  postcro-intenial 

With  a  Short  neck  like  that  OI  the  column  (hypowne)  •  d,  antero-internal  column 

Tapirs,  although   it   has  been  Sub-  (protocone) ;  i,  anterior  intermediate  column 

Sequently     found     that    the     neck  (protoconule);  e,  median  valley;  g,  posterior 

was    considerably   longer.      This 

genus  (which  may  be  taken  to  include  Paloplotherium)  ranges  from 


FIG.  155.— A  half- worn  right  upper  molar  of 
(After  Owen.)     /,  /, 


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  Equidce.  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  Anchitherium  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  EQUID.E. 

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  Palceotheriidce. 
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  Equidce  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 


EQUID^E 


377 


(Fig.   156,  rt),  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  Hymcotherium  of  the  Lower,  and  Pachy- 
nolophus  of  the  Middle  and  Upper  Eocene  Avere  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  Equidce  occur  which 
approach  in  size  the  existing  Horse.  Important  structural  modi- 
fications have  also  taken  place,  with  corresponding  changes  in  the 


FIG.  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  (in),  a,  Pachynolophus  (Eocene) ;  b,  Anchitherium  (Early  Miocene) ;  c,  Anchitherium 
(Late  Miocene) ;  d,  Ilipparion  (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. Macrauchenia. 
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 


FIG.  157. — a,  Grinding  surface  of  unworn  molar  tooth  of  Anchitherlum ;  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 


EQUID^E 


379 


FIG.  158.- 


Outer  vi 


islands  of  the  two  softer  substances,  dentine  and  cement,  separated 
by  variously  reduplicated  and  contorted  lines  of  intensely  hard 
enamel,  resulted  (Fig.  157,  c).  The  crown  continued  lengthening 
until  in  the  modern  Horses  it  has  assumed  the  form  called  "  hyps- 
odont"  (Fig.  158,  b).  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  fl/lfl 
of  the  brachydont  ancestor.  It  is  per-  jft^jg-l^ 
fectly  clear  that  this  lengthening  of  the 
crown  adds  greatly  to  the  power  of  the 
teeth  as  organs  of  mastication,  and  en- 
ables the  animals  in  which  it  has  taken 
place  to  find  their  sustenance  among  the  lwer  raolar  tooth  ?f 

.         ,  .     .         .      ,     '  (brachydont  form) ;  6,  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 
Avell  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 


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.* — In  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- 
therium.  Each  foot  has  three  digits. 

Hipparion? — Upper  cheek-teeth   (Fig.   159),  with  the  antero- 


FIG.  159.— Three  right  upper  cheek-teeth  of  Hipparion.  a,  Antero-external  column ;  6, 
postero-external  column ;  c,  postero-internal  column,  or  posterior  pillar ;  d,  antero-internal 
column,  or  anterior  pillar ;  /,  posterior  intermediate  column  ;  i,  anterior  intermediate  column. 
(From  the  Palceontologia  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.  Either  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. 
2  Christol,  Ann.  Sci.  Indust.  Mid.  France,  vol.  i.  p.  180  (1832). 


EQUID^E  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. 

Equus.1 — 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,  e) ;  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  Hippidium,  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  (E.  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,  c. 

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  Equidce  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,  t  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  Eussia.  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, 

1  Darwin,  Variation  of  Animals  and  Plants  under  Dotnestication,  1868,  vol. 
i.  chap.  ii. 


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  E.  prezevalskii,1  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  E.  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  (E.  asinus),  and  the  wild 
Asses  of  Africa  (E.  asinus,  var.  africanus  and  var.  somalicus 2).  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  3  (E.  hemionus),  they  present  such  marked  variations  in  size 
and  form  that  they  have  commonly  been  divided  into  three — the 
Syrian  Wild  Ass  (E.  hemippus),  the  Onager  (E.  onager)  from  Persia, 
Baluchistan,  the  Punjab,  Sind,  and  the  desert  of  Kach,  and  the 
Kiang  or  Dzeggetai  (E.  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  (E.  quagga),  may  be  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  Eiver,  but  now,  in  common  with  most  of  the  larger  wild 
animals  of  that  region,  is  becoming  extremely  scarce,  owing  to  the 


Fio.  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  of 
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 


EQUIDsE 


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 
(E.  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 


1 


-c3&;i 
FIG.  161.— True  or  Mountain  Ze 


zebra). 

become  very  scarce,  and  is  even,  like  the  Quagga,  threatened  with 
extermination  at  no  distant  date.  The  second  species,  Burchell's 
Zebra  (E,  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. 

E.  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  Avhite  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. 


Fio.  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  strongly 
marked)  is  completely  isolated  in  its  posterior  half,  and  the  upper- 
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 


EQUID^E  387 

black  stripes  of  the  flanks  and  haunches  fainter  stripes  are  generally 
seen. 

E.  grevyi. — 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  Avith.  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 
Biu'chell'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 


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, 


EQUID^E 


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 


\ 


Fio.  103. — Side  view  of  skull  of  Horse,  with  the  bone  removed  so  as  to  expose  the  whole  of 
the  teeth.  PMx,  Premaxilla ;  MX,  maxilla  ;  Na,  nasal ;  Ma,  malar  or  jtigal ;  L,  lachrymal ;  F r, 
frontal ;  Sq,  squamosal ;  Pa,  parietal ;  oc,  occipital  condyle  ;  pp,  paroccipital  process ;  t1,  i2, 
and  i3,  the  three  incisors  ;  c,  the  canine  ;  pm1,  the  situation  of  the  rudimentary  first  premolar, 
which  has  been  lost  in  the  lower,  but  is  present  in  the  upper  jaw ;  pm-,  pm3,  and  pm*,  the 
three  fully  developed  premolars  ;  m1,  ?n2,  and  m3,  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  .  UNGULA  TA 


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  premaxillse.  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  maxillae,  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  bounded  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  betAveen  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  stylif onn 
process ;  and  completely  embraces  the  truncated  cylindrical  tym- 
panohyal,  which  is  of  great  size,  in  correspondence  with  the  large 


EQUID^E  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  vertebrae,  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  vertebras  are  elon- 
gated, strongly  keeled,  and  markedly  opisthoccelous,  or  concave 
behind  and  convex  in  front.  Their  neural  laminae  are  very  broad, 
the  spines  almost  obsolete,  except  in  the  seventh,  and  the  trans- 
verse processes  not  largely  developed.  In  the  trunk  vertebrae  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  nuchce,  or  "pax wax,"  which  passing  forwards  in  the 
middle  line  of  the  neck  above  the  neural  arches  of  the  cervical  ver- 
tebrae, to  which  it  is  also  connected,  is  attached  to  the  occiput  and 
supports  the  weight  of  the  head.  The  transverse  processes  of  the 
lumbar  vertebrae  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 
vertebras,  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 


392  UXGULATA 


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  Avith  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  (os  siiffraginis,  or  large  pastern),  a  very  short  middle  (os 
corona,  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." l 

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  "  pastern,"  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 
to  a  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  oi  the  fore  limb, 
1  This  must  not  be  confounded  with  the  navicular  of  the  tarsus. 


EQUIDsE 


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  011  the  ex- 
tremity of  the  single  digit  of  each  foot,  which  is  kept  habitually  in 
a  position  approaching  to  vertical. 

The  muscles  l  of  the  limbs  are  modified  from  those  of  the  ordi- 
nary mammalian  type  in  accordance  with  the  reduqed  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- 
f orm  these 
functions  in 
other  mammals 
are  absent  or 
rudimentary. 
.  Below..,,  the 


FIG.  164.— Section  of  foot  of  Horse.  1,  Metacarpal  bone  ;  2,  first 
phalanx  (os  suffraginis);  3,  second  phalanx  (as  corona);  4,  third  or 
urigual  phalanx  (os  pedis,  or  coffin-bone) ;  5,  one  of  the  upper  sesamoid 
bones;  6,  lower  sesamoid  or  "navicular"  bone;  7,  tendon  of  anterior 
Carpal  and  tar-  extensor  of  the  phalanges ;  8,  tendon  of  superficial  flexor  (fl.  perforate) ; 
9,  tendon  of  deep  flexor  (fl.  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  13,  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. 


sal  joints  the 
fore  and  hind 
limbs  corre- 
spond almost 
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  extensor  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  calcaneum.  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  papillae  or  laminae.  From 
this  the  horny  material  which  constitutes  the  hoof  is  exuded.  A 


EQUID&  395 

thickened  ring  encircling  the  upper  part,  called  coronary  cushion 
(13),  and  the  sole  (15),  are  covered  with  numerous  thickly  set 
papillae  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  laminae,  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  papillae  of  the  keratogenous  membrane, 
so  that  a  section  near  the  base  of  the  hoof,  cut  transversely  to  the 
long  axis  of  these  papillae,  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  papillae  of  the  skin,  are 
found  on  the  inner  face  of  the  fore  limb,  above  the  carpal  joint,  in 
all  species  of  JEquidce,  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  f ,  c  ^,  p  $, 
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, 

/  /          Id  e      *  from     Avhich     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 

PIG.  165.— Longitudinal  and  transverse  section  of  upper  quite  rudimentary,  often, 

incisor  of  Horse,    p,  Pulp  cavity  ;  d,  dentine  or  ivory  ;  e,  especially     in     the     lower 

enamel ;  c,  outer  layer  of  cement ;  cf,  inner  layer  of  cement,  •  ,    j         ••          -,  , , 

lining  a,  the  pit  or  cavity  of  the  crown  of  the  tooth.  ]aw>  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  in  a 
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 


EQUIDsE  397 

quadrate,  except  the  first  and  last,  which  are  nearly  triangular. 
The  lower  teeth  are  much  narrower  than  the  upper. 

The  milk  dentition  consists  of  i  f ,  c  $,  m  f  =  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  by  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  before  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  five  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 
accidental  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  back  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  elongated  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  papillae  near  the  base,  rather  irregular  in 
form,  about  a  quarter  of  an  inch  in  diameter  and  half  an  inch  apart. 
The  conical  papillae  are  very  small  and  close  set,  though  longer  and 
more  filamentous  on  the  intermolar  portion.  There  are  no  fungi- 
form  papillae  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 
fraenum  linguae.  The  sublingual  is  represented  by  a  mass  of  glands 
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  fraenum  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 


EQUID&  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  sac,  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  latter  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  cesophageal  orifice  is  very  small,  and  is  guarded  by  a  strong 
crescentic  or  rather  horse-shoe-like  band  of  muscular  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  ca3cum  is 
of  conical  form,  about  2  feet  long  and  nearly  a  foot  in  diameter ; 
its  walls  are  sacculated,  especially  near  the  base,  having  four  longi- 
tudinal fibrous  bands  ;  and  its  capacity  is  about  twice  that  of  the 
stomach.  It  lies  with  its  base  near  the  lower  part  of  the  abdomen, 
and  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 
fissures,  which  do  not  extend  quite  to  the  posterior  border  of  the 
organ ;  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  fcetal 
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 
tongue-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  the  spleen, 
and  a  shorter  right 
lobe.  The  principal 
duct  enters  the 
duodenum  with  the 
bile-duct,  and  there 
is  often  a  second 

FIG.  166.— Under  surface  of  the  liver  of  the  Horse,  u,  Umbilical  Small  duct  which 
fissure  ;  U,  left  lateral  lobe ;  Ic,  left  central  lobe ;  re,  right  central  Onens  SCDaratelv 
lobe ;  rl,  right  lateral  lobe ;  s,  Spigelian  lobe  ;  c,  caudate  lobe.  * 

near  to  this. 

Circulatory  and  Respiratory  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 


EQUID&  401 

understood.  The  larynx  has  the  lateral  sacculi  well  developed, 
though  entirely  concealed  within  the  alae  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  vesiculse  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  mammse  (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.  S.  Arloing,  "  Organisation  du  pied  chez  le  cheval,"  Ann. 
Sci.  Nat.  1867,  viii.  pp.  55-81  ;  H.  Burmeister,  Los  caballos  fosiles  de  la  Pampa 
Argentina,  Buenos  Ayres,  1875  ;  Chauveau  and  Arloing,  Traite  d'anatomie  com- 
paree  des  animaux  domestiques,  Paris,  1871,  and  English  edition  by  G.  Fleming, 
1873  ;  E.  Cuyer  and  E.  Alix,  Le  Cheval,  1886  ;  A.  Ecker,  "  Das  Europaische  Wild- 
pferd  und  dessen  Beziehungen  zum  domesticirten  Pferd,"  Globus,  Bd.  xxxiv. 
Brunswick,  1878  ;  Forsyth-Major,  "  Beitrage  zur  Geschichte  der  fossilen  Pferde 
besonders  Italiens,"  Abh.  Schw.  Pal.  Ges.  iv.  pp.  1-16,  pt.  iv. ;  George,  "  Etudes 
zool.  sur  les  Hemiones  et  quelques  autres  especes  chevalines,"  Ann.  Sci.  Nat. 
1869,  xii.  p.  5  ;  E.  F.  Gurlt,  Anatomische  Abbildungen  der  Haussdugethiere,  1824, 
and  Hand,  der  vergleich.  Anat.  der  Haiissaugethiere,  2  vols.  1822;  Huet,  "  Croise- 
ment  des  di verses  especes  du  genre  cheval,"  Nouv.  Archives  du  Museum,  2d  ser. 
torn.  ii.  p.  46,  1879  ;  Leisering,  Atlas  der  Anatomic  des  Pferdcs,  Leipsic,  1861 ; 
J.  M'Fadyean,  The  Anatomy  of  ihe  Horse,  1884  ;  0.  C.  Marsh,  "  Notice  of  New 
Equine  Mammals  from  the  Tertiary  Formation,"  Am.  Journ.  of  Science  and  Arts, 
vol.  vii.  March  1874;  Id.  "Fossil  Horses  in  America,"  Amer.  Naturalist,  vol. 
viii.  May  1874  ;  Id.  "  Polydactyle  Horses,"  Am.  Journ.  of  Science  and  Arts,  vol. 
xvii.  June  1879  ;  Franz  Miiller,  Lehrbuch  der  Anatomic  desPferdes,  Vienna,  1853 ; 
R.  Owen,  "Equine  Remains  in  Cavern  of  Bruniquel,"  Phil.  Trans,  vol.  clix. 
(1870),  p.  535  ;  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  KHINOCEROTIDJE. 

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  Lophiodontidce,  from  some 
members  of  which  they  are,  indeed,  probably  derived.  The  outer 
columns  of  the  upper  molars  (Fig.  167)  are,  however,  so  excessively 
A  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. 
The  skull  is  large,  with 
the  orbit  confluent  with 
the  temporal  fossa. 
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. 

Rhinoceros. l — 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). 


Fio.  167. — A  partially  worn  second  right  upper  molar  of 
Rhinoceros  antiquitatis.  Letters  as  in  Fig.  155  (p.  375),  ex- 
cept fc,  which  indicates  a  prolongation  of  the  median  valley. 
(After  Owen.) 


RHINOCEROTIDA.  403 


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  premaxillae,  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.  Vertebrae:  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.  368),  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 
papillae.  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  papillae.  It  results  from  this  that  the  horn  has  the 
appearance  of  a  mass  of  agglutinated  hairs,  which,  in  the  newly 
growing  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. 

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  molar  teeth  present  some 

amount  of  Specific  Variation  ;  FIG.  16S.— A  partially  worn  second  right  upper 
.1  „  ^.i,:!^  /TV™  molar  of  (A)  Rhinoceros  sondaicus,  and  (B)  R.  uni- 

thus    while    one    type    (Fig.  ^^    fc;  F;ssette  cut  off  from  median  valley ;  m> 

168,  A}  has  Only  a'  simple  crotchet;  n,  crisis,  or  corabing-plate ;  «,  anterior 
"  Crotchet  "  projecting  from  valley :  *>  anterior  intermediate  column.  Other 
,i  .  •  j  letters  as  in  Fig.  155,  p.  375. 

the  posterior  transverse  ridge 

into  the  median  valley,  in  others  (Fig.  168,  £)  this  crotchet  joins  a 
"  crista,"  or  "  combing-plate,"  projecting  from  the  outer  wall  to  cut 
off  a  distinct  f ossette  from  the  median  valley.  Occasionally,  however 


404  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  caecum. 

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. 

Rhinocerotic,  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.  unicornis  (Fig.  169)  of  Linnaeus,1  the 
largest  and  best  known,  from  being  the  most  frequently  exhibited 
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 

1  Many  authors  use  Cuvier's  name,  E.  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. 


RHINOCERO  TID^E 


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.  1 68,  b)  being  charac- 
terised by  the  presence  of  a  combing-plate  joining  the  crotchet,  and 


FIG.  169. — Indian  "Rhinoceros  (Rhinoceros  unicornis).  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,  a)  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,1  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  Ehinoceros ;  and  thereby  indicates  that  the  division  of 
the  existing  Khinoceroses  into  separate  genera  is  not  advisable. 


FIG.  170. — Javan  Rhinoceros  (Rhinoceros  sondaieus). 

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.  lasiotis,  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. 
2  See  Beddard  and  Treves,  Proc.  Zool.  Soc.  1889,  p.  9. 


RHINOCERO  TID^E 


407 


and  reference  has  already  been  made  to  the  resemblance  between 
the  visceral  anatomy  of  these  species.1  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,  f  inch  wide,  and  \  inch  deep.  The 
small  intestine,  in  place  of  the  villi  of  R.  unicornis,  has  throughout 
the  greater  part  of  its  length  a  uniform  series  of  thin  and  nearly  or 
quite  continuous  transverse  foldings,  like  the  valvulae  conniventes 
of  the  human  small  intestine.  There  is  no  gland  behind  the  foot. 


FIG.  171.— Posterior  aspect  of  the  liver  of  Rhinoceros  sumatrensis.  re,  Right  central  lobe  ; 
rl,  right  lateral  lobe ;  Ic,  left  central  lobe ;  II,  left  lateral  lobe ;  e,  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  Group. — 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.  sumatrensis  see  Garrod,  Proc.  Zool.  Soc. 
1873,  p.  92  ;  and  Beddard  and  Treves,  loc.  cit. 


408  UNGULAJA 


The  common  Two-horned  Khinoceros,  R.  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 


FIG.  172. — Common  African  Rhinoceros  (Rhinoceros  bicornis). 

as  great  as,  or  greater  than,  the  anterior  have  been  separated  under 
the  name  of  R.  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.1  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  ft.  simus  are  taken  from  a  paper 
by  Sclater  in  the  Proc.  ZooL  Soc.  1886,  p.  143. 


RHINOCEROTID&  4°9 

auditory  meatus.     Nothing  is  known  of  the  anatomy  of  the  soft 
parts  of  either  of  them. 

Burchell's  or  the  Square-mouthed  Rhinoceros  (It.  simus),  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  are  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.  Mr.  F.  C. 
Selous l  gives  the  following  description  of  its  habits  from  extensive 
personal  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  buffaloes  they 
lie  asleep  during  the  heat  of  the  day,  and  feed  during  the  night 
and  in  the  cool  hours  of  early  morning  and  evening.  Their  sight 
is  very  bad ;  but  they  are  quick  of  hearing,  and  their  scent  is  very 
keen ;  they  are,  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 
a  magnificent  broadside  shot,  while  under  similar  circumstances  a 
prehensile -lipped  rhinoceros  will  usually  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 
calf  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  vice  versd,  the  same 
position  is  always  exactly  maintained.  During  the  autumn  and 
winter  months  (i.e.  from  March  to  August)  the  square-mouthed 
1  Proc.  Zool.  Soc.  1881,  p.  726. 


4io  UNGULATA 


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,  in  a 
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.  unicornis,  so 
that  in  this  respect  R.  simus  has  the  same  relation  to  R.  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  Rhinoceros  in  its  widest 
signification,  a  very  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  R.  sivalensis  appears  to  have 
been  the  direct  ancestor  of  R.  sondaicus ;  while  R.  palceindicus  was 
probably  nearly  related  to  R.  unicornis,  although  the  upper  molars 
had  not  developed  a  combing-plate. 

R.  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 
R.  sansaniensis  was  a  closely  allied  if  not  identical  Jorin. 

•  The  Atelodine  group  was  very  widely  spread  in  past  epochs. 
Thus  the  huge  R.  platyrhinus  of  the  Indian  Pliocene,  and  the  equally 
large  R.  antiquitatis  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.  bicornis.  Several  species, 
such  as  R.  leptorhinus  (Fig.  173),  R.  megarhinus,  and  R.  etruscus, 
occur  in  the  European  Pleistocene  which  do  not.^resent  a  marked 
relationship  to  any  of  the  living  forms.  This  group  is  also  repre- 


RHINOCERO  TID^E  4 1 1 

sented  in  the  Pleistocene  of  Southern  India  by  the  small  11.  deccan- 
ensis  and  R.  Jcarnuliensis. 

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  R. 
sondaicus  (described  by  Lesson  as  R.  inermis)  to  the  same  genus. 
If  these  American  types  be  included  in  Rhinoceros  there  seems  no 
valid  reason  for  separating  the  European  Lower  Pliocene  and  Mio- 
cene forms  described  as  Aceratherium,  at  least  some  of  which  have 


Fio.  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  Elasmotherium,  from 
the  Pleistocene  ol  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 
Khinoceroses  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  loAver  canines  are  either  upright,  or  less  proclivous  than  in  the 
Rhinoceroses  ;  in  Metamynodon  the  premolars  are  reduced  to  2. 
Molar  teeth  from  the  Phosphorites  of  Central  France,  described 


Fio.  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  Cadiircotherium,  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.E. 

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, 


LAMBDOTHERIID^E,  ETC.  413 

while  there  may  be  traces  of  an  anterior  ridge.     The  premolars 
are  simpler. 

Lambdotheriidce. — This  family  is  confined  to  the  Upper  Eocene 
and  Miocene  of  North  America,  where  it  is  represented  by  Lambdo- 
therium,  Palceosyops,  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  loAver  molar  has  a  third  lobe.  Limnosyops  differs  from  Palceosyops 
in  having  two  inner  columns  to  the  last  upper  molar. 

Chalicotheriidce. — 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 vertebrae  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- 
theriurn  and  Moropus.  An- 
cylotherium,  of  the  Grecian 
Pikermi  beds,  is  founded  upon 
phalanges  which  indicate  an  Fio.  irs.-Anterior  and  distal  aspects  of 

allied      genUS.          The      Indian    phalangeal  bone  of  Chalicotherium  sivaknse.   (From 

species  of  Chalicotherium  is  dis-  the  p  <**»*****  ^«-> 

tinguished  by  the  loss  of  the  incisors  and  the  upper  canine ;  while 

all  the  species  want  the  first  premolar. 

Titanotheriidce. — This  exclusively  North  American  family  in- 
cludes gigantic  forms  closely  allied  to  the  Lambdotheriidce,  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,  Titanotherium  (Menodus,1  Brontotherium,  Symborodon, 
Allops,  etc.),  may  either  have  the  full  complement  of  teeth,  or  the 
incisors  may  be  reduced  to  £.  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  Ehinoceroses ;  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  MACRAUCHENIID.E. 

This  extinct  South  American  family  is  best  known  by  the  genus 
Macrauchenia,  as  represented  by  M.  patachonica  and  M.  boliviensis, 
Avhich  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,  M.  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  opisthocoelous  as 
in  the  allied  forms.  In  some  of  the  limb  characters  it  resembles 
the  Equidce,  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 Homalodontothemim  should  be  placed  here. 

Family  PROTEROTHERIHLE. 

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  palaeontologists,  our  knowledge  is 
at  present  necessarily  imperfect,  though  daily  extending.  It  will 


HYRACID^E 


415 


only  be  possible  here  to  give  details  of  some  of  the  more  interest- 
ing or  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^E. 

This  division  is  constituted  to  receive  a  single  family  of  mam- 
mals, the  affinities  of  which  have  long  constituted  a  puzzle  to 


Fio.  170. — Uyrax  capensis. 


zoologists.  They  were  first  placed  among  the  Rodents,  to  which 
animals  their  small  size  and  general  appearance  and  habits  give 
them  much  superficial  resemblance.  Cuvier's  investigations  into 
their  anatomical  structure,  and  especially  their  dental  characters, 
led  him  to  place  them  among  the  Ungulates,  near  the  genus 
fihinoceros,  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  affinities  of  this  anomalous  and  isolated 
group,  as  no  extinct  animals  possessing  their  distinctive  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 
*  !>  c  #>  P  i>  m  t-  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 


FIG.  177.— Skull  and  dentition  of  Dendrohyrax  dorsalis.     x  §. 

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 


HYRACID& 


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 
and  sacculated  caecum 
(cm)  at  the  com- 
mencement of  the 
colon,  there  is,  lower 
down,  an  additional 
pair  of  large,  conical, 
pointed,  supplemen- 
tal caeca  (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.  FlG  178_ Diagrammatic  view  of  the  alimentary  canal  of 
The  Ureters  Open  into  Hyrax  capensis,  the  intestines  being  somewhat  abbreviated, 
the  f  lindus  of  the  ^'  Duodenum  >  i>  ileum  ;  «»>  caecum ;  c,  supplemental  colic  caeca  ; 
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. 

Hyrax.1 — Molar  teeth  having  the  same  pattern   as    those  of 

1  Hermann,  Tab.  AffinU.  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  (Spicilcgia  Zoologica}.  Besides  minor 
memoirs,  two  detailed  accounts  of  its  structure  have  appeared — one  by  Brandt, 
in  Mem.  Acad.  Nat.  Sden.  St.  Petersbourg,  7Ume  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. 
Vertebrae :  C  7,  D  22,  L  8,  S  6,  C  6.  Of  this  form  the  earliest 
known  species,  H.  capensis  (Fig.  176)  is  the  type.  There  are  several 
other  species,  as  H.  liabessinicus  and  syriacus,  from  Eastern  Africa 
and  Syria.  They  inhabit  mountainous  and  rocky  regions,  and  live 
on  the  ground. 

Dendrohyrax.1 — Molar  teeth  having  the  same  pattern  as  Palceo- 
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.  Vertebrae  : 
C  7,  D  21,  L  7,  S  5,  C  10.  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.  blainvillei)  under  the  designation  of  Heterohyrax.2 

Suborder  PROBOSCIDEA. 

m 

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  venae  cavse  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  Annalcs  des  Sciences  Naturclles,  6i4me  ser.  torn.  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. 

1  Gray,  Ann.  Mag.  Nat.  Hist.  ser.  4,  vol.  i.  .p.  48  (1868). 

2  See  a  paper  by  J.  V.  Barboza  du  Bocage,  in  the  Jorn.  Sci.  Phys.  Nat.  Lisbon 
(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.  Eadius 
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  venae  cavae  entering  the 
right  auricle.  Stomach  simple.  A  capacious  caecum.  Testes  per- 
manently abdominal.  Uterus  bicornuate.  Placenta  nondeciduate 
and  zonary.  Mammae  two,  pectoral. 

With  regard  to  the  teeth,  the  incisors,1  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,  striae  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  maxillae  ;  but,  as  in  Rodents,  they  are  originally  developed  in  the 
gum  covering  the  premaxillse,  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  premaxillae 
into  the  great  and  specially  modified  alveolar  processes  of  the  maxillae,  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  Mastodon.  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 


Iff 


Fio.  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,  Ekphas  insignis;  III,  EUphas  africanus;  IV,Elephasprimigenius.  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  triturating 
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  Paleeotherium  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,  III,  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  either  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  (E.  planifrons)  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  (Fig.  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  premaxillae  and  maxillae,  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  lamellae  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 


Fio.  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  inesethmoid  (ME), 
an,  Anterior,  and  pn,  posterior  narial  aperture.  ^  natural  size.  (From  Flower's  Osteology  of 
tJie  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.  It  is  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  calcaneum,  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^E. 

Cheek-teeth  succeeding  one  another  in  an  arc  of  a  circle,  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.1 — Dentition  :  i  ^,  c  $,  dm  f ,  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.  Vertebras:  C  7,  D  19- 
21,  L  3-4,  S  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  (E.  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, 


nearly  parallel. 
Ears  of  moder- 
ate size.  Upper 
margin  of  the 
end  of  the  pro- 
boscis devel- 
oped into  a 
distinct  finger- 
like  process, 
much  longer 
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 
E.  sumatranus,  but  the  distinction  has  not  been  satisfactorily  estab- 
1  Linn.  Syst.  .Nat.  12th  ed.  vol.  i.  p.  48  (1766). 


Fio.  181. — Grinding  surface  of  a  half-worn  lower  molar  of  the  Indian 
Elephant  (Elephas  indicus).  d,  Dentine ;  e,  enamel ;  c,  cement.  (From 
Owen.) 


ELEPHANTID&  42$ 

lished.  The  appearance  of  the  Asiatic  Elephant  is  familiar  to  all. 
Though  rarely  breeding  in  captivity,  it  has  been  domesticated  from 
the  most  remote  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  of  shade  and  solitude  and  the 
neighbourhood  of  water.  The  height  of  the  male  at  the  shoulder 
when  full  grown  is  usually  from  8  to  10  feet,  but  occasionally  as 
much  as  11.  The  female  is  somewhat  smaller. 

In  the  African  Elephant  (E.  africanus)  the  molars  (Fig.  182)  are 
of  coarse  construction,  with  fewer  and  larger  plates  and  thicker 
enamel.  Ridge  -  f  ormula :  3,  6,  7,  7,  8,  10.  The  plates  not 
flattened,  but  thicker  in  the  middle  than  at  the  edges,  so  that  their 
worn  grinding  surfaces  are  lozenge-shaped.  Ears  very  large.  The 


Fio.  182. — Grinding  surface  of  a  partially  worn  right  upper  molar  of  the  African  Elephant 
(Elephas  a/ricanus).  Letters  as  in  the  preceding  figure.  The  left  side  of  the  figure  is  the  front 
of  the  tooth,  and  the  lower  side  the  outer  border.  (From  Owen.) 

upper  and  lower  margins  of  the  end  of  the  trunk  forming  two 
nearly  equal  prehensile  lips.  But  three  hoofs  on  the  hind  foot. 
This  species  now  inhabits  the  wooded  districts  of  the  whole  of 
Africa  south  of  the  Sahara,  except  where  it  has  been  driven  away 
by  human  settlements.  Fossil  remains  of  Pleistocene  age,  undis- 
tinguishable  specifically,  have  been  found  in  Algeria,  Spain,  and 
Sicily.  It  was  trained  for  war  and  show  by  the  ancient  Cartha- 
ginians and  Romans,  and  recent  experience  of  the  species  in  captivity 
in  England  shows  that  it  is  as  intelligent  as  its  Asiatic  relative,  if 
not  more  so,  while  surpassing  it  in  courage,  activity,  and  obstinacy. 
Nevertheless,  in  modern  times,  no  people  in  Africa  have  been 
sufficiently  civilised  or  enterprising  to  care  to  train  it  for  domestic 
purposes.  It  is  hunted  chiefly  for  the  sake  of  the  ivory  of  its 
immense  tusks,  of  which  it  yields  the  principal  source  of  supply  to 
the  European  market,  and  the  desire  to  obtain  which  is  rapidly 
leading  to  the  extermination  of  the  species.  In  size  the  male 
African  elephant  often  surpasses  that  of  Asia,  but  the  female  is 
usually  smaller.  The  circumference  of  the  fore  foot  is  half  the 


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 


FIG.  183. — African  Elephant  (Elephas  africanus).    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 


ELEPHANTIDjE  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  Hyaenas  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 


FIG.  184. — Restored  skeleton  of  the  Mammoth  (Elephas  primigenius).  From  Tilesius  in 
Mem.  Acad.  Imp.  Sc.  St.  Petersbmirg,  vol.  v.  (1815).  s,  Scapula  ;  h,  humerus  ;  r,  radius  ;  u,  ulna  ; 
c,  carpus  ;  rs,  ischium  ;  /,  femur  ;  t,  tibia  ;  fi,  fibula ;  ta,  tarsus. 

Hills,  and  equivalent  deposits  in  the  Punjab,  Perim  Island,1  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  E.  priinigenitis,  the 
Mammoth,1  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  in  situ,  in  the  frozen  soil  of  the  tundras  of 
Northern  Siberia,  has  for  a  long  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,  E.  indicus,  has  retained 
the  slightly  more  generalised  characters  of  the  Mammoth's  con- 
temporaries of  more  southern  climes,  E.  columbi  of  America,  and 
E.  arftieniacus  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  seen  on  the  worn  surface,  or  in  a 
horizontal  section  of  the  tooth.  Dr.  Falconer  gave  the  prevailing 

1  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- 
skiold  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. 


ELEPHANTID^E 


429 


Fio.  185. — Grinding  surface  of  upper  molar  of  the 
Mammoth  (Elephas  primigenius).  c,  Cement ;  d,  dentine  ; 
e,  enamel.  (From  Owen.) 


"ridge-formula"  as  4,  8,  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  c      <?.  p 

the  posterior  end  of  the  ^---^J^zSS^IISJfJflPi 

series,  is  subject  to  very  ^/'^^^^^^jmm  'II W  1 
great  individual  variation, 
ranging  in  each  tooth  of 
the  series  within  the  fol- 
lowing limits :  3  to  4,  6 
to  9,  9  to  12,  9  to  15, 
14  to  16,  18  to  27,  ex- 
cluding the  small  plates 
called  talons  at  each  end 
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  E.  indicus,  is  almost  imperceptible. 

The  bones  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  fossae,  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.1  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  Reliquice 
Aquitanicce,  and  in  many  other  works  bearing  on  the  subject  of  the  antiquity  of 


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  '1  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  Nordenskiold,  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, 


ELEPHANTID&  43 1 

both  among  ignorant  peasants  and  learned  academicians,  will  be 
found  in  Nordenskiold's  Voyage  of  the  Vega  (English  translation, 
vol.  i.  1881,  p.  398  sq.)  and  a  series  of  papers  in  the  Geological 
Magazine  for  1880  and  1881,  by  H.  H.  Ho  worth,  as  well  as  in  a 
separate  work  on  the  Mammoth  by  the  same  writer.  For  the 
geographical  distribution  and  anatomical  characters,  see  Falconer's 
Palceontological  Memoirs,  voL  ii.  1868 ;  Boyd  Dawkins,  "Elephas 
primigenius,  its  Eange  in  Space  and  Time,"  Quart.  Journ.  Geol.  Soc. 
xxxv.  p.  138  (1879);  and  Leith  Adams,  "Monograph  of  British 
Fossil  Elephants,"  part  ii.,  Palceontographical  Society  (1879). 

E.  antiquus,  of  the  European  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  E.  mnaidriensis  and  E.  melitensis  ;  some 
of  the  individuals  of  the  latter  not  exceeding  3  feet  in  height.  The 
European  E.  meridionalis  is  a  southern  form  of  somewhat  earlier 
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  large  dimensions,  its  height  being  estimated  at  upwards  of 
15  feet.  The  ridge -formula  is  lower  than  in  E.  antiquus,  the 
molars  are  broad,  with  the  worn  enamel-discs  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 
E.  namadicus  appears  closely  allied  to  E.  antiquus,  from  which  it  is 
distinguished  by  a  bold  ridge  across  the  forehead.  Among  the  Plio- 
cene forms  E.  hysudricus  may  be  an  ancestral  type  allied  to  the  Indian 
Elephant ;  while  E.  planifrons  is  closely  related  to  E.  meridionalis, 
although  retaining  the  ancestral  feature  of  developing  premolars. 

The  Stegodont  group  is  peculiar  to  the  eastern  parts  of  the 
Old  World,  and,  as  already  observed,  connects  the  true  Elephants 
intimately  with  the  Mastodons.  The  molars  (Fig.  179,  II)  are 
characterised  by  the  lowness  of  the  ridges,  while  the  intervening 
valleys  may  have  but  little  cement,  and  there  may  be  a  more 
or  less  distinct  longitudinal  groove  in  the  crown  dividing  each 
ridge  into  an  inner  and  an  outer  moiety.  In  species  like  E.  insignis 
the  ridge-formula  is  nearly  the  same  as  in  E.  meridionalis,  but  in 
E.  difti  some  of  the  molars  carry  only  six  ridges,  and  premolars 
were  present,  so  that  we  thus  have  such  a  complete  transition  to 
the  next  genus  that  it  is  very  difficult  to  know  where  to  draw  the 
line  between  the  two. 

Mastodon.1 — Dentition  :  i  — ^-  c  %,  dm  f ,  m  f .     Upper  incisors 

1  Cuvier,  Ann.  du  Mustum,  vol.  viii.  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  betAveen  the  ridges. 
The  third,  fourth,  and  fifth  cheek-teeth  (i.e.  the  last  milk-molar, 
and  the  first  and  second  molars)  having  the  same  number  of  ridges,1 
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  M.  angustidens,  M.  pentdid,  and  M. 
longirostris).  In  the  common  North  American  species  (M.  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  M.  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  (Palaxmt. 
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. 


ELEPHANTID^K 


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  M.  americanus  of  North  America,  no  vertical  succession  of  the 


FIG.  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  M. 
americanus)  the  ridges 
are  simple  transverse 
wedge  -  shaped  eleva- 
tions, with  straight  or 
concave  edges.  In 
other  species  the  sum- 
mits of  the  ridges  are 
more  or  less  subdivided  into  conical  cusps,  and  may  have  accessory 
cusps  clustering  around  them  (as  in  M.  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 


FIG.  187.— Oblique  side  and  crown  view  of  the  last  upper 
molar  of  Mastodon  arvernensis.    (From  Owen.) 


434 


UNGULATA 


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.  Falconer,  and  are 

FIG.  188.— Grinding  surface  of  the  partially  worn  last  three  ill  number,  name- 
left  lower  milk -molar  of  Mastodon  angustidens,  from  the  }y  the    last    milk-molar 
Upper  Miocene  of  India.     The  lower  side  of  the  figure  is  j  .-,      r- 
the  outer  border  of  the  tooth.  and  the  first  and  S6COnd 

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  Trilophodon  (of  which  an  intermediate  molar  is  shown 
in  Fig.  188),  and  the  latter  that  called  Tetralophodon  by  Dr.  Falconer. 
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  by  the 
formulae  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  Mastodon  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  M.  arnericamis,  M.  ohioticus,  and  M.  giganteus 
— survived  to  a  late  Pleistocene  period. 

The  range  in  space  will  be  best  indicated  by  the  following  list 
of  some  of  the  better  known  species.     (1)  Trilophodont  series — 
M.  angustidens,1  borsoni,  pentelici,  turicensis,  from  Europe  ;  M.  falconen 
1  Also  found  beyond  the  extreme  north-western  frontier  of  India. 


DINO  THERIID& 


435 


and  pandionis,  from  India  ;  M.  americanus,  obscurus,  and  productus, 
North  America ;  and  M.  cordillerum  and  humboldti,  South  America. 
(2)  Tetralophodont  series — M.  arvernensis,  M.  longirostris,  from 
Europe  ;  M.  latidens,  sivalensis,  and  perimensis,  from  India ;  M.  miri- 
faus,  from  North  America.  Mastodon  arvernensis  and  M.  longirostris, 
together  with  a  trilophodont  species,  occur  in  the  crag-deposits  of 
Norfolk  and  Suffolk. 


Family  DINOTHERIID^E. 

An  extinct  family  distinguished  from  the  Elephantidce  by  the  whole 
series  of  permanent  cheek-teeth  being  in  use  at  the  same  time, 

Dinotherium.1 — Dentition  of  adult :  i  &,  c  ft,  p  f,  m  f  =  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  greatly 
thickened  and  deflected  beak  or  pro- 
longation of  the  symphysis.  In  section 
they  do  not  show  the  decussating  striae 
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 
of  the  permanent  molar  series :  2,  2,  3, 
2,  2.  The  three  ridges  of  the  first  true 
molar  are  constant  in  both  upper  and  *>  2>  3> 
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). 


Fie.  189.— Skull  of  Dinotherium 
gigaiiteum,  from  the  Lower  Pliocene 
of  Eppelsheim,  Hessen- Darmstadt. 
(After  Kaup.)  p,  3,  4,  premolars ; 


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  Dinothenum  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. 

Subvrder  AMBLYPODA. 

Uintatherium.1 — 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  Avere  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 

1  Leidy,  Proc.  Ac.  Nat.  Sci.  Philad.  1872,  p.  169. 


AMBLYPODA 


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  Avas  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 
*  §>  c  T>  P  I'  m  f  =  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  Uintatherium.  mirabile.    ^  natural  size.    (From  Marsh, 
Am.  Journ.  Set.  vol.  xii.  pi.  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  described  by  Leidy  in  1872,  under  the 
name  of  Uintatherium  (from  the  Uinta  mountains,  near  which  they 
were  found).  Subsequently  the  names  Dinoceras,  Tinoceras,  Loxo- 
lophodon,  etc.,  have  been  applied  to  various  members  of  the  group, 
but  the  characters  by  which  they  are  distinguished  do  not  seem  of 
sufficient  importance  to  allow  of  their  separation  from  the  type 
genus  Uintatherium.1 

Coryphodon.2 — Another  interesting  form  referred  to  this  suborder 
is  Coryphodon,  which  appears  to  connect  the  Uintatheriidce  with  the 
most  primitive  Perissodactyla.  It  was  first  described  by  Owen  in 

1  For  detailed  descriptions  and  figures  of  this  group,  see  Marsh,  ' '  Monograph 
of  the  Dinocerata,"  Rep.  U.S.  Geol.  Surv.  vol.  x.  (1884). 
-  Owen,  Brit.  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. 


FIG.  191.— Palatal  aspect  of  the  cranium  of  Coryphodon  hamatus,  from  the  Wasatch  Eocene  of 
New  Mexico.   f  natural  size.    (After  Cope.) 

The  genus  should  be  regarded  as  the  type  of  a  distinct  family 
Coryphodontidce. 

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  palaeontologists  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.  Periptychus,  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  Phenacodiis,  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- 
therium  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  Hyracothenum ;  all  the  feet  had 
five  fully  developed  digits,  and  the  tail,  was  long.  Meniscotherinm 
and  Hyracodontotherium  are  more  specialised  forms  of  somewhat 
later  age,  with  a  lophodont  dentition;  the  latter  genus  being 
European. 

Suborder  TOXODONTIA. 

In  addition  to  the  Macraucheniidce  and  certain  other  forms 
noticed  under  the  head  of  the  Perissodactyla,  the  Tertiaries  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  Toxodan. — 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- 
iheriwn  (p.  412)  and  the  following  genus  Toxodon.  The  typical 
species  of  Nesodon  was  about  as  large  as  a  Sheep,  but  nothing 
more  is  known  of  it  than  the  teeth  and  portions  of  the  skull. 

Toxodon  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  i  |,  c  %,p  ^,  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  Protoxodon  and  Adinothemim  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. 


Fio.  192. — Cranium  and  Lower  Jaw  of  Typotherium  crtetatum.    J  natural  size.     From  Gervais. 

Typotherium.  —  Typotherium  (Fig.  192),  also  called  Mcsotherium, 
from  the  same  locality  as  Toxodon,  was  an  animal  rather  larger  than 


TILLODOXTIA  441 


a  Capybara,  and  of  much  the  same  general  appearance.  Its  skeleton 
is  completely  known,  and  shows  a  singular  combination  of  characters, 
resembling  Toxodon  or  a  generalised  Ungulate  on  the  one  hand,  and 
the  Rodents,  especially  the  Leporidce,  on  the  other.  In  the  presence 
of  clavicles  it  differs  from  all  known  Ungulates,  and  in  having  two 
pairs  of  lower  incisors  from  all  Rodents.  The  teeth  are  i  4-,  c  %,p  |, 
m  f  =  24. 

From  the  Tertiaries  of  various  parts  of  South  America  a  number 
of  forms  more  or  less  closely  allied  to  Toxodon  and  Typotherium  have 
been  recently  described,  but  as  many  of  them  are  very  imperfectly 
known,  and  there  is  much  doubt  as  to  their  generic  position,  it  will 
be  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  curious 
chain  of  affinities,  which  would  seem  to  unite  the  Ungulates  on  the 
one  hand  with  the  Rodents  on  the  other ;  but  further  materials 
are  required  before  we  can  establish  with  certainty  so  important  a 
relationship,  one  which,  if  true,  would  alter  materially  some  of  the 
prevailing  views  upon  the  classification  of  mammals. 


Group  TILLODONTIA. 

Here  may  be  noticed  a  remarkable  group  of  animals,  called  by 
Marsh,  Tillodontia,  the  remains  of  which  are  found  abundantly  in 


FIG.  193.—  Skull  of  Tillotheriumfodiens.     |  natural  size.     From  .Marsh. 

the  Lower  and  Middle  Eocene  beds  of  North  America.  They  seem 
to  combine  the  characters  of  the  Ungulata,  Rodentia,  and  Carnivora. 
In  the  genus  Tillotherium  of  Marsh  (probably  identical  with  the  pre- 
viously described  A nchippodus  of  Leidy)  the  skull  (Fig.  193)  resembled 
that  of  the  Bears,  but  the  molar  teeth  were  of  the  Ungulate  type, 
while  the  large  incisors  were  very  similar  to  those  of  the  Rodents. 
The  dental  formula  is  i  •§• ,  c  -},  p  f ,  m  f .  The  first  pair  of  incisors 


442  UNGULATA 


was  very  small ;  the  upper  molars  were  tritubercular,  while  the 
lower  ones  had  crescentoid  ridges  as  in  PaUeothenwn.  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  Tseni- 
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  Eodent 
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  Ungulata. — In  addition  to  the  works  and  memoirs  mentioned 
under  the  different  sections  of  the  order,  the  following  may  be  referred  to : — 
"\V.  Kowalevsky,  "Monographic  des  genus  Anthracotherium,"  Palceontographica 
1873;  Id.  "Sur  1'Anchitherium  aurelianense  et  sur  1'histoire  paleontologique 
des  Chevaux,"  Mim.  de  I'Acad.  Imp.  des  Sciences  de  St.  Pttersbourg,  1873  ;  Id. 
"On  the  Osteology  of  the  Hyopotamidre, "  Philosophical  Transactions,  1873; 
L.  Riitimeyer,  "Versuch  einer  natiirlichen  Geschichte  des  Rindes,"  etc.,  Neue 
Denks.  der  allgem.  Schweiz.  Gesellsch.  fur  Naturwissenschaften,  1867;  Id.  "Die 
Kinder  der  Tertiar-Epoche, "  Abhand.  der  Schweiz.  Paldont.  Gesellsch.  1877  and 
1878  ;  Id.  "Beitrage  zu  einer  Natiirliche  Geschichte  der  Hirsche,"  ibid.  1880-1881 ; 
C.  J.  Forsyth-Major,  "Beitrage  zur  Geschichte  der  Fossilen  Pferde,"  ibid.  1880  ; 
M.  Schlosser,  "Beitrage  zur  Kenntniss  der  Stammesgeschichte  der  Hufthiere 
und  Versuch  einer  Systematik  der  Paar-und  Unpaarhufer, "  Morph.  Jahrb.  1886  ; 
E.  D.  Cope,  "The  Perissodactyla,"  Amer.  Natural.  1887;  M.  Pavlow,  "Etudes 
sur  1'histoire  paleontologique  des  Ongules,"  Bull.  Soc.  Imp.  Naturalistcs  Moscoic, 
1887-1890.  W.  B.  Scott  and  H.  F.  Osborn,  "The  Mammalia  of  the  Uinta  For- 
mation," Trans.  Amer.  Phil.  Soc.  vol.  xvi.  (1889). 


CHAPTEE  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  ^,  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  imperfect  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  fossae ;  the  condyle  of  the 
mandible  is  elongated  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  Myoxidce)  has  a  large  caecum ;  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  include  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- 


444  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  structural  importance.  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- 
matic arch  varies  in  its 

FIG.  194.— Skull  of  Hystrix  cristata  (juv.)    t,  Temporal   degree     of     development, 
injiscle  ;  m,  inasseter ;  m.',  portion  of  masseter  transmitted   ^nd    ^Jjg    position    of    the 
through  the  infraorbital  foramen,  the  superior  maxillary    .         ,     , 
nerve  passing  outwards  between  it  and  the  maxillary  bone.    Juga^  therein  IS  USCd  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  (Bathyergince),  it  is  extremely  narrow 
transversely,  its  width  being  less  than  that  of  one  of  the  molar  teeth. 
Auditory  bull*  are  always  present,  and  generally  large ;  in  some 
genera,  as  in  the  Gerbilles  and  Jerboas,  there  are  also  supplemental 


RODENTIA  445 


mastoid  bullse  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  *f»6$»j>-f»fli£  (total  28) 
in  the  Duplicidentata  to  i  ^,  c  %,  p  #,  m  |-  (total  12)  in  Hydromys, 
Xeromys,  and  one  species  of  HeterocepJuilus ;  but  in  the  great 
majority  of  forms  it  is  very  constant,  i  y,  c  §,  p  ^  -^L  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         FIG.  195.— Vertical  aud  longitudinal  section  through  skull  of 

circle    Or   more  has  *^e  •^eaver  (Castor  fiber)  showing  the  cerebral  cavity,  the  greatly 

,        ,  developed  turbinal  lamellse,  the  mode  of  implantation  of  the  large 

been     lOrmeci,     tne  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  (Dasyproctidce),  the  milk-teeth  are  long  retained, 
Avhile  in  the  allied  Cavies  (Caviidce)  they  are  shed  before  birth. 

There  are  generally  nineteen  dorso-lumbar  vertebrae  (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  vertebra?  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  Dipodidce  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  (Geomyidcv)  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 
papillae  at  the  base ;  and  the  apical  portion  is  generally  covered 
with  small  filiform  papillae,  some  of  which  in  the  Porcupines 
(Hystrix}  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  amphibius)  and  the  Agouti  (Dasypi-oda.  aguti)  it  is 
strongly  constricted  between  the  oesophagus  and  pylorus.  In  the 
common  Dormouse  the  oesophagus  immediately  before  entering  the 


RODENTIA 


447 


in    some    other    species,    as   in 
Beaver,    glandular   masses   are 


stomach  is  much  dilated,  forming  a  large  egg-shaped  sac  with 
thickened  glandular  walls ;  and 
Lophiomys  imhausi  and  in  the 
attached  to  and  open  into  the 
cardiac  or  pyloric  pouches.  The 
alimentary  canal  (Fig.  196)  of 
all  Rodents,  with  the  exception 
of  the  Dormice  (Myoxidce),  has 
a  caecum,  which  is  often  of  great 
length  and  sacculated,  as  in  the 
Hares,  Water- Voles,  and  Porcu- 
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 


Fio.  196.—  Alimentary  canal  of  Rat  (Mus  decu- 

penis   (which    is    generally  pro-  ™a'"'s)'  thf!  »fater ,  >?rt  °f  the  T"  i!:testine 

.-,•..  i          i  ••  being  omitted,    o,  (Esophagus ;  d,  duodenum ; 

vided  with  a  bone)  can  be  more  it  neum  •  cm,  csecum ;  c,  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,  vesiculae  seminales  are  present  in  all.  The  uterus 
may  be  double,  each  division  opening  by  a  separate  aperture  into 
a  common  vagina,  as  in  Leporidce,  Sciuridce,  and  Hydrochoerus,  or 
completely  two-horned,  as  in  most  species.  The  mammae  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  Ododontidce  the  mammae  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  Mus,  Arvicola,  Cricetus,  etc.,  or  into  the  rectum,  as  in 
Arctomys  and  Aulacodus,  or  into  the  passage  common  to  both,  as  in 


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- 
Eats  and  Porcupines ;  in  several  genera,  as  in  Xerus,  Acanthorniis, 
Platacanthomys,  Ecliinothrix,  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  Porcupines  and  Hares  the  tendons 
of  the  flexor  digitorum  longus  and  flexor  hallucis  longus  are  con- 
nected in  the  foot,  while  in  the  Eats  and  Squirrels  they  are  separate, 
and  the  flexor  digitorum  longus  is  generally  inserted  into  the 
metatarsal  of  the  hallux.1 

Eodents  are  tolerably  well  represented  in  a  fossil  condition  from 
the  period  of  the  Upper  Eocene,  while  if  Decticadapis,  of  the  Lower 
Eocene  of  Eheims,  is  rightly  referred  to  it  the  order  dates  from  the 
oldest  Tertiary.  All  the  fossil  forms  at  present  knoAvn  are,  however, 
essentially  true  Eodents,  and  afford  no  clue  as  to  the  relations  of 
the  order  with  other  mammals.  The  remote  affinities  of  the 
Eodents  to  the  Proboscidea,  as  well  as  their  more  marked  resem- 
blances to  Typother'mm,  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  calcaneum.  Testes  abdominal,  and 
descending  periodically  only  into  a  temporary  sessile  scrotum. 

Section  SCIUROMORPHA. 

Zygomatic  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.  Anat.  Phys.  vol.  xvii. 


ANOMALURID& 


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^E. 

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 


Pro.  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  ^ ;  molars  not  tuberculate,  with 
transverse  enamel-folds.  Confined  to  the  Ethiopian  region. 

Anomalurus,1  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 ;  A.  orientalis  occurs 
near  Zanzibar,  and  A.  pusillus  is  from  the  equatorial  regions  of  that 

1  Waterhouse,  Proc.  Zool.  Soc.  1842,  p.  124. 
29 


450 


RODENT1A 


continent.  According  to  Mr.  0.  Thomas,1  the  latter  "  little  animal 
is  most  nearly  allied  to  the  West-African  A,  beecrofti,  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 


FIG.  198.— Lateral  view  of  skull  of  American  Mai-mot 
(Arctomys  monca). 


Arboreal  or  terrestrial  forms,  with  cylindrical  hairy  tails,  with- 
out scales,  and  with 
twelve    or    thirteen 
Skull 


or 

pairs  of  ribs. 
(Figs.  198, 199)  with 

distinct  postorbital 
processes;  infra- 
orbital  opening 
small;  palate  broad; 
p  f ;  first  upper  pre- 
molar  very  small  or 
deciduous ;  molars 
rooted,  tubercular. 

Subfamily  Seiur- 
inae. — Incisors  com- 
pressed ;  form  slen- 
der ;  tail  long  and 

hairy.     Cosmopolitan  (excluding  Australian  region). 

This    subfamily  includes   the  true    Squirrels,  of  which  seven 

existing  genera  are  usually  recognised. 

Sciurus.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.    Mammae, 

from  four  to  six.   Skull 

(Fig.  199)  lightly  built, 

with    long    postorbital 

processes.  Penultimate 

upper  premolar,  when 

present,  minute. 

1  Proc.  Zool.  Soc.  1882,  p.  8.          "  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  86  (1766). 


Fio.  199. — Palatal  Aspect  of  cranium  of  Squirrel  (Sciurus 
bicolor).    Natural  size. 


SCIURID&  45i 

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 


Pio.  200. — Burmese  Squirrel  (Sclurus  pygerythnis).    After  Anderson. 

English  Squirrel  (S.  vulgaris)  is  found  over  the  whole  of  the  Palsearctic 
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.  palmarum),  which  in  large  numbers  runs 
about  every  Indian  village.  Another  Oriental  species  (S.  canicep*) 
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  Palsearctic,  15  to  the  Ethiopian,  about  40  to  the 
Oriental,  and  1 6  to  the  combined  Nearctic  and  Neotropical  regions  " 
(Thomas). 

Fossil  species  referred  to  Sciurus  are  found  in  the  European 
Tertiaries  down  to  the  Phosphorites  of  Central  France,  while  others 
occur  in  the  White  Eiver  Miocene  of  the  United  States. 

fihithrosciurus.1 — A  very  striking  Squirrel,  confined  to  Borneo, 
and  as  yet  only  known  from  three  or  four  examples,  has  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  i,  and  its  molars  are  simpler  and  less  ridged 
than  in  the  other  genera.  This  Squirrel  (B.  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  4=- ; 
molars  as  in  Sciurus.  Mammae  two.  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). 

2  Hemprich  and  Ehrenberg,  Symbol.  Phys.  Mamm.  vol.  i.  (1832). 

3  Illiger,  Prodromus  Syst.  Mamm.  p.  83  (1811). 


SCIURID&  453 

true  Squirrels,  from  which  they  differ  in  several  unimportant 
details.  There  is  only  one  functional  premolar — the  small  anterior 
one  usually  found  in  Sciurus  being  either  absent  altogether  or  quite 
small  and  functionless.  There  are  some  four  well-defined  species, 
all  found  in  North  America,  one  (T.  asiaticus)  extending  also  through 
Siberia  into  Eastern  Europe.1  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  Spermophihts, 
so  that  the  division  of  the  Sciuridce  into  two  subfamilies,  although 
convenient  for  classification,  is  rather  artificial. 

Remains  of  Tamias,  probably  belonging  to  existing  species,  occur 
in  the  Pleistocene  deposits  of  Europe  and  Nebraska. 

Pteromys2  and  Sciuropterus.3 — 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  Sciuropterus  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,  Lemons  d' Anatomic  Comp.  (1800). 

3  Cuvier,  Ann.  du  Museum,  vol.  x.  p.  126  (1825). 


454  RODEXTIA 


natives  of  the  Oriental  region,  except  that  one  of  SctNFQfttnu  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  Sciuridce  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. 

Extinct  Genera. — The  genera  Pseudosciurus  and  Sciuroides,  from 
the  Upper  Eocene  of  Europe,  have  the  molar  teeth  more  elongated 
than  in  Sciurus.  Gymnoptychus  with  p  ^,  from  the  North  American 
Miocene,  approximates  in  the  structure  of  its  molars  to  Tamias. 
Meniscomys  (p  f ),  from  the  latter  deposits,  together  with  Sciurodon 
of  the  French  Phosphorites,  are  regarded  as  Squirrels  showing  signs 
of  affinity  with  the  Haplodontidce. 

Subfamily  Aretomyinse. — 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  Palse- 
arctic  and  Nearctic  regions. 

Arctomys.2 — External  form  stout  and  heavy,  ears  short,  tail 
short  and  hairy,  cheek-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.  A.  bobac  occurs  in  Eastern  Europe 
and  Siberia.  Several  species  (e.g.  A.  monax,  Fig.  198)  are  found 
in  the  Nearctic  region,  and  many  in  Kashmir  and  Central  Asia. 
The  long-tailed  Red  Marmot  (A.  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, 

1  0.  Thomas,  Journ.  As.  Soc.  Bengal,  vol.  ML  p.  256  (1888). 
2  Schreber,  Sdugethiere,  vol.  iv.  p.  721  (1792). 


SCIURID& 


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 


fe^Bev 

.     — Vr  -'- 

Fio.  201.— Alpine  Marmot  (Arctomys  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,  Amur.  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, 

FIG.  202.— Palatal  aspect  of  the  cranium  of  the  Prairie  Marmot    as  they  are  of  ten  Called 
(Cynomys  ludovicianus).  ,,  TJ      •   •       -r%          j> 

"  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  cunicularia)  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.1 — 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  Arctomys,  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  (S.  citillus) : 
1  F.  Cuvier,  Mem.  du  Museum,  vol.  vi.  p.  293  (1822). 


CASTORJD^E  457 

"  It  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  Eussian  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  Ardomys 
and  Sciurus,  but  with  tritubercular  upper  molars  and  no  postorbital 
processes  to  the  skull ;  in  the  latter  respect  agreeing  Avith  the  next 
family.  In  the  size  of  the  preorbital  vacuity  the  skull  resembles  the 
Hystricomorpha. 

Family  HAPLODONTID.E. 

Distinguished  from  the  Sciuridce  by  the  absence  of  postorbital 
processes  to  the  frontals,  the  depressed  skull,  and  the  rootless  cheek- 
teeth. Premolars  f ;  the  penultimate  upper  one  small. 

Haplodon.1 — H.  rufus  and  H.  major,  of  North  America,  west  of 
the  Rocky  Mountains,  are  the  only  representatives  of  the  family; 
their  habits  are  similar  to  those  of  Cynomys. 

Family  CASTORID^E. 

Skull  massive,  without  postorbital  processes,  the  angle  of  the 
mandible  rounded,  and  the  cheek-teeth  rootless,  with  re-entering 
enamel-folds.  Premolars  y.  Habits  natatorial. 

Castor? — 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  O3sophageal  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,  Palaearctic  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  Eussia  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  Eucastor. 

Extinct  Genera. — A  very  large  Beaver  known  as  Trogontherium 
(Didbroticus),  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. 


MYOXIDsE 


459 


Section  MYOMORPHA. 

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  Bathyerginop,  rising 
from  the  inferior  sur- 
face of  the  incisive 
alveolus.  Clavicles 
well  developed,  except 
in  Lophiomys.  Tibia 
and  fibula  united. 

FIG.  203.— Side  view  of  skull  of  Fiber  zibethicus,  natural  size. 


Family  MYOXID/E. 

Small  arboreal  forms,  with  long  hairy  tails,  large  eyes  and  ears, 
and  short  fore  limbs.  No  caecum  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  i ;  molars  rooted,  with  transverse  enamel-folds. 

The  Dormice  form  a  natural  family  allied  to  the  Squirrels  in 
form  and  habits,  and  confined  to  the  Palaearctic  and  Ethiopian 
regions.  The  absence  of  the  caecum  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.1 

Myoxus.2 — Represented  by  the  European  M.  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? — Tail  tufted  and  distichous ;  stomach  simple  ;  and 
the  molars  small,  with  concave  crowns  and  indistinct  enamel-folds. 
Some  seven  species,  Ethiopian  and  Palaearctic. 

Gmphiurus.4 — Tail  short,  cylindrical,  and  tufted  at  the  end ; 

1  For  a  monograph  of  the  Myoxidce,  see  C.  L.  Reuvens,  Die  Myoxidcc,  etc., 
4to,  Leyden,  1890.  2  Schreber,  Saugethiere,  vol.  iv.  p.  824  (1792). 

3  Wagner,  Abh.  baier.  Akad.  vol.  iii.  p.  179  (1843). 

4  F.  Cuvier,  Mammiferes,  60me  livr.  (1845). 


460 


RODENTIA 


molars  very  small,  with  the  enamel -folds  almost  absent.  Some 
three  Ethiopian  species. 

Claviglis.1 — 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  (M.  avellanarius) 
of  Europe,  distinguished  by  the  cylindrical  bushy  tail,  and  thickened 
glandular  walls  of  the  cardiac  extremity  of  the  oesophagus ;  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  LOPHIOMYID^:. 
The    genus  Lophiomys,3  represented    only  by  L.  imhausi  (Fig. 


FIG.  204. — Lophiomys  imhausi.    From  Milne-Edwards. 

204)  of  North-East  Africa,  differs  from  the  typical  Muridce  in 
having  the  temporal  fossae  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 

1  Jentink,  Notes  Leyd.  Mus.  vol.  x.  p.  41  (1888). 

2  Kaup,  Enturickl.  Europ.  Thierwelt,  p.  139  (1829). 

3  A.  Milne-Edwards,  L'Institiit,  vol.  xxxv.  p.  46  (1867). 


MURID^E  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 

Skull  (Fig.  203)  with  contracted  f rentals ;  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, ttiberculate,  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  Hydromyinse. — Molars  f  in  number,  rooted,  and 
divided  into  transverse  lobes.  Represented  by  two  Australasian 
genera. 

Hydromys2 — 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  Avail  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.3 

Xeromys.* — External  form  Murine.  Tip  of  muzzle  as  in  Mm, 
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  Plataeanthomyinse. — Molars  rooted,  with  transverse 

1  Sminthus  is  referred  to  the  Dipodidce. 
"  Geoffrey,  Ann.  du  Mustum,  vol.  vi.  p.  81  (1805). 

3  For  the  anatomy  of  this  animal  see  B.  C.  A.  Windle,  Proc.  Zool.  Soc.  1887, 
p.  53.  *  0.  Thomas,  Proc.  Zool.  Soc.  1889,  p.  247. 


462  RODENTIA 


laminae.      Flattened  spines  mingled  with  the   hair ;    tail    thickly 

haired.     Represented  by  one  genus. 

Platacanthomys.1 — 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  Gerbillinae. — Incisors  narrow;  molars  with  transverse 

laminae   (Fig.    205).     Auditory  bullae  very  large   in   most   cases. 

Hind  limbs  elongated.     Tail  usually  long  and  hairy.     Ranges  over 

the  Palaearctic,  Oriental,  and  Ethiopian  regions. 

Gerbillus.'2 — Upper  incisors  grooved;  first  molar  with  three 
laminae,  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. 

205) 

"  P^yuramys .'-The  African  genus  Pachy- 
view  of  the  molars,  from  a  uromys  is  distinguished  by  the  very  large  size 
cavern  deposit  in  Madras.  of  ^e  auditory  bulla,  as  well  as  by  the  short 

(From    the    Pakeontologia  j     «      i.        A    -T         v  v     •          I-LT.I          mi 

Jn<iica.)  anc*  neshy  tail,  which  is  club-shaped.  The 

incisors  are  narrow  and  faintly  grooved. 

Mystromys*  Otomys,5  and  Dasymys.6 — 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  Phlseomyinse.8 — This  subfamily  is  represented  only 
by  Phlceomys  9  cumingi,  of  the  Philippine  Islands,  in  which  the  incisors 
are  very  broad,  the  molars  are  divided  into  transverse  laminae,  and 
the  claws  are  large.  The  muzzle  is  blunt ;  the  ears  are  hairy 

1  Blyth,  Proc.  As.  Soc.  Bengal,  vol.  xxviii.  p.  289  (1859). 
2  Desmarest,  Nouv.  Diet.  cVHist.  Nat.  vol.  xxiv.  p.  22  (1804). 

3  Lataste,  Le  Nat.  vol.  i.  p.  314  (1880). 

4  Wagner,  Wiegmanris  Archiv,  1841,  p.  132. 

5  F.  Cuvier,  Dents  des  Mammiferes,  p.  168  (1825). 

6  Peters,  Monatsber.  Ak.  Berlin,  1875,  p.  12. 

7  A.  Milne-Edwards,  Bull.  Soc.  Philom.  ser.  6,  vol.  xi.  p.  9  (1877). 

8  Nesocia  was  included  by  Alston  in  this  subfamily. 

9  Waterhouse,  Proc.  Zool.  Soc.  1839,  p.  108. 


MURID^E  463 

externally ;  the  tail  is  moderate,  and  thickly  haired ;  and  the 
auditory  bullae  are  very  small.  The  first  upper  molar  has  three, 
and  the  others  two  laminae. 

Subfamily  Dendromyinse. — Incisors  convex  in  front ;  molars  £, 
rooted  and  tuberculated.  Ears  hairy ;  claws  long.  Confined  to 
the  Ethiopian  region. 

Dendromys.1 — 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 2  and  Lophuromys 3  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  Crieetinse. — Molars  3,  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  Murince  have  been 
evolved. 

CricettU.* — According  to  the  arrangement  pro- 
posed by  Mr.  0.  Thomas  5  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.  FIG.  206.— Left  upper 
The  first  molar  (Fig.  206,  .B)  generally  has  six  molars  of  J/IM  (^)  and 
tubercles.  The  tail  may  be  very  short. 

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  vertebrae.  The  tail  is  remarkable  for  its  shortness.  The 
best-known  species  is  C.  frumentarius,  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.  Rhipidomys,  including  Dormouse -like 
forms  with  long  tails  and  a  dentition  like  that  of  the  typical 
group ;  Oryzomys,  represented  by  Murine  species ;  Calomys,  with 
short  tail  and  Hamster-like  body ;  Vesperimus,  with  only  five  tuber- 
cles on  the  first  molar ;  Onychomys,  in  which  the  tail  is  extremely 

1  Andrew  Smith,  S.  African  Qiiart.  Journ.  vol.  ii.  p.  158  (1834). 
2  Peters,  Reisc  n.  Mossambiquc,  vol.  i.  p.  162  (1852). 

3  Peters,  Monatsber.  Ak.  Berlin,  1874,  p.  234. 

4  Cuvier,  Regne  Animal,  vol.  i.  p.  198  (1817). 

5  Proc.  Zool.  Soc.  1888,  p.  133. 


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 ;  Habrothrix,  an  Arvicoline  group,  with  a  short  and  thinly  haired 
tail ;  and  Oxymydems,  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.  Thomas1  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  Ehipidomys,  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  Cncetus  are  abundant  in  the  Pleistocene  cavern- 
deposits  of  Brazil,  where  a  numbel  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. 

Holochilus2  (Nedomys). — The  Rats  of  this  genus  are  allied  to 
the  American  forms  of  Cncetus,  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 3  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. 

Rhithrodonf  and  Ochetodon.5 — 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? — A  peculiar  North  American  genus,  in  which  the 
teeth  simulate  the  prismatic  appearance  of  those  of  the  Arvicolince. 
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 

Rat-like  tails. 

1  Proc.  Zool.  Soe.  1884,  p.  451. 

2  Brandt,  Mtm.  Acad.  Imp.  St.  Pttersbourg,  ser.  3,  vol.  iii.  p.  428  (1835). 

3  Say  and  Ord,  Journ.  Acad.  Philad.  vol.  iv.  p.  352  (1825). 

4  Waterhouse,  Proc.  Zool.  Soc.  1837,  p.  29. 

8  Coues,  Proc.  Acad.  Philad.  1874,  p.  184. 

6  Say  and  Ord,  Journ.  Acad.  Philad  vol  iv.  p.  346  (1825). 


MURIDsE 


465 


Fossil  remains  of  Neotoma  from  cavern-deposits  in  Pennsylvania 
are  not  improbably  referable  to  the  existing  Florida  Rat  (N. 
floridana).  Paciculus,  from  the  Miocene  of  the  United  States,  is 
regarded  as  an  allied  extinct  genus  with  enamel-folds  to  the  molars. 

Hypogeomys.1 — 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. 

JBrachytarsomys.3 — Represented  only  by  B.  albicauda,  a  pretty 
velvety-haired  fawn-coloured  Rat,  with  short  feet  and  a  long  tail. 

Hallomys* — 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  Arvieolinse. — 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 
Palaearctic  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 
of  molar  dentition.  The 
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.  ser.  6,  vol.  xx.  art.  1,  bis,  p.  1  (1886). 

30 


FIG.  207.— Upper  (A)  and  lower  (B)  molars  of  the 
Water- Vole  (Arvicola  amphibius). 


466  RODENTIA 


Phenacomys.1 — 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. 

Arvicola.2 — 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.  0.  Thomas  : — 

The  common  Water- Vole  (A.  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  (A.  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). 

2  Lacepede,  M6m.  de  I'Institut,  vol.  iii.  p.  495  (1801).     Many  writers  employ 
the  earlier  name  Microtus  for  the  true  Voles. 


MURID&  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  (A.  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  Moray  shire,  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  A.  gapperi  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.1 — 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  (M.  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- 


Fio.  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  Tromso-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 


MURID^E  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  RODENT! A 


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.1 — 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.2 — 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  1 2  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- 

1  Wagler,  Isis,  1832,  p.  1220. 
2  Cuvier,  lAyms  d' Anatomic  Compar.  tab.  1  (1800). 


MURID& 


47i 


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 


FIG.  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. 

Neoftber.1 — This  genus,  while  agreeing  with  Fiber  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  N.  alleni,  commonly  known  as 
the  Round-tailed  Musk-Eat,  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  Siphneinse.  —  Includes  two  genera  of  Mole  -  like 
Rodents  with  an  Arvicoline  dentition,  but  with  the  body  thoroughly 


FIG.  210.— Stpftwews  armandi.    (From  Milne-Edwards.) 

adapted  for  a  subterranean  life,  the  limbs  and  tail  being  very  short, 
and  the  external  ears  rudimentary.  Both  are  Palaearctic. 

Ellobius? — The  Russian  E.  talpinus,  the  typical  representative 
of  the  genus,  has  short  claws,  and  comes  nearest  to  the  Arvicolince. 
E.  fuscocapillus  is  from  Afghanistan. 

Siphneus.3 — 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). 

2  Fischer,  Zoognosia,  vol.  iii.  p.  72  (1814). 
3  Brants,  Het.  Oeslact  der  Muizen,  p.  20  (1827). 


MURID^E 


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  Cricetince  and  that  of 
the  Murince. 

Deomys.1 — Externally  as  in  Mus.  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,  D.  ferrugineus,  from  the  Lower  Congo,  an  animal  about  the 
size  of  the  Common  Mouse. 

Subfamily  Murinse. — Molars  rooted  and  tuberculated  ;  those 
of  the  upper  jaw  with  three  longitudinal  rows  of  tubercles  (Fig. 
206,  A). 

This  group  includes  the  true  Rats  and  Mice,  and  may  be 
regarded  as  more 
specialised  than 
the  Cricetince. 
All  the  members 
of  the  group 
closely  resemble 
one  another,  and 
are  light  and 
active,  with  large 
ears,  bright  eyes, 
and  long  and 
scaly  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. 

Mus.2 — Incisors 
narrow,    without 

grooves.     Structure  of  molars  as  in  Fig.  206,  A  (p.  463).     Incisive 
foramina  of  skull  long ;  coronoid  process  of  mandible  well  developed. 

1  0.  Thomas,  Proc.  Zool.  Soc.  1888,  p.  130. 
2  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  79  (1766). 


Fio.  211.— The  Australian  Brown-footed  Rat  (Mus  fitscipes). 
After  Gould. 


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.  Vertebrae :  C  7,  D  1 3,  L  6,  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  M.  decumanus 
or  smaller  than  M.  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 Acanthomys  gaudryi  of  the  Lower  Pliocene  Pikermi  beds  of  Attica) 
occur  in  the  Pleistocene  of  Europe. 

The  Brown  or  Norway  Rat  (M.  decumanus)  is  a  heavily  built 
A  animal,  growing  to  8  or  9 

inches  in  length,  with  a 
bluff  rounded  head,  small 
ears  (Fig.  212,  A},  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 
and  proportions  form  a 
ready  means  of  distinguish- 
ing the  two.  The  Brown 

Rat  is  believed  to  be  a  native  of  Western  China,  where  a  race 
(M.  humiliatus)  has  been  discovered  so  like  it  as  to  be  practically 
indistinguishable.  Both  this,  and  the  next  species  agree  in  their 


Fio.  212. — A,  Head  of  Brown  Rat  (M.  decumanus). 
B,  Head  of  Black  Rat  (Mus  rattus). 


MURID&  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  Eat  (M.  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  M. 
alexandrinus  the  general  colour  is  gray  or  rufous,  and  the  belly 
white.  The  disposition  of  the  Black  Rat  is  milder  than  that  of 
M.  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  M.  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  M.  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  2£  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  Phlce&mys,  but  Mr.  0.  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.  scullyi  is  found  in  Turkestan.  The  great  Indian 
Bandicoot-Rat  (N.  bandicota)  is  the  largest  representative  of  the 
subfamily,  often  exceeding  a  foot  in  length.  N.  bengalensis  is 
remarkable  for  possessing  no  less  than  eighteen  mammse.  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  Mm,  but  Avith  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  from  Eastern 
Africa. 

Uromys.2 — Differs  from  Mus  in  having  the  scales  of  the  tail  not 
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.3 — Externally  like  Mus,  but  with  the  terminal 
portion  of  the  tail  without  scales  above,  quite  naked,  transversely 
wrinkled,  and  prehensile.  Scales  of  remainder  of  tail  more  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.4 — 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  Jerboas,  whose  place  they  seem  to  take  in 
the  sandy  Australian  deserts.  Remains  of  H.  albipes  occur  in  the 
Pleistocene  of  New  South  Wales. 

Mastacomys.5 — Like  Mus,  bufc  with  the  molars  remarkably 
broadened,  and  with  only  four  mammse.  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.6 — 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, 

1  Gray,  Charlesworth's  Mag.  Nat.  Hist.  vol.  i.  p.  586  (1837).     Syn.  Pelomys, 
Peters  (1852). 

2  Peters,  Monatsber.  Ak.  Berlin,  1867,  p.  343. 

3  O.  Thomas,  Proc.  Zool  Soc.  1888,  p.  237. 

4  Liechtenstein,  Darst.  neu.  Saugethicre,  pt.  iv»  pi.  29  (1829). 

5  0.  Thomas,  Ann.  Mag.  Nat.  Hist.  ser.  5,  vol.  ix.  p.  413  (1882). 

6  Geoffroy,  Ann.  Sci.  Nat.  ser.  2,  vol.  x.  p.  126  (1840).     Acomys. 


SPALACID&  477 

Palestine,  and  Eastern  Africa  as  far  south  as  Mozambique.  A. 
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  (A.  gaudryi) 
referred  to  this  genus  occurs  in  the  Lower  Pliocene  of  Attica. 

Echinothrix.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  E.  leucura,  an 
animal  of  about  the  size  of  the  Brown  Rat,  with  its  fur  thickly 
mixed  with  spines.  It  is  found  in  Celebes. 

Typhlomys. 2 — 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. 

Cricetomys3  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  SPALACID^E. 

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  Spalaeince. — Angular  part  of  the  mandible  arising 
from  the  lower  edge  of  the  socket  of  the  lower  incisor.  No  pre- 
molars. 

Spalax.5 — Represented  by  the  great  Mole -Rat  (S.  typhlus)  of 
South-Eastern  Europe,  in  which  the  eyes  are  completely  covered  by 
the -skin. 

Ehizomys.6 — 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,  Bull.  Soc.  Philom.  ser.  6,  vol.  xi.  p.  9  (1877). 

3  Waterhouse,  Proc.  Zool.  Soc.  1840,  p.  2. 

4  Peters,  Monatsber.  Ak.  Berlin,  1846,  p.  258. 

5  Giildenstadt,  Nov.  Comment.  Petrop.  vol.  xiv.  art.  i.  p.  409  (1770). 

6  Gray,  Proc.  Zool.  Soc.  1830,  p.  95. 


478  RODENTIA 


Subfamily  Bathyerginse. — 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  \,  m  f ;  no 
ear-conchs  ;  very  powerful  claws.  One  species  (B.  maritimus),  from 
South  Africa,  attaining  a  length  of  about  10  inches. 

Geary chus*  and  Myoscalops.3 — Upper  incisors  without  grooves. 
Georychus,  with  some  half  dozen  species,  generally  has  p  ^ ;  Myo- 
scalops, with  one  species,  usually  has  p  f ,  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.* — 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  $,  m  f^^L  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.  P  \ ;  molars 
rooted  or  rootless,  with  transverse  laminae.  Nearctic  and  Neo- 
tropical regions. 

Subfamily  Geomyinse. — Incisors  broad ;  mastoid  not  appearing 
on  the  top  of  the  skull ;  eyes  small ;  ear-conch  rudimentary ;  limbs 
short,  subequal.  Habits  fossorial. 

Geomys.® — Upper  incisors  deeply  grooved.  The  common  North 
American  Pouched-Rat  or  "Pocket-Gopher"  (G.  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.7 — Upper  incisors  plain.    Represented  by  two  species, 

1  Illiger,  Prodromus  Syst.  Mamm.  p.  86  (1811). 

2  Illiger,  loe.  tit.  p.  87. 

3  0.  Thomas,  Proc.  Zool  Soc.  1890,  p.  ^8-Heliophobius ;  Peters,  Monatsber. 
Ak.  Berlin,  1846,  p.  243. — Preoccupied. 

4  Riippel,  Mus.  Senkenb.  vol.  i.  Siiugeth.  p.  99  (1834). 

5  Including  the  Saccomyidce  of  Coues. 

6  Eafinesque,  Amer.  Monthly  Mag.  vol.  ii.  p.  45  (1817). 

7  Wied,  Nova  Ada  Ac.  Cces.  Leop.-Car.  vol.  xix.  pt.  i.  p.  383  (1839). 


DIPODID^E  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.  Entoptychus,  from  the 
Miocene  of  the  United  States,  is  an  allied  genus,  with  broad  incisors 
and  rootless  molars. 

Subfamily  Heteromyinse. — 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.1 — This  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  2  and  Heteromys.3 — 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  DIPODIDJE. 

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  Sminthince. — Molars  rooted ;  p  £,  m  f .  Skull  with 
the  infraorbital  vacuity  widest  below,  and  the  incisive  palatal 
foramina  long.  Limbs  short.  Palaearctic. 

Sminthw.4 — 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 
Muridce,  but  was  transferred  in  1887  to  the  present  family  by 
Dr.  H.  Winge. 

1  Gray,  Ann.  Mag.  Nat.  Hist.  vol.  vii.  p.  521  (1840). 
2  Wied,  Nova  Acta  Ac.  Cces.  Leop.-Car.  vol.  xix.  pt.  i.  p.  369  (1839). 

3  Desmarest,  Mammalogie,  p.  313  (1820). 
4  Keyserling  und  Blasius,   Wirbelthiere  Europ.  p.  38  (1840). 


480  RODENTIA 


Subfamily  Zapodinse. — Molars  rooted  ;  p  ±,  m  f ;  cervical  ver- 
tebrae free  ;  hind  limbs  elongated  ;  metatarsals  separate  ;  hind  feet 
with  five  digits.  Nearctic  region. 

Zapus.1 — The  American  Jumping-Mouse  (Z.  hudsonianus)  ex- 
tends over  almost  the  whole  North- American  continent  from  Labra- 
dor to  Mexico. 


Subfamily  Dipodinse. —  Molars  rooted ;  p  ^  r, ', 


m 


~5  ) 


vertebrae  more  or  less  ankylosed ;  hind  limbs  elongated ;  metatarsals 
united ;  hind  feet  with  only  three  functional'  digits.  Palaearctic 
and  Ethiopian  regions. 

This  subfamily  includes  the  true  Jerboas,  and  contains  three 
genera  :  Dipus  2  with  three  toes,  and  Alactaga  3  and  Platycercomys 4 
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. 

Eemains  of  the  existing  Alactaga  decumana  5  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  Pedetinse.  ^-Molars  rootless ;  cervical  vertebras  free  ; 
hind  limbs  elongated ;  metatarsals  separate ;  hind  feet  with  four 
digits.  Vertebra  :  C  7,  D  12,  L  7,  S  3,  C  30.  Ethiopian  region. 

Pedetes,6  the  Cape  Jumping-Hare  (P.  coffer),  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  zygoma  tic  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  OCTODONTID^E. 

Clavicles  complete.  Skull  with  long  incisive  foramina  extend- 
ing into  the  maxillae ;  and  usually  an  inferior  angle  to  the  jugal. 
Molars  with  external  and  internal  enamel -folds ;  p  ^,  except  in 
Ctenodactylus.  Mammae  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.  Petersbaurg,  1844,  p.  209. 
5  =  A.  jaculus,  Auct.  6  Illiger,  Prodromus  Syst.  Mamm.  p.  81  (1811). 


OCTODONTID^E 


481 


tion  of  one  species  of  Echinomys  which  ranges  into  Central  America. 
Habits  mostly  terrestrial,  but  occasionally  fossorial  or  natatorial. 

Subfamily  CtenodaetylinsB. — Molars  semi-rooted ;  jugal  as  in 
Dipodidce  ;  the  two  inner  toes  of  the  hind  feet  with  a  horny  comb 
and  rigid  bristles.  Ethiopian  region. 

Ctenodactylus.1 — 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  to  a  stump.  This  animal  is  about 
the  size  of  the  Water- Vole,  and  dwells  on  rocky  ground,  its  habits 


FIG.  213. — Skull  of  Hydrochcerus  capybara  (reduced). 

being  diurnal.  The  peculiar  comb-like  inner  toes  are  employed  for 
dressing  the  fur. 

Pectinator? — Closely  allied  to  the  preceding,  but  with  a  minute 
premolar  in  each  jaw ;  and  a  moderately  long  and  bushy  tail.  One 
species  (P.  spekei),  from  Somali-land. 

Subfamily  Oetodontinse. — 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.3 — Upper  and  lower  molars  alike  ;  ears  moderate  ;  tail 
of  medium  length  and  tufted.  Vertebrae :  C  7,  D  1 2,  L  7,  S  4,  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,  Spicilcgia  Zoologica,  p.  10  (1830). 

2  Blyth,  Journ.  As.  Soc.  Bengal,  vol.  xxxiv.  p.  294  (1855). 

3  Bennett,  Proc.  Zool.  Soc.  1832,  p.  46. 

31 


RODENTIA 


Habrocoma.1 — Lover  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.3 — 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.5 — 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  Eehinomyinse. — 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.  Vertebra? : 
C  7,  D  13,  L  6,  S  4,  C  25.  The  well-known  Coypu  (M.  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.'1 — 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  Hystricidce.  The  incisors  are 
smaller  than  in  the  Coypu,  and  the  upper  molars  have  one  internal 

1  Waterhouse,  Proc.  Zool.  Soc.  1837,  p.  30.     Amended  from  Abrocoma. 

2  Waterhouse,  Proc.  Zool.  Soc.  1841,  p.  91. 
3  De  Blainville,  Bull.  Soc.  Philom.  1826,  p.  62. 

4  Wagler,  ibid.  p.  1219. 
8  Andrew  Smith,  S.  African  Quart.  Journ.  vol.  ii.  p.  2  (1831). 

6  Geoffroy,  Ann.  du  Museum,  vol.  vi.  p.  81  (1805). 
7  Desmarest,  Mitn.  Soc.  d'Hist.  Nat.  vol.  i.  p.  44  (1822). 


OCTODONTID&  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.1 

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  (A.  swinderianus), 
from  Western  and  Southern  Africa,  which  attains  a  length  of  nearly 
2  feet,  and  dwells  in  burrows. 

Plagiodon? — 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.  cedium  of  Hayti 
and  Jamaica. 

Loncheres*  and  Echinomys.5 — 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  Echinomys  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. 

Dadylomys.7 — 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  Dadylomys  recalls  Chiromys 
among  the  Primates. 

Cercomys.8 — 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.  Soc. 
1884,  p.  233. 

2  Temminck,  Monographies  des  Mammifires,  vol.  i.  p.  245  (1827). 

3  Cuvier,  Ann.  Sci.  Nat.  ser.  2,  vol.  vi.  p.  347  (1836).     Amended. 

4  Illiger,  Prodromus  Syst.  Mamm.  p.  90  (1811). 

5  Desmarest,  Nouv.  Diet.  d'Hist.  Nat.  vol.  x.  p.  45  (1817).     Amended  from 
Echimys.  6  Wagner,   Wiegmann's  Archiv,  1845,  pt.  2,  p.  145. 

7  Geoffroy,  Ann.  Sti.  Nat.  ser.  2,  vol.  x.  p.  126  (1838). 

8  F.  Cuvier,  Mammiferes,  6me  livr.  (1829). 


484  RODENTIA 


Carterodon.1 — 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  Ododontidce  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  Morenia,  Orthomys,  and 
Trilodon  show  affinity  to  Myopotamus.  Pellegrinia,  from  the  Pleisto- 
cene of  Sicily,  may  be  allied  both  to  Ctenodactylus  and  Octodon. 


Family  THERIDOMYED^E. 

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  Ododontidce,  although  connected  by  Archceomys 
with  the  Chinchillidce.  The  dental  formula  is  the  same  as  in  the 
Ododontidce.  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  Archceomys  of  the  same  deposits  the  molars  are  rootless, 
with  the  enamel-folds  dividing  their  crowns  into  laminae,  as  in  the 
Chinchillas. 

Family  HYSTRICID^E. 

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  Synetherinae. — Molars  rooted ;  clavicles  complete ; 
upper  lip  not  cleft;  soles  tuberculated;  pollex  absent ;  four  mammae; 
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  Mamm.  vol.  ii.  p.  351  (1848). 
2  F.  Cuvier,  Dents  des  Mammiferes,  p.  256  (1825). 


HYSTRICIDA: 


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.  Eemains  of  Erethizon  occur  in  cavern-deposits 
in  Pennsylvania. 

Synetheres.1 — This  genus  contains  some   eight  or   ten  species, 
known  as  Tree  Porcupines  (Fig.  2 1 4),  found  throughout  the  tropical 


FIG.  214. — The  Tree  Porcupine  (Synetheres  prehensilis). 

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  a  hand.  Vertebrae  :  C  7,  D  17,  L  5,  S  3,  C  36.  An  extinct 
species  of  this  genus  has  been  described  from  the  cavern-deposits  of 
Brazil. 

1  F.  Cuvier,  Mem.  du  Museum,  vol.  ix.  p.  413  (1822).     "Sinethere." 


486 


RODENTIA 


Cfuetomys.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  Hystpieinse. — Molars  semi -rooted;  clavicles  incom- 
plete ;  soles  smooth  ;  a  rudimentary  pollex ;  six  mammae  ;  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. 

Hystrix.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.  Vertebrae:  C  7,  D  15,  L  4,  S  4,  C  12.  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.  africce-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  :  "  Hystrix  leucura  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. 
2  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  ]>.  76  (1766). 


CHINCHILLID&  4»7 

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  Hyslrix,  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  Hystrix  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. 
Eemains  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  Porcupines  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. 

Tricky s.2 —  This  genus  contains  but  one  Bornean  species  (T. 
guentheri),  externally  very  like  an  Atherura,  but  differing  from  the 
members  of  that  genus  in  many  important  cranial  characters. 

Family  CHINCHILLID.E. 

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  laminae.  Neo- 
tropical region.  This  family  includes  only  three  existing  species, 
divided  into  as  many  genera. 

Chinchilla.3 — In  this  genus  the  fore  feet  have  five  and  the  hind 
four  digits,  the  tail  is  long  and  bushy,  and  the  auditory  bullae  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  Eodent, 

1  Cuvier,  Regne- Animal,  2d  ed.  vol.  i.  p.  215  (1829).     "  Atherure." 

-  Giinther,  Proc.  Zool.  Soc.  1876,  p.  739. 
3  Bennett,  Gardens,  etc.  Zool.  Soc.  pt.  i.  p.  i.  (1829). 


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. 

Lagidiiim1  and  Lagostomus.2 — Lagidium  has  four  digits  in  both 
fore  and  hind  feet,  and  Lagostomus  three  only  in  the  hind  feet, 
and  the  auditory  bullae  are  much  smaller  than  in  the  preceding 
genus.  Lagidium  has  the  same  distribution  as  Chinchilla;  while 
Lagostomus,  as  represented  by  the  Viscacha  (L.  trichodactylus),  is 
found  in  the  Pampas  from  the  Uruguay  Kiver  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  Lagostomus  have  been 
described  by  Dr.  Ameghino  under  the  names  of  Prolagostomus, 
Pliolagostomus,  etc.  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  Epiblema  and  Tetra- 
stylus. 

Family  CASTOROIDID^E. 

Castoroides.3 — 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  Hydrochwrus,  but 
some  of  the  cranial  characters  are  like  those  of  the  Castoiida.  The 
genera  Amblyrhiza  and  Loxomylus,  from  the  Pleistocene  of  the 
Antilles,  appear  to  be  allied  types. 

Family  DASYPROCTID^E. 

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.4 — 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,  Nova  Ada  Ac.  Cccs.  Leop.-Car.  vol.  xvi.  p.  576  (1833). 

2  Brooks,  Trans.  Linn.  Soc.  vol.  xvi.  p.  102  (1828). 

3  Foster,  Second  Sep.  Geol.  of  Ohio,  p.  81  (1838). 

4  Illiger,  Prodromus  Syst.  Mamm.  p.  93  (1811). 


DINOMYID&—CA  VIID&  489 

Ccelogenys.1 — This  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.  taczanowskii. 

Family  DINOMYID^E. 

Distinguished  from  the  Dasyproctidce  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 
laminae. 

Dinomys.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 
Octodontidce,  Chinchillidce,  Dasyproctidce,  and  Caviidce. 


Family  CAVIID^E. 

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  laminae  ;  milk-teeth  shed  before  birth.  Other  characters 
as  in  Dasyproctida>.  Neotropical  region. 

Cavia.s — Limbs  and  ears  short,  subequal ;  tail  none.  Vertebrae  : 
C  7,  D  1 3,  L  6,  S  4,  C  7.  This  genus  includes  several  species  widely 

1  F.  Cuvier,  Ann.  du  Musium,  vol.  x.  p.  203  (1807). 

2  Peters,  Monatsber.  Ak.  Berlin,  1873,  p.  551. 

3  Pallas,  Misc.  Zool.  p.  30  (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 
1 0  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.  cutleri.  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.1 — 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. 
Vertebrae :  C  7,  D  12,  L  8,  S  3,  C  10. 

The  Patagonian  Cavy  (D.  patachonica) — 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. 

Hydrochcerus." — 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  Desinarest,  Mammalogie,  p.  360  (1822). 
-  Erxleben,  Syst.  Etg.  Animal,  p.  191  (1777)  ;  ex  Brisson. 


LAGOMYID&  49i 

one  having  some  twelve  transverse  laminae.  Upper  incisors  grooved. 
Vertebrae  :  C  7,  D  14,  L  6,  S  3,  C  8. 

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  Hydrochcerus  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  Caviidce.  Thus  Plexochoerus, 
from  the  Tertiary  of  Argentina,  differs  from  Hydrochcerus  in  having  only 
nine  laminae  in  the  last  upper  molar ;  Cardiomys,  Cardiatherium,  etc., 
from  the  infra-Pampeans  are  also  stated  to  be  allied  to  Hydrochcerus, 
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  confluent ;  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  LAGOMYID.E. 

Complete  clavicles,  subequal  limbs,  no  external  tail,  and  short 
ears.  Skull  depressed,  frontals  contracted  and  without  postorbital 
processes  ;  p  -j-  or  f- ;  molars  rootless,  Avith  transverse  enamel-folds. 
Palsearctic  and  Nearctic. 

Lagomys.1 — Represented  by  about  a  dozen  species  of  small 
1  Cuvier,  Tabl.  fiUmeiit.  de  I'Hist.  Nat.  p.  132  (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  f  as  Myolagus,  and  those  with  p  \  as  Titanomys,  but  this 
seems  scarcely  advisable. 


Family  LEPORID^E. 

Imperfect  clavicles,  elongated  hind  limbs,  short  recurved  tail, 

and  long  ears.  Skull 
(Fig.  216)  com- 
pressed, frontals 
with  large  wing- 
shaped  post -orbital 
processes  p  f ;  molars 
as  in  the  Lagomyidcp. 
Cosmopolitan  (ex- 
cept Australasia). 
Vertebrae:  C  7,  D 
12,  L  7,  S  4,  C  13- 
15. 

Lepus.1  —  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  Palsearctic  and  Nearctic  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 2)  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). 

2  From  the  absence  of  the  Common  Hare  in  Scandinavia  it  is  considered 
probable  that  the  name  L.  timidus  was  really  applied  to  the  Mountain  Hare, 
and  some  writers  accordingly  use  the  name  L.  europceus  for  the  former. 


FIG.  216.— Skull  of  Hare  (Lepus  timidus). 


LEPORID& 


493 


FIG.  217.— The  Common  Hare  (Leptis  timidits). 


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 
to  become  white 

during  winter  in  northern  countries,  while  assuming  a  reddish- 
yellow  hue  in  the  more  genial  climate  of  southern  Europe.  The 
Hare  is  a  nocturnal  animal,  remaining  during  the  day  on  its  "  form," 
as  the  slight  depression  is  called  which  it  makes  in  the  open  field, 
usually  among  grass. 

The   Mountain  Hare  (L.  variabilis)  is   found    throughout   the 

northern  part  of 
the  Pala5arctic 
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 
smaller  than  the 


common    species, 
with     a    smaller 
and  more  rounded 
In  cold  climates  the 


FIG.  218. — The  Mountain  Hare  (Lepus  variabilis). 


head,  and  shorter  ears,  tail,  and  hind  limbs. 


colour  of  the  whole  animal  changes  in  the  winter  to  a  pure  white 


494  RODENT/A 


(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 


PIG.  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  2|  to  3  Ibs.,  although 
individuals  perfectly  wild  have  been  recorded  up  to  more  than  5  Ibs. 
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. 
It  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&  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  Babbits  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  Eabbits,  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  bounds. 
In  other  cases  Eabbits  introduced  into  islands  have  become  or 
remained  more  or  less  distinct  from  their  parent  stock ;  thus  the 
Eabbits  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,1  the  Eabbits  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  Eabbits,  and  differing  in  certain  slight 
details  of  colour  and  habits. 

Bibliography  of  Rodentia,. — G.  R.  Waterhouse,  "  Observations  of  the  Rodentia," 
Mag.  Nat.  Hist.  iii.  (1839) ;  Id.  Ann.  Nat.  Hist.  viii.  and  x.  (1839-42) ;  Id. 
"On  the  Geographical  Distribution  of  the  Rodentia,"  Proc.  Zool.  Soc.  1839,  pp. 
162-174;  Id.  Natural  History  of  the  Mammalia,  vol.  ii.  "Rodentia"  (1848); 
Gervais,  Die.  Univ.  d'Hist.  Nat.  xi.  p.  202  (1848);  Brandt,  "  Untersuchungen 
iiber  die  craniologischen  Entwickelungsstufen  und  Classification  der  Nager  der 
Jetzwelt,"  Mem.  de  VAcad.  Impir.  de  St.  Petersbourg  (1855)  :  Lilljeborg, 
Systematisk  CEfversight  af  de  Gnagnde  Daggdjuren,  Upsala,  1866  ;  Alston,  "On 
the  Classification  of  the  Order  Glires,"  Proc.  Zool.  Soc.  1876,  pp.  61-98  ;  Trouessart, 
"  Catal.  de  Rongeurs,  Vivants  et  Fossiles,"  Bullet.  Soc.  d'fitudes  Scient.  d' Angers, 
1880-81  ;  Coues  and  Allen,  "Monographs  of  North  American  Rodentia,"  United 
States  Geol.  Surv.  of  Territories,  vol.  xi.  (1877);  Winge,  "Rodentia  pa  Lagos 
Santa,  Brazil,"  Mus.  Lund.  vol.  iii.  (1887) ;  various  papers  by  Peters  in  Monatsber. 
Ak.  Berlin,  and  by  Alston,  Anderson,  Blanford,  Dobson,  Milne -Ed  wards, 
Thomas,  and  others,  in  Proc.  Zool.  Soc.,  Journ.  Asiat.  Soc.  Beng.,  Ann.  Mag. 
Nat.  Hist.,  etc. 

1   Variations  of  Animals  and  Plants,  2d  ed.  vol.  i.  p.  119. 


CHAPTEE   XI 

THE    ORDER    CARNIVORA 

THOUGH  the  existing  Carnivora  as  at  present  restricted1  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  caecum  is  either  absent  or  short  and  simple  (Fig.  235),  and 
the  colon  is  not  sacculated,  or  greatly  wider  than  the  small  intestine. 
Vesiculse  seminales  are  never  present.  Cowper's  glands  are  present 

1  The  Ferse  of  Linnaeus  included  all  the  then  known  species  of  the  modern 
orders  Carnivora,  Insectivora,  and  Marsupialia. 

2  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. 


CARNIVORA    VERA  497 

in  some,  absent  in  other  groups.  The  uterus  is  bicornuate.  The 
mammae  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  f  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 


Fio.  220.— Left  upper  carnassial  teeth  of  Camivora.  I,  Fells;  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  Ursldce. 

in  extent,  but  is  generally  placed  near  the  anterior  end  of  the 
blade,  though  sometimes  it  is  median  in  position.  In  the  Ursidce 
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  Felidce  (I),  the  blade  alone  is  developed,  both 
talon  and  inner  cusp  being  absent  or  rudimentary.  In  others,  as 


CARNIVORA    VERA 


499 


Meles  (V)  and  Ursus  (VI),  the  heel  is  greatly  developed,  broad,  and 
tuberculated.  The  blade  in  these  cases  is  generally  placed  obliquely, 
its  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  (Latax)  than 
in  any  other  form,  but  even  in  it  they  can  be  traced. 

The  homology  of  the  various  parts  of  the  Carnivorous  carnassial 


FIG.  221.— Left  lower  carnassial  teeth  of  Carnivora.  I,  Felis ;  II,  Canis;  III,  Herpestes; 
l\,Lutra;  V,  Meles;  VI,  Ursus,  1,  Anterior  lobe  (paraconid)  of  blade  ;  2,  posterior  (protoconid) 
lobe  of  blade ;  3,  inner  cusp  (metaconid)  ;  4,  talon  (hypoconid).  It  will  be  seen  that  the 
relative  size  of  the  two  roots  varies  according  to  the  development  of  the  portion  of  the  crown 
they  have  respectively  to  support. 


with  the  primitive  tritubercular  type  (p.  30)  is  indicated  in  the 
figures.  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 
(not  seen  in  the  figure)  is  the  entoconid. 

The  toes  are  nearly  always  armed  with  large,  strong,  curved, 
and  tolerably  sharp  claws,  ensheathing  the  ungual  phalanges,  and 
held  more  firmly  in  their  places  by  broad  laminae  of  bone  reflected 
over  their  attached  ends  from  the  bases  of  the  phalanges.  In  some 
forms,  most  notably  the  Felidce,  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  Fetidce,  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 
^Eluroidea,  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  Carnivora,"  Proc.  Zool.  Soc.  1869,  p.  4  ;  Mivart, 
"On  the  Classification  and  Distribution  of  the  ^luroidea,"  ibid.  1882,  pp.  135 
and  459  ;  see  also  The  Cat,  an  Introduction  to  the  Study  of  Backboned  Animals, 
especially  Mammals,  by  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  palaeontological  classification  the  Canidce 
cannot  be  satisfactorily  separated  from  the  Ursidce  ;  while  in  another 
direction  the  Canidce  were  closely  allied  to  the  ancestral  Vivenidce. 
The  most  important  objection  against  this  classification  is,  however, 
the  apparent  intimate  connection  exhibited  by  fossil  forms  between 
the  Viwrridce  and  the  Mustelidce,  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  Ursidce  and  Mustelidce  have  been  independ- 
ently acquired,  in  the  course  of  the  descent  of  the  one  family  from 
a  Ganoid,  and  of  the  other  from  a  Viverroid  stock. 


Section 


The  ^Eluroidea  or  Cat-like 
Viverridce,  Proteleidce,  and  Hycenidce. 
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  Hycenidce)  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.  Caecum  small, 
rarely  absent  Os  penis  generally 

Small      and      irregular      (large      in 

CrilptOproda\         Cowper's      glands 
" 


Carnivores   include    the   Felidce, 


F]o  2,,._Left  side  of  the  1>alatal  .^  of 

the  cranium  and  mandible  of  the  Suricate(Sim- 
"«  Mradactyla).  c,  Carotid  foramen  ;  /,  fissure 
j.     .      ,,     i   ,      ,     in  floor  of  auditory  meatus.     From  Mivart, 

present  ;  prostate  distinctly  lobed.  pnCt  ZooL  Soc-  1882>  p>  184> 

Some  details  of  the  anatomy  of 

the  soft  parts  will  be  found  under  the  head  of  Genetta. 


502 


CARNIVORA 


Family  FELID^E. 

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  vertebrae  13. 

Felis.1 — The  whole  structure  of  the  animals  of  this  genus  ex- 
hibits the  Carnivorous  type  in  its  fullest  perfection.  Dentition : 
*  f '  c  T)  P  f '  m  T  >  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 

1  Linn,  Syst.  Nat.  12th  ed.  vol.  i.  p.  60  (1766). 


FIG.  223.— Front  view  of  skull  of  Lion  (Felis  leo). 


FELID&  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  bullae  are  large,  rounded,  and  smooth.  Vertebrae : 
C  7,  D  13,  L  7,  S  3,  C  13-29.  Clavicles  better  developed 
than  in  other  Carnivora,  but  not  articulating  with  either  the 
scapulae  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  papillae.  Caecum 
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  Mivart1  gave  the  number  of 
existing  species  of  Felis  as  48,  but  by  Mr.  Blanford's  reduction  of 
the  number  of  Indian  species 2  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  (F.  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  Pseonians  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). 
2  Fauna  of  British  India,  "  Mammalia,"  pp.  56-90  (1888). 


FELIDsE 


5°5 


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 
on  "  Felts  spelcea  "  in  Boyd  Dawkins  and  Sandford's  British  Pleisto- 
cene Mammalia  (1868).  Fossil  remains  attest  a  still  wider  range,  as 
it  is  shown  in  the  same  work  that  there  is  absolutely  no  osteologi- 


\ 


Fio.  224. — Lion  and  Lioness,  after  a  drawing  by  Wolf  in  Elliot's  Monograph  of  the  Felicke. 

cal  or  dental  character  by  which  the  well-known  Cave  Lion  (F. 
spelwa),  so  abundantly  found  in  cave-deposits  of  the  Pleistocene  age 
in  Western  Europe,  can  be  distinguished  from  the  existing  F.  leo. 

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,1  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  oj  Eastern  Persia  (1876). 


5o6  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  Eajputana, 
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.1 

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,2  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.  Selous3  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 
Felidce  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,  Fauna  of  British  India,  "Mammalia,"  p.  57  (1888). 

2  Transactions  of  the  Zoological  Society,  vol.  i.  p.  165  (1835). 

3  A  Hunter's  Wanderings  in  Africa,  1881,  p.  258. 


FELIDsE  507 

surface  of  the  body,  wanting,  however,  in  some  examples,1  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  Felidce ;  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  1 0  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  papillae  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  lies  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  Gumming : — 

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  usiial  pace  of  a  Lion,"  C.  J.  Andersson l  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. 


FELID.-E  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-Avitness 
gives  a  good  idea  of  Lion  family  life x : — 

"I  once  had  the  pleasure  of,  unobserved  myself,  watching  a 
lion  family  feeding.  I  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.  Druminond,  The  Large  Game  and  Natural  History  of  South 
and  South-East  Africa,  1875,  p.  278. 


5io  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  Felidce  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 


FELID^E  511 

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 : — 

"It  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,1  adult  males  measure  from  5£  to  6|  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 
are  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 


FIG.  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, 


FELIDsE  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  an  old  Tiger  whose  vigour  is  passed,  and  whose 
teeth  are  worn  and  defective ;  it  takes  up  its  abode  in  the  neigh- 
bourhood of  a  village,  the  population  of  which  it  finds  an  easier 
prey  than  the  larger  or  wilder  animals  named  above.  Though 
chiefly  affecting  grassy  plains  or  swamps,  it  is  also  found  in  forests, 
and  seems  to  be  fond  of  haunting  the  neighbourhood  of  old  ruins. 
As  a  rule,  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 
extract  on  the  habits  of  the  Tiger  is  taken  from  Sir  J.  Fayrer's 
Royal  Tiger  of  Bengal  (1875) : — 

"The  tigress  gives  birth  to  from  two  to  five,  even  six  cubs; 
but  three  is  a  frequent  number.  She  is  a  most  affectionate  and 
attached  mother,  and  generally  guards  and  trains  her  young  with 
the  most  watchful  solicitude.  They  remain  with  her  until  nearly 
full  grown,  or  about  the  second  year,  when  they  are  able  to  kill  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  eat  them  when  starved.  As  soon 
as  they  begin  to  require  other  food  than  her  milk,  she  kills  for 
them,  teaching  them  to  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  cruelty,  killing  apparently 
for  the  gratification  of  her  ferocious  and  bloodthirsty  nature,  and 
perhaps  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  far  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 

33 


5  M  CARNIVORA 


ground  in  the  vicinity  of  a  village  or  gowrie,  continue  its  ravages, 
destroying  one  or  two  cows  or  buffaloes  a  week.  It  is  very  fond  of 
the  ordinary  domestic  cattle,  which  in  the  plains  of  India  are 
generally  weak,  half-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  some  other  countries 
they  appear,  however,  to  be  on  the  increase ;  thus  according  to 
one  of  the  administration  reports  of  Java  laid  before  the  Dutch 
Chambers,  portions  of  that  island  are  being  depopulated  through 
Tigers.  In  1882  the  population  of  a  village  in  the  south-west  of 
the  Bantam  province  was  removed  and  transferred  to  an  island  off 
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  1887,  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  where  Tigers  are  not  so  common,  and  where 
they  can  pursue  their  agricultural  occupations  with  a  greater 
degree  of  security.  At  present  they  fear  to  go  anywhere  near 
the  borders  of  the  forest.  The  people  seem  disinclined,  or  they 
lack  the  means  and  courage,  to  attack  and  destroy  their  enemy, 
although  considerable  rewards  are  offered  by  Government  for  the 
destruction  of  beasts  of  prey.  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  otherwise 
does  injury 

The  Leopard  (F.  pardus,  Fig.  226),  although  belonging  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  (TrapSos  and 
TrapSaXts)  or  Panther  (iravd-^p)  by  the  ancients.  Leopard  (leo-pardus) 
is  a  later  term,  originally  applied,  it  is  believed,  to  the  Cheeta  or 
Hunting  Leopard,  upon  the  supposition  that  it  was  a  creature 
intermediate  between  the  Lion  and  the  true  Pard.  If  so  it  has 
been  completely  transferred  to  the  more  common  species,  and 
though  in  this  sense  a  perfectly  unnecessary  and  unmeaning  term, 
has  gradually  superseded  those  by  which  this  was  originally  known. 
Pard,  so  commonly  used  by  Elizabethan  authors,  is  now  nearly 
obsolete  in  the  English  language,  and  Panther  has  either  become 


FELID^E  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 


FIG.  226.— The  Leopard  (Fells  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  be  assigned 
definitely  to  either  one  or  the  other  form.1  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  vertebrae  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). 


5i6  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  4J  feet  in  length, 
and  the  tail  from  2|  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, 


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  British  Pleistocene 
Mammalia* 

The  Ounce,  or  Snow  Leopard  (F.  uncia),  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  Felidce ;  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.  nebulosa 2)  is  a  beautifully  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 
(Fig.  227)  is  a  good  example, 
are  found  throughout  the 
warmer  parts  of  Asia  and 
Africa.  The  Wild  Cat  of  Europe,  F.  catus,  still  inhabits  the 
mountainous  and  wooded  parts  of  Great  Britain. 

The  Caffre  Cat  (F.  caffra  3),  of  Africa  and  Southern  Asia,  was  the 
species  held  in  veneration  by  the  ancient  Egyptians,  and  immense 

1  Monograplis  of  the  Palccontographical  Society,  1872. 
2  Syn.  F.  inacrocelis.  :J  Syn.  F.  maniculata  and  caligata. 


FIG.  227.— The  Marbled  Cat  (Fdis  marnwrata). 
From  Blanford,  Mammalia  of  British  India,  p.  74, 
after  Elliot. 


5i8  CARNIVORA 


numbers  of  its  mummified  remains  have  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  F.  caffra  is  black,  and 
that  the  same  feature  obtains  in  the  darker  varieties  of  the  Domestic 
Cat;  while  in  F.  catus  there  are  only  spots  of  black  upon  this 
portion  of  the  limb.  Remains  of  the  Caftre  Cat  occur  in  the 
Pleistocene  cave-deposits  at  Gibraltar.  The  Indian  F.  rubiginosa  is 
the  smallest  species  of  Cat. 

The  Caracal  or  Persian  Lynx  (F.  caracal)  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  true  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  hair  that  grows  between  them. 
The  skull  and  skeleton  do  not  differ  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  are  exactly  those  of  the  other  Wild 
Cats,  and  they  are  exceeded  by  none  in  the  untameable  savageness 
of  their  disposition.  They  capture  their  prey  in  the  same  manner, 
either  lying  in  wait,  or  noiselessly  stealing  within  reach,  and  then 
making  a  sudden  rush  or  spring  upon  it.  Their  food  consists  of 
any  mammals  or  birds  which  they  can  overpower.  In  inhabited 
countries  they  commit  extensive  ravages  upon  sheep,  lambs,  and 
poultry.  Lynxes  generally  frequent  rocky  places  and  forests,  being 
active  climbers,  and  passing  much  of  their  time  among  the  branches 
of  the  trees.  Their  skins  are  of  considerable  commercial  value. 

Zoologists  are  by  no  means  agreed  at  present  as  to  the  specific 
distinctions,  if  any  really  exist,  between  the  various  modifications 
of  this  group.  As  many  as  eight  species  are  sometimes  recognised, 
four  belonging  to  the  Old  and  four  to  the  New  World.  The  former 


FELID& 


519 


are  F.  lynx,  of  Scandinavia,  Kussia,  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.  cervana,  Siberia ;  F.  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 


FIG.  228.— European  Lynx  (t'elis  lynx).    From  a  drawing  by  Wolf  in  Elliot's 
Monograph  of  the  Felidce. 

southern  California  by  F.  'maculata,  and  in  northern  Oregon  and 
Washington  territory  by  F.  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  (F.  isdbellina) 
is  inseparable  from  F.  lynx  ;  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. 


FIG.  2-J9. — The  Pinna  (Fells  concolor). 

B.  New  Wovlcl  Species. — The  Puma  or  Couguar  (F.  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, 


FELID^E  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  Xorth  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  monkej^s.  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  (F.  pardcdis)  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. 

F.  yaguanmdi,  rather  larger  than  the  Domestic  Cat,  with  an 
elongated  head  and  body,  and  of  a  uniform  brownish -gray  colour, 
ranges  from  Matamoras  to  Paraguay.  F.  eyra  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 


FIG.  -.'SO.— The  Ocelot  (Felis  pardulis). 

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  cavern- 
deposits  in  Madras.  In  the  Pliocene  Siwaliks  of  Northern  India 
the  huge  extinct  F.  wistata  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, 


FELID&  523 


some  of  which  were  as  large  as  a  Leopard.     F.  alrox  and  /'. 

of  the  Pliocene  of  the  United  States,  were  of  the  dimensions  of  the 

Lion. 

Cyncelurus.1  —  The  Cheeta  or  Hunting  Leopard  ((.'.  jubatus)  is  dis- 
tinguished from  the  other  Felidce  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  Genera.  —  A  number  of  forms  are  gradually  becoming 
known,  especially  through  the  researches  of  American  palaeonto- 
logists, which,  though  evidently  animals  of  the  same  general  type, 
and  therefore  to  be  placed  in  or  near  the  family  Felidce,  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  Procelurus,  from  the 
Upper  Eocene  and  Lower  Miocene,  with  p  £,  m  |,  ,an  alisphenoid 
canal,  and  a  third  trochanter  to  the  femur.  Dinictis,  of  the  North 
American  Miocene,  is  a  larger  allied  form,  with  p  j},  m  |  ;  the  upper 
carnassial  having  no  anterior  lobe,  and  the  lingual  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.  Psewlct'licnis,  from  the  French  Miocene,  is  another 
very  generalised  Feline,  in  which  there  may  be  either  three  or  four 
1  Wagler,  Syst.  Amphib.  etc.  p.  30  (1830). 


524  CARNIVORA 


premolars,  and  the  lower  carnassial  may  retain  its  inner  cusp. 
.Elurictis,  of  the  French  Phosphorites,  with  p  /^r^j,  in  gzsjp  together 
with  several  American  Miocene  genera,  such  as  Nimravus  (p  f, 
m  i),  Archcdurus  (p  ~^y  m  |),  Pogonodon  (p  f ,  m  }•),  and  Hoplo- 
phoneus  (p  ^^,  m  \},  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  Felidte  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  Machcerodus  nceogeus,  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  "  machaerodont "  type 
have  been  met  with  both  in  the  Old  and  New  World.  In  M. 
cultridens,  of  the  Upper  Pliocene  of  Italy  and  France,  the  upper 
canine  is  long  and  narrow,  with  smooth  cutting  edges ;  the  smaller 
form  described  as  M.  meganthereon  being  apparently  the  female 
of  this  species.  M.  creiiatidens,  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  M.  latidens  of  the  English  cavern-deposits.  The  Italian 
Pliocene  form  described  as  M.  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.  M.  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,  Eusmiliis,  from  the 
Upper  Eocene  Phosphorites  of  Central  France,  differs  from  all  other 
known  Felines  in  having  only  two  pairs  of  incisors  in  the  lower  jaw, 
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- 


VJVERRID^E 


525 


—  »«  l 


ment  of   the  sabre-toothed  or  machaerodont  dentition,   the   most 
specially  carnivorous  type  of  structure  known. 

Other  species  of  MacJuerodus  are  found  in  the  Pliocene  de- 
posits of  Europe  and  Asia.  The  accompany- 
ing woodcut  exhibits  the  last  two  upper  teeth 
of  the  Indian  M.  sivalensis,  from  which  it  will 
be  seen  that  the  inner  tubercle  of  the  carnassial 
is  much  reduced  in  size,  while  the  molar  is 
very  minute. 

Family 


Premolars  §  or  |.     Molars  ^  or  f  .     Upper 
carnassial  usually  without  an  anterior  lobe,  and 

J    .  .  ill  FlG-  281.—  Oral  surface 

the  lower  one  with  a  well  -developed  talon;  Of  the  left  tipper 
second  lower  incisor  (as  in  all  the  following  and  molar 
families)  raised  above  the  level  of  the  first  and  8 
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 
vertebrae  13  or  14.  Limited  in  distribution  to  the  Old  World.' 

The  subfamily  Cryptoproetinse  contains  the  single  genus  Crypto- 
procta.1  Dentition  :  i  §,  c  ^,  p  £  ,  m  \  ;  total  36.  The  teeth 
generally  closely  resemble  those  of  the  Felidce.  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  transversely. 
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.  Vertebra  :  C  7,  D  13,  L  7,  S  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.  ferox,  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 


Bennett,  Tram.  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  Fiverm  or  Civet  Cats  for  its  most  typical  repre- 
sentative, and  the  other  Herpestes  or  the  Ichneumons. 

Subfamily  Viverrinse. — Auditory  bulla  oval,  or  rather  conical, 
broad  and  truncated  and  not  everted  behind,  narrow  in  front  and 
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  genera,  has  the  follow- 
ing characters.  Dentition :  i  f,  c  ^,  p  |,  m  -|  (|  in  Prionodori)  ; 
total  40.  Skull  elongated ;  facial  portion  small  and  compressed. 
Orbits  well-defined  but  incomplete  behind.  Teeth  always  sectorial, 
never  very  small.  Vertebrae:  C  7,  D  13,  L  7  (or  D  14,  L  6), 
S  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. 

Fwerra.1 — 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 
1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  63  (1766). 


VIVERRIDsE  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  V.  civetta,  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, 


FIG.  232. — The  left  upper  dentition  of  the  Indian  Civet  (Viverra  zibetJuC).     From  the 
Palceontologia  Indica. 

and  V.  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  Fiverra,  although 
often  referred  to  a  separate  genus  (Fiverricula).  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  Fiverra  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  Ictitherium, 
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  V.  antiqua  has  an  alisphenoid  canal,  and  all  the 
other  cranial  characters  of  the  typical  forms. 

Fossa.1 — 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  Viveftra  is  absent ;  and  in  the  tail  the  spots  only  tend  to 
form  rings,  Avhich  are  not  complete.  The  skull  has  an  alisphenoid 
canal,  and  a  large  bulla  as  in  the  typical  group  of  Viverra. 

Genetta.1 — 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 


FIG.  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 

1  Cuvier,  Regne- Animal,  vol.  i.  p.  156  (1817). 


VIVERRID^E 


529 


oc 


FIG.  234.  —Stomach  of  Genet  cut  open,  a-,  (Esophagus  pv, 
pyloric  valve ;  x,  sudden  bend  where  the  internal  folds  are  inter- 
rupted. (From  Mivart,  Proc.  Zool.  Soc.  1882,  p.  505.) 


^Eluroidea)  of  the  anatomy  of  the  soft  parts  of  the  Genet  may  be 
given  as  illustration  of  these  parts  in  the  Carnivora  generally,  and 
of  this  family  and  genus  in  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 
its  (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- 
acteristic of  the 
Carnivora,  is 
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  pyloric  valve  is 
small.  The  allied  genera  present 
modifications  from  this  form  of 
stomach.  The  caecum  (Fig. 
235)  is  short,  thick,  and 
pointed.  The  liver  (Fig.  236) 
much  resembles  that  of  the 
Cat,  but  differs  in  that  the  left 
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  longer,  has  a  deep 
concavity,  and  runs  uninter- 
ruptedly into  the  Spigelian; 
the  latter  being  relatively  somewhat  larger  than  in  the  Cat, 
with  a  deep  groove  dividing  the  proximal  third  from  the  distal 
two-thirds.  In  Viverm  the  right  lateral  and  right  central  lobes 
are  nearly  equal  in  size.  The  variations  in  the  form  of  the  liver 

34 


Fin.  -235.— Caecum  of  Genet.    (After  Mivart, 
toe.  cit.  p.  508.) 


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- 


LL 


RC 


Pio.  236. — Abdominal  aspect  Of  the  liver  of  the  Genet,  c,  Caudal  lobe ;  gb,  gall-bladder  ;  lia, 
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 ;  vc,  vena  cava.  (From 
Mivart,  Proc.  Zool.  Soc.  1882,  p.  510.) 

tinguishes  the  brains  of  the  Felidce  from  those  of  all  other  members 

of  the  JEluroidea. 

Prionodon. x — 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- 
pletely haired.  The  pupil  is  round.  The 
caecum  (Fig.  237)  is  remarkably  small. 
This  genus  is  exclusively  Oriental,  and 

comprises  P.  gracttis  from  Borneo,  Java,  and  (?)  Sumatra,  P.  pardi- 
1  Horsfield,  Zool.  Research.  Java  (1824). — Pritmodontidcv. 


Fio.  237. — Caecum  of  Prionodon. 
(From  Mivart,  Proc.  Zool.  Soc.  1882, 
p.  508.) 


VIVERRIDA?  531 


color  from  Nipal,  and  P.  maculosm  from  Tenasserira ;  the  head  and 
body  of  the  latter  measuring  from  18  to  20  inches  in  length. 
Speaking  of  P.  pardicol&r,  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 


Fio.  238.— The  African  Linsang  (Poiana  poisnsis).    From  Mivart,  Proc.  Zool.  Soc.  18S2,  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  two 
litters  each  year." 

Poiana.1 — This  African  genus,  represented  solely  by  one  species, 
P.  poensis  (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  Genetfa.  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. 

Paradoxurus.1 — Dentition:  i  f,  c-^-,  p  £,  m  f-;  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.  Vertebrae:  C  7,  D  13,  L  7,  S  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  species2  are  P.  niger,  P.  hermaphroditus,  P.  jerdoni,  P.  aureus, 
P.  grayi  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.  Blanford3  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). 

2  See  W.  T.  Blanford,  Proc.  Zool.  Soc.  1885,  p.  780. 
3  Fautia  of  British  India,  "Mammalia,"  p.  108  (1888). 


VIVERRIDsE 


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." 

Ardogale.1 — This  genus — represented  only  by  A.  trivirgata  of 
Java,  and  A.  leucotis  of  Burma,  Tenasserim,  Sumatra,  Java,  etc. — 
is  chiefly  distinguished  from  Paradoxurus  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  2 6 '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 
considerably  lighter. 

Hemigale?  another 
modification  of  the 
Paradoxure  type,  contains  one  species,  H,  hardurickei,  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. 
2  Jonrdan,  Comptes  Rcndus,  vol.  v.  p.  442  (1837).     Amended. 


Fio.  239.— Palatal  aspect  of  the  left  side  of  the  cranium 
anil  mandible  of  Arctogale  leucotis.  a,  Anterior  opening  of 
alisphenoid  canal ;  o,  foramen  ovale ;  c,  carotid  canal  \. 
(From  Mivart,  Proc.  Zool.  Soc.  1882,  p.  165.) 


534  CARNIVORA 


characters    are   those    of   Paradoxurus,  but   the    auditory  bulla  is 
ankylosed  into  a  single  piece. 

Arctidis.1 — Dentition:  i  |,  c  $,  p  £,  m  f ;  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.  Vertebrae:  C  7,  D  14,  L  5,  S  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.  Caecum 
extremely  small.  But  one  species  is  known,  A.  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  Ardogale.  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,  N.  binotata,  a  somewhat 
aberrant  Paradoxure,  from  West  Africa.  It  is  rather  smaller  than 
the  true  Paradoxures,  with  smaller  and  more  pointed  molar  teeth, 
and  no  caecum.  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.3 — This  remarkable  genus  is  regarded  by  Professor 
Mivart  as  representing  a  third  section  of  the  F'iverrince;  it  contains 
one  species,  C.  bennetti  (described  by  S.  Mtiller  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  dcs  Mammiftrcs,    March    1824  ; 
Monographics,  vol.  i.  p.  xxi.  (1827). 

2  Gray,  List  of  Mamm.  Brit.  Mus.  p.  54  (1843). 

3  Gray,  Proc.  Zool.  Soc.  1836,  p.  88. 


VIVERRID^E  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  Herpestinse. — 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  Avail  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  Palsearctic  regions. 

Herpestes.1 — Dentition  :  i  f ,  c  ^,  p  £,  sometimes  f ,  m  f ;  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.  Vertebra:  C  7,  D  13,  L  7,  S  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  Illiger,  Prodromus  Syst.  Mamm.  p.  135  (1811). 


536  CARNIVORA 


snakes.  H.  ichneumon  was  a  sacred  animal  to  the  ancient 
Egyptians.  They  vary  considerably  in  appearance,  some,  as  H. 
galera  and  H.  win  (Fig.  240),  are  larger  and  heavier,  with  stouter 
body,  longer  limbs,  and  stronger  teeth.  The  common  Indian 
Mungoose  (H.  mungo)  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 


FIG.  240.— The  Crab-eating  Mungoose  (Herpestes  urva).     From  Blanford,  Mammalia  o/ 
British  India,  p.  130. 

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.  albicaudatus.  The  Oriental  H.  urva  (Fig.  246)  is  stated  to 
be  somewhat  aquatic  in  habits,  and  to  feed  on  frogs  and  crabs. 


VIVERRID.-E 


537 


Skull  shorter 


Kemains  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,1  premolars  §,  without  diastema  between  first  and 
second ;  soles  of  feet  completely  naked.  Contains  two  small 
South-African  species,  H.  parvula  and  H.  undulata. 

Bdeogale 2  contains  also  two  small  Ichneumon-like  animals,  11. 
crasskauda  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, 
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  caecum 
(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. 

Rhinogale* — Toes  5-5.     Claws  of  fore 
feet  short,  compressed,  acute.     Under  sur- 
face of  tarsus  hairy.     Palate  flat.     Founded  on  a  single  specimen 
from  East  Africa,  R.  melleri. 

Crossarchus.5 — Dentition:  i  f,  c  \,  p  |,  m  -§-;  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. 


FIG.  241. — Csecum  of  Cynictis 
penicillata.  (From  Mivart,  Proc. 
Zool.  Soc.  1882,  p.  508.) 


1  Gray,  Proc.  Zool.  Soc.  1861,  p.  308. 
2  Peters,  Mith.  Ges.  Nat.  Freunde  Berlin,  19th  November  1850. 

3  Ogilby,  Proc.  Zool  Soc.  1833,  p.  48. 

4  Gray,  Proc.  Zool.  Soc.  1864,  p.  573. 

5  F.  Cuvier,  Hist.  Nat.  des  Mammifires,  No.  199  (1825). 


538  CARN1VORA 


obscurus,  the  Kusimanse,  a  small  burrowing  animal  from  West 
Africa,  of  uniform  dark  brown  colour ;  C.  fasciatus ;  C.  zefoa ;  and 
(7.  gambianus. 

Suricata.1 — 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.  Vertebrae :  C  7,  D  15,  L  6,  S  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  4-4,  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.  tetradactyla,  a  small  gray-brown  animal, 
with  dark  transverse  stripes  on  the  hinder  part  of  the  back,  from 
South  Africa.  The  caecum  is  short. 

Galidictis,2  Galidea,5  and  Hemigalidea  4  are  names  of  three  slight 
generic  modifications  of  the  Viverrine  type, 
allied  to  the  Herpestince,  but  placed  by 
Mivart  in  a  distinct  subfamily,  Galidictiince. 
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,  Galidea  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  caecum  (Fig.  242)  is 
remarkable  for  its  comparative  length  and 
pointed  termination.  Hemigalidea  is  dis- 
tinguished by  the  absence  of  rings  on  the 
FIG.  242.-c*cum  of  Gaiuiea  tail  Cfalidictis  vittata  and  striata  chiefly 

ekgans.     (From     Mivart,    Proc.  .ITT.  •      Ai_    •          I 

zooi.  soc.  1882,  p.  508.)  diner  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  Fiverridce  in  the  weak  development  of  the  jaws  and  the 

1  Desmarest,   "Tabl.  Meth.  Mamm."  in  Nouv.  Diet.  d'Hist.  Nat.  vol.  xxiv. 
(1804).  2  Geoffrey,  Comptes  Jiendus,  1837,  p.  578. 

3  Geoffrey,  Mag.  de  Zool.  1839,  pp.  27,  37. 

4  Doyere,  Ann.  Sci.  Nat.  vol.  iv.  p.  281  (1835). 

5  Jourdan,  Comptes  Hendtcs,  1837,  p.  422.     Amended. 


PROTELEID&  539 


small  size  of  the  teeth  (Fig.  243),  in  consequence  of  which  it  was, 
when  first  discovered,  placed  in  the  order  Insectivora.  Dentition  : 
*  -$>  c  T>  .P  T>  ™  f  5  total  *0.  Vertebrae :  C  7,  D  13,  L  7,  S  3,  C  20. 
No  alisphenoid 
canal ;  an  entepi- 
condylar  fora- 
men to  the  hum- 
erus.  But  one 
species  is  known, 
E.  goudoti. 

Extinct    Gen- 
era.—The    Ter- 

,  •      •          r  J.T.      r\tj  Flo.  243. — Skull  of  Eupleres  goudoti.     t  natural  size. 

tianes  of  the  Old  Mug_  Roy-  Coll_  Surgeons. 

World        have 

yielded  several  genera  allied  to  the  existing  Viverroids,  some  of 
which  s.how  decided  signs  of  affinity  with  other  families.  Of  these 
the  Lower  Miocene  Amphidis  appears  to  be  nearly  related  to  Fiverra, 
but  is  distinguished  by  the  form  of  the  second  lower  molar,  which  is 
longer  and  has  two  distinct  roots.  Palaoprionodon,  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  Viverridce,  but  resembles  the  Mustelidce 
in  the  presence  of  a  glenoid  foramen  and  in  the  position  of  the 
condylar  foramen.  In  Stenoplesidis,  of  the  same  deposits,  the  dental 
formula  is  i  f ,  c  y,  p  ^,  m  -f- ;  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  Mustelidce,  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  Stenoplesidis. 
The  Lower  Pliocene  Iditherium  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  Hycenidce  that  it  is  practically  impossible  to  draw  up  a  defini- 
tion which  will  distinguish  the  two  families. 

The  North  American  Eocene  genera  Miacis  and  Didymictis  are 
generally  regarded  as  representing  a  separate  family — Miacidce — 
with  affinities  both  to  the  Viverridce  and  Canidce. 

Family  PROTELEIDJG. 

Skull  with  no  alisphenoid  canal;  and  the  auditory  bulla  divided 
into  two  distinct  chambers.  Dorsal  vertebrae  15.  Molars  ^.  Pre- 
niolar  and  molar  teeth  very  small  and  simple  in  character. 

Proteles.1 — This  genus  contains  but  a  single  species,  P.  cristatus, 
1  Geoffrey,  Mtm.  du  Museum,  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  Hyaenas,  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 

*  §>  c  \>  P  an(^  m 
dhs  total  30  or 
32.  Vertebrae : 
C  7,  D  15,  L  5, 

Fio.  244.—  Skull  and  Dentition  of  the  Aard-Wolf  (Proteles  cristatus).      g2    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  Hyaena,  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,  larvae,  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 
Hycenidce.1 

Family  HY^ENID^E. 

Skull  with  no  alisphenoid  canal ;  and  the  auditory  bulla  not 
divided  by  a  septum  into  two  chambers.  Dorsal  vertebrae  15. 
Molars  usually  ^,  but  in  some  fossil  forms  ^-,  or  f ,  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  Fiverridce,  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  Hycenidce  very  closely  with  the 
Fiverridce. 

Hycena.z — Dentition  in  existing  forms  usually  i  § ,  c  },  p  ^,  m 
1  For  Anatomy  of  Proteles  see  Flower,  Proc.  Zool.  Soc.  1869,  p.  474. 
2  Zimmermann,  Specimen  Zoologice  Geographical,  p.  365  (1777). 


HY^ENID^E  541 

\ ;  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  Felidce. 
Lower  carnassial  con- 
sisting of  little  more  than 
the  bilobed  blade.  Zygo- 
matic  arches  of  cranium  A  ^^  B 

Very     wide     and     Strong.          FIG.  245.— Outer  (A)  and  palatal  (B)  aspects  of  the  right 

Sao-ittal  rrpst,  hiVh    e-ivine1    uPPer  c41™188"11  ^°th  of  the  Striped  Hyaena  (Hyaxui 
Ulgn,  giving    gtrjata      From  tte  Quart.  Journ.Geol.Soc. 

attachment  to  very  power- 
ful biting  muscles.  Orbits  incomplete  behind.  Vertebrae :  C  7, 
D  15,  L  5,  S  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  Hya3na  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  Euhycenine  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  Hyaena  (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  Hyaena  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  3|  feet,  and  that  of  the  tail,  with  its  hair, 
1|  feet.  It  occurs  throughout  peninsular  India,  where  it  is  most 
common  in  open  hilly  districts,  and  in  North  Africa.  Mr.  Blanford l 
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 
Avide,  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  ItwLia,  "Mammalia,"  p.  133  (1888). 


542  CARNIVORA 


in  shape,  but  with  the  marks  of  the  hind  feet  conspicuously  smaller 
than  those  of  the  fore  feet.  Unlike  the  Spotted  Hyaena,  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  Hysena 
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 


FIG.  246. — The  Striped  Hysena  (Hyaena  striata). 

Hyaenas,  after  the  flesh  has  been  picked  off  by  vultures  and  jackals. 
Occasionally  sheep  or  goats,  and  more  often  dogs,  are  carried  off  by 
Hyaenas,  and  the  latter  at  all  events  are  often  taken  alive  to  the 
animal's  den."  The  Striped  Hyaena  is  essentially  a  cowardly  animal, 
and  one  that  is  much  more  silent  than  H.  crocuta.  Remains  of  H. 
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 
a  mane.  H.  crocuta,  the  Spotted  Hyaena  (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,1  this  species  may  be  considered  as  by  far  the  more 

1  The  anatomical  peculiarities  of  Hycena  crocuta  have  been  fully  elucidated  in 


543 


specialised  form.  The  Spotted  Hysena  is  a  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.  Eemains  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  Hycena  spelcea ;  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- 


FIG.  247.— The  Spotted  Hyaena  (Hyaena  crocuta). 

ence  has  been  already  made,  there  were  numerous  extinct  forms  of 
Hycena  in  the  upper  Tertiaries  of  Europe,  from  the  horizon  of  the 
Lower  Pliocene  Pikermi  beds  of  Greece  upwards.  In  the  Crocutine 
group  H.  colvini  of  the  Pliocene  of  India  (Fig.  248),  and  H.  robusta  of 
that  of  Italy,  appear  to  have  been  ancestral  forms  allied  to  H.  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. 
peirieri  makes  a  farther  step  towards  the  Crocutine  group  by  the 
loss  of  the  inner  cusp  in  the  lower  carnassial.  The  extinct  Hycenic- 

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.  grceca,  connects  H.  striata  with  Palhycena.  Both  are 
characterised  by  the  presence  of  a  small  second  lower  molar  behind 
the  carnassial ;  while  H.  grceca  also  has  four  lower  premolars.  Still 
more  generalised  is  the  Lychycenine  group,  comprising  H.  macrostoma 
of  India  and  H.  chceretis  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 
Viverridce.  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 


FIG.  248. — Outer  view  of  part  of  the  right  ramus  of  the  mandible  of  Hyceiia  colvini,  showing 
the  third  and  fourth  premolars  and  the  carnassial.    (From  the  Palceontologia  Indica.) 

carnassial.  The  Lower  Pliocene  Palhycena  hipparionum.  in  which 
the  dental  formula  is  i  §,  c  ^,  p  £,  m  f ,  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 
Hyaenas  have  been  gradually  modified  from  that  stock. 


Section  CYNOIDEA. 

Family  CANID^E. 

This  section  contains  the  single  family  of  the  Canidce,  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  ^Eluroid  and  Arctoid  forms.  In 
the  number  and  arrangement  of  the  teeth  they  more  nearly  approach 
the  primitive  heterodont  type  than  any  other  existing  Carnivora. 


CANID& 


545 


A  caecum  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 


FIG.  249.—  Right  lateral  aspect  of  the  skull  of  the  Dog  (Canis  familiaris). 

(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  caecum  (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-csecal  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  colon,  in  the  nat- 
less  aberrant  forms  to  which  the  names  of  Icticyon  ural  P08'*'*"1  *>>e 

-  .  ,         ,.     .       ,.        .      ~,        colon     is     upper- 

and  Cyon  have  been  given,  and  a  diminution  in  the  most. 
number  of  toes  in  the  Cape  Hunting  Dog  (Lycaori), 
which  has  4-4,  instead  of  5-4  as  in  the  remainder  of  the  family. 

35 


FIG.     250.— 
caecum  of  the  AM- 


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  Avhole  convenient  to  retain  all  the 
species,  with  the  exception  of  Otocyon,  Icticyon,  and  Lyceum,  in  the 
old  genus  Canis,  the  most  prominent  characters  of  which  are  the 
following.  Teeth,  usually  i  -f,  c  -J-,  p  f,  m  -f ;  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- 
't. \  crivorus).  In  certain 
P%  P 4  m  *  m  2  Asiatic  species  (C.  pri- 

FIG.  251.— The  last  four  left  upper  teeth  of  an  extinct  Wolf     <m(KVUS    and    its     allies) 
(Canis  cautleyi).    From  the  Palceontologia  Indica.  -,  .  i  ,  i  . 

which  on  this  account 

have  been  separated  to  form  the  genus  Cyan  of  Hodgson,  the  last 
lower  molar  (m  3)  appears  to  be  constantly  absent.  The  milk- 
dentition  is  di  -|,  dc  \,  dm  f ;  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, 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  56  (1766). 


CANID&  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- 
brae: C  7,  D  13,  L  7,  S  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.1 
All  the  toes  are  provided  with  exserted,  non- re  tractile,  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  holloAved  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." 


548  CARN1VORA 


Huxley1  has  based  his  division  of  the  group  into  two  parallel  series, 
the  Thooids  or  Lupine  forms  and  Alopecoids  or  Vulpine  forms, 
which  he  characterises  by  the  presence  of  frontal  air-sinuses  in  the 
former,  Avhich  not  only  affect  the  external  contour  but  to  a  still  greater 
degree  the  shape  of  the  anterior  part  of  the  cranial  cavity,  and  the 
absence  of  such  sinuses  in  the  latter.  The  pupil  of  the  eye  when 
contracted  is  round  in  most  members  of  the  first  group,  and  vertic- 
ally elliptical  in  the  others,  but  more  observations  are  required 
before  this  character  can  be  absolutely  relied  upon.  The  form  and 
length  of  the  tail  is  often  used  for  the  purposes  of  classification, 
but  its  characters  do  not  coincide  with  those  of  the  cranium,  since 
many  of  the  South  American  Canidce  have  the  long  bushy  tails  of 
Foxes  and  the  skulls  of  Wolves.  Taking  into  account  various 
combinations  of  these  and  other  minor  characters,  the  species  may 
be  arranged  in  the  following  groups,  which  some  authors  have 
considered  as  of  generic  importance. 

A.  TJwoid  or  Lupine  Series. — The  typical  group,  or  Canis  proper, 
contains  the  largest  members  of  the  genus,  the  true  Wolves  of  the 
northern  parts  of  both  Old  and  New  Worlds  (C.  lupus,  etc.),  the 
Jackals  of  Southern  Asia  and  Africa  (C.  aureus,  mesomelas,  etc.),  and 
the  various  breeds  of  the  domestic  Dog  (C.  familiaris).  The  true 
Wolves  are  (excluding  some  varieties  of  the  domestic  Dog)  the 
largest  members  of  the  genus,  and  have  a  wide  geographical  range, 
extending  over  nearly  the  whole  of  Europe  and  Asia,  and  North 
America  from  Greenland  to  Mexico,  but  they  are  not  found  in 
South  America  or  Africa,  being  replaced  in  both  of  these  continents 
by  various  species  of  Jackals  and  Foxes.  As  might  be  expected 
from  this  extensive  range,  and  the  varied  character  of  the  climatic 
conditions  of  the  countries  they  inhabit,  they  present  great  diversi- 
ties of  size,  length  and  thickness  of  fur,  and  coloration,  although 
resembling  each  other  in  all  important  structural  characters.  These 
differences  have  given  rise  to  a  supposed  multiplicity  of  species, 
expressed  by  the  names  of  C.  lupus,  C.  lycaon  (Central  Europe), 
C.  laniger  and  C.  niger  (Tibet),  C.  pallipes  (India),  C.  occidentalis, 
C.  nubUis,  C.  mexicanus,  etc.,  of  North  America,  but  it  is  very  doubt- 
ful whether  some  of  these  ought  to  be  distinguished  as  other  than 
local  varieties.  Mr.  W.  T.  Blanford,  in  his  recent  work  on  the 
mammals  of  India,  regards  the  two  forms  from  Tibet  mentioned 
above  as  inseparable  from  C.  lupus.  In  North  America  there  is 
a  very  distinct  smaller  species,  called  the  Coyote  or  Prairie  Wolf 
(C.  latrans)  ;  and  perhaps  the  Japanese  Wolf  (C.  Iwdophylax)  may  also 
be  distinct,  although,  except  for  its  smaller  size  and  shorter  legs,  it 
is  scarcely  distinguishable  from  the  common  species.  Though 
generally  distributed  throughout  the  Indian  peninsula,  the  Indian 
1  Proc.  Zool.  Soc.  Land.,  1880,  p.  238.  See  also  Mivart,  Dogs,  Jackals,  Wolves, 
and  Foxes;  a  Monograph  of  the  Canidce  (1890). 


CANID^E  549 

Wolf  (C.  pallipes),  which  is  rather  smaller  and  slighter  than  6'.  /w////.s, 
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  Extinct  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  GARNI  VORA 


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  II'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 
cfintury  (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.  lupus  are  common  in  the  European  Pleistocene ; 
while  the  Indian  Pliocene  C.  cautleyi,  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.  aureus,  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. 


CANID& 


55i 


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 — 


FIG.  252.— The  Jackal  (Cams  aureus). 

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  placenta!  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 
Fiverra,  and  is  precisely  paralleled  by  some  of  the  species  of  the 
extinct  genus  Cynodictis  mentioned  below.  The  muzzle  is  shorter 


552  CARNIVORA 


than  in  other  species,  and  the  facial  profile  is  slightly  convex 
instead  of  concave.  The  mammae  are  also  12  or  14  instead  of  the 
normal  1 0  ;  while  there  is  long  hair  between  the  f ootrpads.  Wild 
Dogs  inhabit  not  only  the  whole  of  the  Oriental  region,  but  extend 
into  Central  Asia  as  far  north  as  the  Altai  and  Amur-land  (C.  alpinus). 
C.  dukhunensis  ranges  from  the  forest  regions  of  peninsular  India  to 
Gilgit  and  Western  Tibet,  where  it  must  inhabit  open  country.  In 
their  general  form,  and  more  especially  the  shortness  of  the  legs, 
these  animals  come  nearer  to  the  Jackals  than  to  the  Wolves.  They 
hunt  their  prey  in  packs.  Eemains  of  species  of  this  group  occur 
in  the  cavern-deposits  of  the  Continent,  and  have  been  described 
under  the  name  of  C.  europceus. 

A  group  for  which  the  name  Lycalopex  has  been  proposed  com- 
prises certain  South  American  Canidce,  distinguished  from  Cam's 
proper  by  their  longer  tails  and  Fox-like  aspect : — C.  cancrivorus, 
C.  brasiliensis,  C.  melampus,  C.  vetulus,  C.  fulvicaudus,  C.  azarce,  (7. 
magellanicus,  C.  griseus.  The  last  three  have  been  further  separated 
(under  the  name  of  Pseudalopex)  on  account  of  slight  differences  in 
the  relative  size  of  the  molar  teeth,  and  of  their  pupil  being  ellip- 
tical when  contracted.  Nyctereutes  (one  species,  C.  procyonides,  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,  which  give  it  some  superficial  resemblance  to  a  Raccoon. 

B.  Alopecoid  or  Vulpine  Series. — The  Vulpine  group  (Vulpes) 
includes  the  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  C.  vulpes  (Vulpes  alopex), 
the  common  Fox  of  Europe ;  C.  niloticus,  adustus,  and  variegatus, 
Africa  ;  C.  flavescens,  montanus,  bengalensis,  japonicus,  corsac,  Asia ; 
C.  fulvus,  macrurus,  velox,  North  America.  Mr.  Blanford1  con- 
cludes, however,  that  the  Asiatic  C.  flavescens  and  C.  montanus,  and 
very  probably  the  North  American  Cross-Fox  (C.  fulvus)  are  merely 
local  races  of  C.  vulpes,  distinguished  by  certain  peculiarities  of 
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  its  bushy  tail  is  white.  Its  long,  sharp  muzzle,  erect  pointed 
ears,  and  sharp  eye,  give  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 
1  Fauna  of  British  India,  "Mammalia,"  pp.  153,  154  (1888). 


CANID^E  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.  Kemains  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 
bullse,  have  been  separated  under  the  name  of  Fennecus,  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.  ceningensis.  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  (7.  cwvipalatus,  of  the  size  of  a  small 
Fox,  which  appears  to  have  certain  resemblances  to  Otocyon. 

Lycaon.1 — 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  Hycena.  The  one  species,  L.  pictus,  the 
Cape  Hunting  Dog  (Fig.  253)  from  South  and  East  Africa,  is  very 
distinct  externally  from  all  the  other  Canidce.  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  Hycena  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  (/.  venaticus),  from  Guiana  and  Brazil, 
is  a  species  about  the  size  of  a  Fox,  with  close  hair,  and  short  legs 

1  Brookes,  Griffith's  Animal  Kingdom,  vol.  v.  p.  151  (1827). 

2  Lund,  K.  Danks.  Vid.  Selsk.  Afhand.  vol.  xi.  p.  62  (1845). 


554  CARNIVORA 


and  tail,  distinguished  from  all  other  Dogs  by  the  reduction  of  the 
molar  teeth  to  ^,  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  Speothos. 

Otocyon.1 — Dentition:  i  f,  c  ^-,  p  -f-,  m  ^^ ;  total  46  or  48. 
The  molar  teeth  are  thus  in  excess  of  any  other  living  heterodont 


FIG.  253.— The  Cape  Hunting  Dog  (Lycaon  pictus). 

mammal.  They  have  the  same  general  characters  as  in  Canis, 
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  bullae  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.  Vertebrae  :  C  7,  D  13,  L  7,  S  3,  C  22. 
Ears  very  large.  Limbs  rather  long.  Toes  5-4.  One  species, 
0.  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 

1  Lichtenstein,  Wiegmann's  Archiv.  1838,  vol.  i.  p.  290. 


CANID^E  555 

of  the  four  molar  teeth  as  a  survival  of  a  condition  of  the  dentition 
exhibited  by  the  common  ancestors  of  the  existing  Canidce  and  the 
existing  carnivorous  Marsupials.  There  is,  however,  at  present  no 
palaeontological  proof  of  this,  as  none  of  the  numerous  fossil  forms 
of  Canidce  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  Canidce,  although,  as  already  men- 
tioned, connecting  the  latter  so  closely  on  the  one  hand  with  the 
Fiverridce  and  on  the  other  hand  with  the  Ursidce,  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  Idicyon  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  Cynodidis,  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  Viverridce,  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  Cynodidis  is  not  far  removed  from 
the  ancestral  type  of  many  of  the  Viverroids  and  Ganoids,  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  Cynodidis  we  ought  to  regard 
the  Foxes  and  Jackals  as  the  descendants  of  Cynodidis,  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 
Daphcenus.  It  is  characterised  by  the  presence  of  three  upper 
molars — thus  bringing  up  the  dental  formula  to  the  full  Eutherian 
mimber;  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 l  the  minute  foramina 
1  Arch.  Mus.  Lyan.  vol.  iii.  art.  1,  p.  85  (1881). 


described  by  Professor  Cope1  in  the  postparietal  and  mastoid  which 
occur  in  Ursus,  but  are  said  to  be  absent  in  Canis,  are  present  in 
Amphicyon.  So  far,  however,  as  we  can  see,  the  presence  or  absence 
of  those  foramina  cannot  be  regarded  as  diagnostic  of  Ursus  and 
Canis,  although  they  are  generally  more  strongly  developed  in 
the  former.  Amphicyon  may,  indeed,  be  considered  as  a  very 
generalised  Dog,  with  affinities  to  the  Bears  in  the  structure  of 

its  limbs.  Dinocyon  is  a  still  larger  form, 
from  the  Middle  Miocene  of  France, 
which,  so  far  as  its  teeth  are  concerned, 
connects  Amphicyon  with  the  Ursoid 
genus  Hycenarctus  so  closely  as  to  render 
it  absolutely  impossible  to  indicate  any 
characters  of  family  importance  by  which 
they  can  be  distinguished.  The  upper 
carnassial  of  Dinocyon  is  unknown.  For 
other  genera,  see  p.  562. 

Section  ARCTOIDEA. 

This  section  includes  a  considerable 
number  of  forms  which  agree  in  the 
essential  characteristics  of  the  struc- 
tures of  the  base  of  the  cranium  and 
reproductive  organs,  and  in  the  absence 
of  a  caecum  to  the  intestinal  canal. 
They  have  no  Cowper's  glands,  but 
there  is  a  rudimentary  prostate  and  a 
large  cylindrical  penial  bone ;  while  all 
the  members  of  the  group  have  five 
completely  developed  toes  on  each  foot. 
Considerable  diversity  is  found  in  the 
characters  of  the  base  of  the  skull  in 
the  various  forms,  but  the  following 
features  are  common  to  all.  The  cavity 
.  .«.  ,  , ,  of  the  auditory  bulla  is  simple,  and  has 

Fio.  254.— Right  half  of  the  palatal  J      ...    ....  , 

aspect  of  the  cranium  of  the  Raccoon  no  trace  of  a  dividing  septum ;  the 
(Procyon  lotor).  Letters  as  in  Fig.  s,  inferior  lip  of  the  auditory  meatus 

fwTp.  io7m  the  PrOC'  ZO°L  SOC'   (am>    FiS'    254>    is    considerably    pro- 
longed ;  the  paroccipital  process  ( p)  of 

the  exoccipital  is  more  or  less  triangular,  directed  backwards, 
outwards,  and  downwards,  and  standing  quite  apart  from  the 
bulla ;  the  mastoid  process  (m)  of  the  periotic  is  always  widely 
separated  from  the  paroccipital,  and  generally  very  prominent; 
the  carotid  foramen  (car)  is  large,  and  placed  on  the  inner  margin 

1  Proc.  Amcr.  Phil.  Soc.  vol.  xviii.  p.  452  (1880). 


URSID&  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  ^lurus. 

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,  Ursidce,  Procyonidce, 
and  Muslelidce.1 

Family  URSIDCE. 

In  existing  forms  the  true  molars  -f ,  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  jEluropus).  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* — Dentition  :  i  f ,  c  ^,  p  £ ,  m  f ;  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.  Vertebrae:  C  7,  D  14,  L  6,  S  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.  Ears  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  earth'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 


FIG.  255.— Head  of  the  Brown  Bear  (Ursus  arctos).    From  Sclater,  Proc.  Zool.  Soc.  1867,  p.  817. 

Bear,  usually  purely  carnivorous  or  piscivorous,  devours  grass  with 
avidity  in  summer.  The  various  species  may  be  arranged  in  the 
following  groups  : — 

Thalassarctine  Group. — Head  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  few 
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  (17.  arctos)  is  the  best  known 
example.  This  species  is  an  exceedingly  variable  one,  and  has  a 
very  wide  range  in  the  Palsearctic  region ;  the  Syrian  form  described 
as  U.  syriacus,  as  well  as  the  Hairy-eared  Bear  (U.  piscator,  Fig. 
255)  of  North-Eastern  Asia,  and  the  Snow-Bear  (U.  isdbdlinus)  of 


URS1D/E  559 

Kashmir  and  Nipal,  not  being  specifically  separable.  The  Brown 
Bear  hibernates  in  cold  regions,  and  in  the  Himalaya  keeps  to 
comparatively  high  regions,  emerging  from  its  winter  lair  in  March, 
April,  or  May,  according  to  the  season  and  elevation,  to  feed  on 
the  numerous  bulbous  plants  which  abound  in  the  regions  it  inhabits. 
Both  the  Syrian  and  Himalayan  varieties  are  generally  of  lighter 
colour  and  smaller  size  than  the  typical  European  form.  Bears 
were  at  one  time  found  in  the  British  Isles,  from  which,  how- 
ever, they  have  been  long  since  exterminated.  They  are  still  found 
in  the  Pyrenees,  and  are  comparatively  abundant  in  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  seen  no  less  than  seven  examples 
at  one  time  from  the  top  of  a  mountain  ridge ;  of  late  years  their 
numbers  have,  however,  been  greatly  diminished.  The  Brown 
Bear,  although  with  strong  powers  of  smelling,  is  very  slow  of 
sight  and  hearing,  and  in  the  Himalaya  it  is  easy  to  approach  so 
near  that  they  may  be  shot  with  a  smooth-bore  gun.  The  Grizzly 
Bear  (U.  h&rribilis)  of  North  America  is  so  closely  allied  to  the 
Brown  Bear  that  some  writers  think  it  should  only  rank  as  a  very 
well-marked  local  variety.  The  Black  Bears  of  the  Himalaya  (U. 
torquatus),  Japan  ( U.  japonicus),  and  North  America  ( U.  americanus) 
belong  to  this  group.  The  Himalayan  species  ranges  from  Persia 
to  Assam,  and  thence  to  China  and  Formosa.  In  the  greater  part 
of  this  area  it  is  essentially  a  forest  animal,  and  may  be  found  in 
autumn  in  the  forests  of  the  Kashmir  valley  feeding  upon  chestnuts 
and  other  fruits.  It  is  also  exceedingly  fond  of  maize,  mulberries, 
and  walnuts ;  and  a  few  years  ago  it  was  no  very  uncommon 
sight  to  see  three  or  even  five  of  these  bears  up  a  single  mulberry 
or  walnut  tree  in  Kashmir.  The  Spectacled  Bear  (U.  ornatus)  of 
the  Peruvian  Andes  is  another  member  of  this  group. 

The  Helarctine  group  is  represented  only  by  the  Malay  Bear  or 
Sun  Bear  ( U.  malayanus),  in  which  the  head  is  short  and  broad  ;  the 
molar  teeth  are  comparatively  broad  (but  the  length  still  exceeding 
the  breadth),  the  tongue  is  very  long  and  extensile,  and  the  fur 
short  and  smooth.  This  small  species  inhabits  the  Malay  Peninsula, 
Sumatra,  Java,  Borneo,  Tenasserim,  Arakan,  Chittagong,  and  the 
Garo  hills  of  India ;  it  inhabits  forest  districts,  and  is  an  expert 
climber. 

The  earliest  known  occurrence  of  the  genus  is  in  the  Lower 
Pliocene  of  the  Indian  Siwalik  Hills ;  where  it  is  represented  by 
U.  theobaldi,  which  was  probably  the  ancestor  of  the  existing 
Melursus.  The  genus  is  represented  in  the  Upper  Pliocene  of 
Europe  by  the  small  U,  etruscus  ;  and  in  the  Pleistocene  by  the  exist- 
ing U.  arctos,  as  well  as  by  the  great  extinct  Cave-Bear  ( U.  spdmis\ 
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. 
Eemains  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.1 — 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,  M.  Idbiatus,  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. 


FIG.  256. — JEluropus  melanoleucus.    (From  Milne-Edwards.) 

2 — Dentition  :  i  f ,  c  j-,  p  £,  m  f ;  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.  neu.  Zool.  Entdeckimgen,  etc.  p.  155  (1793). 

2  A.  Milne-Edwards,  Nouv.  Arch,  du  Museum,  vol.  viL  Bull.  p.  88  (1871). 
Amended  from  "  Ailuropus." 


URSIDA: 


561 


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,  A.  melanoleucus  (Fig. 
256),  discovered  by  Pere  David  in  1869,  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  eyes,  shoulders  and  limbs  black.  In  the  large  size  and 
complex  crowns  of  the  upper  premolars  this  genus  differs  very 
markedly  from  the  true  Bears.  The  fourth  upper  premolar  (car- 
nassial)  makes  no  approach  to  the  markedly  sectorial  type  presented 
by  the  corresponding  tooth  of  Hycenarctus,  its  structure  being,  on 
the  whole,  more  like  that  of  dElurus. 

Extinct  Genera. — The  genus  Arctotherium  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 
crowns  of  the  upper 
molars  are  broader  and 
shorter.  The  humerus 
is  said  to  have  an 
entepicondylar  fora- 
men. Hycenarctus,  of 
the  Miocene  and  Plio- 
cene of  Europe  and 
Southern  Asia,  has  the 
crowns  of  the  upper 
molars  either  square  or 
triangular ;  the  upper 
carnassial  having  three 
distinct  lobes  to  the 
blade,  while  the  lower 
carnassial  is  practically  indistinguishable  from  that  of  the  Dog-like 
Dinocyon  (p.  556).  The  proximal  extremity  of  the  ulna  differs 
from  that  of  Ursus  in  having  a  long  olecranon,  and  thereby  re- 
sembles the  corresponding  bone  of  the  Dogs.  Indeed  all  the 
characters  at  present  available  tend  to  show  a  complete  passage 
from  the  Tertiary  Dog-like  animals,  through  Dinocyon,  Hycenarctus, 

36 


Fio.  257. — Palate  of  Arctotherium  bonariense,  Pleistocene, 
South  America — J  natural  size.  (From  the  Palceontologia 
Indica.) 


562 


CARNIVORA 


and  Arctvtlierium,  to  the  true  Bears.  Most  of  the  species  of  Hyce- 
narctus  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  Hycenardus.  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  Procyonida.  Among  these  are  Simocyon  from 
the  Pliocene  of  Europe,  with  p  ^£5,  m  f ;  and  Enliydrocyon  of  the 
North  American  Miocene,  with  p  f ,  m  f ,  a  secant  talon  to  the 
lower  carnassial,  and  a  very  short  skull.  The  Miocene  JElurodon 
comprises  several  large  North  American  forms,  having  a  trilobed 
upper  carnassial  like  that  of  Hycenarctus,  and  a  dental  formula 
similar  to  that  of  the  latter  and  Canis  Prohycena  is  founded  upon 
a  much- worn  jaw  of  ^lurodon.  Hycenocyon,  of  the  Miocene  of  the 
United  States,  with^>  f,  m  4>  appears  to  be  an  allied  form,  also 
having  a  trilobed  upper  carnassial. 

Family  PROCYONID.*:. 

True  molars  f ,  tuberculated  or  multicuspid ;  upper  carnassial 
short  and  broad.  Alisphenoid  canal  absent,  except  in  JElwrus. 
Feet  plantigrade.  Tail  generally  annulated.  In  some  cases  an 
entepicondylar  foramen  to  the  humerus.  Typically  American,  but 
with  the  outlying  Oriental  genus  jElurus. 

j^Elurus,1 — Dentition  :  i  f ,  c  ^,  p  f ,  m  f- ;  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.  Vertebra? :  C  7,  D  U,  L  6,  S  3,  C  18.  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,  ^.  fulgens,  the  Panda  (Fig.  258),  an  animal  rather  larger 
than  a  Cat,  found  in  the  South-East  Himalaya,  at  heights  of  from 
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  jElurus  has  been  made  the  type  of  a  distinct  family, 


1  F.  Cuvier,  Hist.  Nat.  dcs  Mammiftrcs  (1825).     Amended  from 
For  anatomy,  see  Flower,  Proc.  Zool.  Soc.,  1870,  p.  752. 


'Ailurus." 


PROCYONIDsE 


563 


but  its  relationship  to  the  Raccoons  is  regarded  by  Mr.  W.  T. 
Blanford l  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 


Fio.  258.— The  Panda  (/EZttnts  /uZj/ens).     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  dSlurus  (&.  anglicus)  have  been 
obtained  from  the  English  Pliocene  Crag  deposits  which  indicate  an 
animal  of  about  one  and  half  times  the  size  of  dE.  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  JElurus  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  are 
characterised  by  the  absence  of  an  alisphenoid  canal  in  the  skull. 

Procyon.1 — Dentition  :  i  f,  c  ±,  p  £ ,  m  f ;  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 ; 


Fio.  259. — Lateral  view  of  skull  and  right  half  of  palate  of  sElurus  fulgens.    (From  Blanford, 
Mammalia  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,  but  narrower  behind,  with  five 
obtuse  cusps.  Vertebrae:  C  7,  D  14,  L  6,  S  3,  C  16-20.  Body 
stout.  Head  broad  behind,  but  with  a  pointed  muzzle.  Limbs 

1  Storr,  Prodromus  Meth.  Mamm.  p.  35  (1780). 


PROCYONW^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 x 
(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 


FIG.  200.— The  Raccoon  (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  Mammals  of  the  Adirondack  fiegion : — 

"  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 

1  A  corruption  of  the  North  American  Indian  "arrathkune"  or  "arathcone." 
The  French  raton  or  raton  laveur,  German  Waschbar,  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.1 — 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  Viverridce.  Two  species  : — B.  astuta,  from  the  southern 
parts  of  the  United  States  and  Mexico,  and  B.  sumichrasti,  from 
Central  America. 

Bassaricyon.2 — 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.  alleni  They  much  resemble  the  Kinkajou  (Cercoleptes)  in  external 
appearance,  but  the  skull  and  teeth  are  more  like  those  of  Procyon 
and  Nasua. 

Nasua? — 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- 
brae:  C  7,  D  14,  L  6,  S  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  tAventy,  are  chiefly  arboreal,  and  feed 
on  fruits,  young  birds,  eggs,  insects,  etc.  Recent  researches  have 
reduced  the  number  of  supposed  species  to  two,  N.  narica  of  Mexico 

1  Lichtenstein,  Isis,  1831,  p.  512. 
2  Allen,  Proc.  Ac.  Nat.  Sci.  Philad.  1876,  p.  20. 
3  Storr,  Prodromus  Meth.  Mamm.  p.  35  (1780). 


MUSTELID^E  567 


and  Central  America,  and  N.  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.1 — Dentition  :  i  f ,  c  ^,  p  f ,  m  f ;  total  36.  Molars 
with  low  flat  crowns,  very  obscurely  tuberculated.  Skull  short  and 
rounded,  with  flat  upper  surface.  Vertebrae :  C  7,  D  1 4,  L  6,  S  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,  C.  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  MUSTELID^E. 

True  molars  J-  (or  ^  in  Mellivora  2).  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  Procyonidce  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  Lutrinse. — Feet  short,  rounded  (except  the  hind  feet  of 
Latax).  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. 

Lutm.s — Dentition  :    i  f ,   c  1,  p  £,   m  | ;    total   36.      Upper 

1  Illiger,  Prodromus  Syst.  Afamm.  et  Avium,  p.  127  (1811). 
"  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  im- 
mediately behind  the 
orbits.  Facial  portion 
very  short ;  brain  case 
large.  Vertebrae :  C  7, 
D  14-15,  L  6-5,  S  3,  C 

FIG.  261.— Palate  of  Lutra  cinerea.    (From  the  20-26.       Body  very   long. 

Palceontologia  Indica.)  1^1.  j  i     i 

Jbars  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.  Tail  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  Avebs,  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  3£  feet,  of  which  the  tail  occupies 
1  foot  3  or  4  inches.  The  weight  of  a  full-sized  male  is  from  1 8  to 
24  Ibs.,  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 


MUSTELIDsE 


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.1  Besides  those  mentioned  above,  the 
following  may  be  noticed.  In  the  Oriental  region  there  are  L. 
ellioti 2  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  Aonyx.  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.  bmsiliensis.  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.  palceindica, 
which  is  closely  allied  to  L.  sumatrana,  and  a  larger  form  described 
as  L.  bathygnathus.  The  Pliocene  of  Hessen- Darmstadt  yields 
L.  hessica ;  while  L.  diibia,  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  described  as  Trochictis,  which  are  believed  to  connect  Lutra 
with  the  Mustelince.  Two  very  large  Otters,  respectively  from  the 
Indian  Siwaliks  and  the  Italian  Miocene,  named  L.  simlensis  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.  valetoni)  from  the  Lower  Miocene  of  the 
Continent,  which  differs  from  all  other  known  Mustdidce  in  having 
a  minute  second  upper  true  molar.  This  species  is  evidently  a 
very  generalised  form  approximating  to  the  Viverridce-  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  Lufni 
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  Viverridce,. 

Latax.1 — Dentition  :  i  f ,  c  i,  p  f ,  m  4- ;  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  Ada  Ac.  Cces.  Leap. -Car.  vol.  xiii.  pt.  2,  p.  511  (1827) :  Syn. 
Enhydra  ;  Fleming,  Philosophy  of  Zoology,  vol.  ii.  p.  187  (1822).  Preoccupied  by 
Enhydris,  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. 


' 


FIG.  2(52.— The  Sea-Otter  (late  lutris).    From  Wolf,  Proc.  Zool.  Soc.  18(55,  pi.  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  Melinse. — 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.1 — Dentition:  i  f,  c  \,  p  %,' m  ^;  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.  Vertebrae :  C  7,  D  16, 
L  6,  S  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  (M.  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  insectreater, 

1  Cuvier,  "Tabl.  de  Classif."  in  Lemons  d'Anat.  Compar.  vol.  i.  (1800). 


MUSTEUD& 


573 


he  destroys  more  beetles,  grasshoppers,  and  the  like  than  all  our 
other  mammals  together,  and  in  addition  to  these  he  devours  vast 
numbers  of  mice.  He  does  not  evince  that  dread  of  man  that  is  so 
manifest  in  the  great  majority  of  our  mammals,  and  when  met  during 
any  of  his  circumambulations  rarely  thinks  of  running  away.  He 
is  slow  in  movement  and  deliberate  in  action,  and  does  not  often 
hurry  himself  in  whatever  he  does.  His  ordinary  gait  is  a  measured 
walk,  but  when  pressed  for  time  he  breaks  into  a  low  shuffling 
gallop.  It  is  hard  to  intimidate  a  skunk,  but  when  once  really 
frightened  he  manages  to  get  over  the  ground  at  a  very  fair  pace. 


FIG.  263.— The  Common  Skunk  (Mephitis  mephitica). 

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  influence  whatever 
upon  their  movements.  Skunks,  particularly  when  young,  make 
very  pretty  pets,  being  attractive  in  appearance,  gentle  in 
disposition,  interesting  in  manners,  and  cleanly  in  habits — rare 
qualities  indeed !  They  are  playful,  sometimes  mischievous,  and 
manifest  considerable  affection  for  those  who  have  the  care  of  them. 
Their  flesh  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  papillae  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  (M.  macruni),  from  Central  and  Southern 
Mexico,  has  two  lateral  stripes,  and  a  longer  and  more  bushy  tail 
than  the  common  species.  M.  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.1 — 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  Mephitis  being  almost 
invariably  absent,  so  that  the  dental  formula  is  i  f ,  c  ^,  p  -|, 
m  % ;  total  32. 

Remains  of  Conepatus,  which  have  been  referred  to  three  species, 
are  found  in  the  cavern-deposits  of  Brazil. 

Arctonyx? — Dentition :  i  f ,  c  ^,  p  £ ,  m  £ ;  total  38.  Incisor 
line  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  an tero- 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.  Vertebrae :  C  7, 
D  16,  L4,  S4,  C  20.  Snout  long,  naked,  mobile,  and  truncated, 
with  large  terminal  nostrils,  much  like  that  of  a  Pig.  Eyes  small. 
Ears  very  small  and  rounded.  Body  compressed  rather  than 
depressed.  Limbs  of  moderate  length  and  digitigrade  in  walking. 

1  Gray,  Ann.  Mag.  Nat.  Hist.  ser.  2,  vol.  i.  p.  581  (1837). 
2  F.  Cuvier,  Hist.  Nat.  des  Mammiferes  (1825). 


MUSTELID^E  575 

Tail  moderate,  tapering.  A  full  soft  under  fur,  with  longer,  bristly 
hairs  interspersed.  The  best-known  species  is  A.  colhiris,  the  Sand- 
Badger,  or  Bhdlu-soor l  (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  A.  taxoides  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 
Arctonyx  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. 

Mydaus.'2 — 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.  Vertebrae  :  C  7, 
D  14-15,  L  6-5,  S3,  C  12.  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,  M.  meliceps, 
the  Teledu,  a  small  burrowing  Badger,  found  in  the  mountains  of 
Java  at  an  elevation  of  7000  or  more  feet  above  sea-level. 

Meles.3 — Dentition  :  i  -f ,  c  ^,  p  $,  m  £ ;  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  strongly  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. 
Vertebrae :  C  7,  D  15,  L  5,  S  3,  C  18.  Muzzle  pointed.  Ears  very 
short.  Body  stout,  broad.  Limbs  short,  strong,  subplantigrade. 
Tail  short.  The  best-known  species  is  the  common  Badger  (M.  taxus) 
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,  M.  leucurus  and  M.  chinensis,  are  found  in 
continental  Asia,  M.  canescens  in  Persia,  and  M.  anakuma  in  Japan. 

The  appearance  of  the  common  Badger  is  too  well  known  to 

1  Possibly  the  name  should  be  Balu-soor  (Sand-pig). 

2  F.  Cuvier,  Hist.  Nat.  des  Mammiferes  (1825). 

3  Storr,  Prodromus  Meth.  Mamm.  p.  34  (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  2J  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.1 — Dental  formula  as  in  Meles,  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  Meles. 
Vertebrae:  C  7,  D  15,  L  5,  S  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.  americaiia  is  the  common  American  Badger  of  the  United  States ; 
T.  berlandieri,  the  Mexican  Badger,  is  perhaps  only  a  local  variety. 

Mellivwa.'2 — Dentition  :  if,  c  y,  _p  •§-,  m  \;  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.  Vertebrae  :  C  7,  D  14,  L  4,  S  4,  C  15. 
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.  M.  indica  from 
India,  and  M.  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,  Proc.  Zool  Soc.  1838,  p.  154. 
2  Storr,  Prodromus  Meth.  Mamm.  p.  34  (1780). 


MUSTELIDA-:  $77 


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 


FIG.  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  Circars.  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 
Melliwrodon. 

37 


578 


CARNIVORA 


Helictis.1 — Dentition  :  i  f ,  c  |,  p  •£-,  m  £  •  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 

^IIS^V  "^    especially     nar- 

row. Infra- 
orbital  foramen 
very  large. 
Head  rather 
small  and  pro- 
duced in  front, 
with  an  elon- 
gated, obliquely 
truncated,  naked 
snout.  Ears 
small.  Body 

elongated.  Limbs  short.  Tail  short  or  moderate,  bushy.  Several 
species  are  described  (H.  orientalis,  personata  [Fig.  265],  moscJmta, 
subaurantiaca),  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.  mientalis  also  ranging  into  Java ;  the  Chinese 
H.  subaurantiaca  is  brilliantly  coloured  in  the  region  of  the  throat.2 


FIG.  265. — Helictis  personata.    (From  Blanford,  Mammalia  of  British 
India,  p.  175.) 


FIG.  266. — Left  lateral  and  superior  aspect  of  the  brain  of  Helictis  sabaurantiam.    (From 
Garrod,  Proc.  Zool.  Soc.  1879,  p.  307.) 

The  brain  of  Helictis,  represented  in  the  accompanying  figure, 
shows  the  general  type  of  cerebral  structure  characteristic  of  the 
Mustelidce.  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.  2  Garrod,  ibid.  1879,  pi.  xxix. 


MUSTELID&  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  Mustela,  ceases  at 
the  superior  posterior  angle  of  the  hemisphere. 

Ictonyx.1 — Dentition  :  i  f ,  c  \,  p  f ,  ra  | ;  total  34.  In  general 
characters  the  teeth  much  resemble  those  of  the  Polecats  (Mustela), 
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,  /.  zorilla,  the  Cape 
Polecat,  was  placed  by  Cuvier  in  the  genus  Mustela,  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, 
/.  frenata,  from  Sennaar  and  Egypt,  has  been  described. 

Subfamily  Mustelinse. — 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. 

Galidis.- — Dentition:  i  f,  c  ^,  p  f,  m  |;  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 
Grison  (genus  Grisonia,  Gray),  and  G.  barlara,  the  Tayra  (genus 
Galera,  Gray),  both  South  American  ;  G.  allamandi  is  an  inter- 
mediate form. 

Remains  of  Galidis  occur  in  the  Pleistocene  cavern-deposits  of 
Brazil,  and  also  in  the  Pleistocene  of  North  America. 

M '/stela.3 — Dentition :  i  f,  c  |,  p  -|J,  m  \ ;  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.  Vertebrae :  C  7,  D  1 4,  L  6,  S  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). 


58o  CARNIVORA 


rounded  ;  toes  short,  with  compressed,  acute,  semiretractile  claws. 
Tail  moderate  or  long,  more  or  less  bushy. 

The  genus  Mustela,  as  restricted  by  Cuvier  (fiegne  AniinnL 
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  Mustela,  and  to  the 
second  assigned  that  of  Putorius.  Three  years  later  Nilsson  (Skand. 
Fauna,  1820)  definitely  constituted  the  two  groups  into  genera, 
applying  to  the  first  the  name  of  Maries,  by  which  the  animals 
composing  it  had  been  generally  designated  by  the  Latin-writing 
zoologists  of  the  preceding  century,  and  keeping  Mustela  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  Mustela  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 a  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  bullse  large,  but  not  very 
globose.  Mandible  with  a  strong  triangular  vertical  coronoid 
process  and  a  well -developed  angular  process.  Premolars  £. 
Upper  incisors  in  a  straight  transverse  line,  rather  long  and 

1  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. 


MUSTELID&  58 1 


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 
eggs,  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 
Muxtelidce.  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.1  In  the  same  way,  according  to  Hodgson,  the  Yellow- 
bellied  Weasel  (M.  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  (M.  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  (M.  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 

1  See  Rolleston,  "On  the  Domestic  Cats,  Fclis  domesticus  and  Mustela foina, 
of  Ancient  and  Modern  Times,"  Journal  of  Anatomy  and  Physiology,  vol.  ii.  p. 
47,  1868. 


MUSTELID^E 


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  : — 

M.  foina,  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 


FIG.  267.— The  Pine  Marten  (Mustela  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. 

M.  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  CARNIVORA 


common  in  most  parts  of  Great  Britain  and  Ireland.  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  regard  to  its  recent  occurrence  in  the  British  Isles, 
Mr.  Alston  writes  in  the  Proc.  Zool.  Soc.  1879  : — 

"Although  greatly  reduced  in  numbers  by  persecution,  it  still 
maintains  its  ground  in  the  wilder  districts  of  Scotland,  the  north 
of  England,  Wales,  and  Ireland;  and  occasionally  specimens  are 
killed  in  counties  where  the  species  was  thought  to  have  been  long 
extinct.  In  Scotland  it  is  still  found,  though  comparatively  rarely, 
in  the  Lews  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  fine  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  Lancashire,  and  in  Lincolnshire  several  have  been 
recorded,  the  latest  killed  in  1865,  by  Mr.  Cordeaux.  In  Norfolk 
one  was  shot  last  year;  and  I  have  myself  examined  a  fine 
example  which  was  shot  in  Hertfordshire,  within  20  miles  of 
London,  in  December  1872.  In  Dorsetshire  the  last  is  said  to 
have  been  killed  in  1804;  but  a  specimen  occurred  in  Hampshire 
about  forty  years  ago,  and  another  in  Surrey  in  1847.  In  Ireland 
the  following  counties  were  enumerated  by  Thompson  as  habitats 
of  this  species :  Donegal,  Londonderry,  Antrim,  Down,  Armagh, 
Fermanagh,  Longford,  Galway,  Tipperary,  Cork,  and  Kerry.  The 
Cat-crann  is  probably  now  a  rarer  animal  in  Ireland  than  it  was 
when  Thompson  wrote ;  but  it  still  exists  in  various  districts, 
especially  in  County  Kerry,  whence  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." 
&>  M.  zibellina,  the  Sable  (German,  Zobel  and  Zebel ;  Swedish, 
sabel ;  Russian,  sobel,  a  word  probably  of  Turanian  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  chiefly  in  Eastern  Siberia. 

M.  flamgula,  the  Indian  Marten. — Inhabits  the  southern  slopes 
of  the  Himalaya,  the  Nilgiri  Hills,  the  interior  of  Ceylon,  the 
Malay  Peninsula,  and  Java.  The  coloration  of  this  species  is  very 
striking,  the  upper  parts  being  blackish -brown,  and  the  throat 
and  breast  yellow  or  orange,  in  the  bright  coloured  variety.  It 
differs  from  the  other  species  in  having  the  soles  of  the  feet  more 
or  less  naked. 

M.  mdampus. — Japan. 


MUSTELIDsE  585 

M.  americana,  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,  from  October  to  April,  when  the  fur  is 
in  good  condition,  as  it  is  nearly  valueless  during  the  shedding  in 
summer.  Dr.  Coues  tells  us  that,  notwithstanding  the  persistent 
and  uninterrupted  destruction  to  which x  the  American  Sable  is 
subjected,  it  does  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  because  it  is  so  prolific,  bringing  forth  six  to  eight  young 
at  a  litter.  Its  home  is  sometimes  a  den  under  ground  or  beneath 
rocks,  but  oftener  the  hollow  of  a  tree,  and  it  is  said  frequently  to 
take  forcible  possession  of  a  squirrel's  nest,  driving  off  or  devouring 
the  rightful  proprietor. 

.]/.  pennanti,  the  Pekan  or  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  Fox  than  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. 
It  was  generally  distributed  in  wooded  districts  throughout  the 
greater  part  of  North  America,  as  far  north  as  Great  Slave  Lake, 
63°  N.  lat.,  and  Alaska,  and  extending  south  to  the  parallel  of  35°; 
but  at  the  present  time  it  is  almost  exterminated  in  the  settled  parts 
of  the  United  States  east  of  the  Mississippi. 

Fossil  remains  of  a  Marten  from  the  Pliocene  Siwaliks  of  India 
indicate  a  species  which  cannot  be  distinguished  from  those  now 
inhabiting  the  same  region  ;  while  remains  of  M.  martes  occur  in 
European  cavern-deposits,  and  in  the  fens  of  Cambridgeshire. 

With  the  Putoriine  group  (genus  Putorius)  we  come  to  those 


586  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.  lulreola,  the  Nors  or  Sump/otter  (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,  M.  sibirica,  from  Eastern  Asia,  of  which  much  less  is 
knoAvn,  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 


MUSTELID^E 


587 


on  the  under  parts  of  the  body.  In  very  rare  instances  the  tail  is 
tipped  Avith  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  Polecat  (Mustela  putorius). 

molluscs  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  Coues,  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  can  be  readily  tamed,  and  lately  Minks  have 
been  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 
(M.  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  comparatively  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  Blasius,  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  every- 
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  either  in  the  open  air,  in  holes, 
fox-earths,  rabbit-warrens,  under  rocks,  or  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  birds  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  early,  may  be  easily  trained,  like  Ferrets,  for  rabbit-catching. 
The  Polecat  is  very  tenacious  of  life,  and  will  bear  many  severe 
wounds  before  succumbing ;  it  is  also  said  to  receive  with  impunity 
the  bite  of  the  adder.  Its  fetid  smell  has  become  proverbial. 

Four  other  species  of  Polecats  are  known,  viz. — The  Siberian 
Polecat  (M.  eversmanni)  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  M.  larvata  is  distinguished  from  the  last 
by  the  presence  of  a  process  connecting  the  pterygoid  with  the 
auditory  bulla,  and  by  a  difference  in  the  shape  of  the  upper 
molar.  The  American  Polecat  (M.  nigripes),  inhabiting  the  central 
plateau  of  the  United  States,  and  extending  southwards  into  Texas, 
is  another  closely  allied  species,  although  some  zoologists  have  made 
it  the  type  of  the  genus  Cynomyonax.  Finally,  the  Mottled  Polecat 
(M.  sarmatica)  is  a  species  sparsely  distributed  in  Eastern  Europe 
and  parts  of  Western  Asia,  but  common  in  Southern  Afghanistan. 


MUSTELID&  589 


Its  skull,  although  smaller,  resembles  that  of  the  common  species; 
but  the  coloration  is  very  different,  all  the  upper  parts  being 
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  (M.  vulgar  is,  Fig.  269)  the  upper  parts,  outside  of 
limbs  and  tail,  are  a  uniform  reddish-brown,  the  under  parts  pure 


FIG.  269. — The  Common  Weasel  (Mustda  vulgar  is). 

white.  In  very  cold  regions,  both  in  Europe  and  America,  it  turns 
completely  white  in  winter,  but  less  regularly  and  at  a  lower 
temperature  than  the  Stoat,  from  which  it  is  easily  distinguished  by 
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  8  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  of  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  Avith  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  (M.  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  Mustela  in  its  wider  sense,  and  serve  to 
confirm  the  propriety  of  this  use  of  the  term.  Thus  M.  sedoria  is 
a  species  of  somewhat  larger  size  than  the  Stoat,  with  p  \,  while  in 
M.  angustifrons  the  number  of  premolars  is  -£,  and  in  M.  mustelina 
-£ ;  the  latter  species  agreeing  very  closely  in  size  with  the  Stoat. 
The  extinct  Plesictis,  in  which  there  are  p  -f-  and  the  lower  car- 
nassial  has  a  large  inner  cusp,  is  distinguished  from  Mustela  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  Mustela}  is  shorter 
in  an  antero-posterior  direction  than  the  secant  outer  moiety;  and 
the  auditory  bulla  is  more  inflated  than  in  Mustela,  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.  Mustela  larteti,  of  the  Middle 
Miocene  of  France,  should  perhaps  be  referred  to  Ictonyx. 

Pcedlogale.1 — This  genus  has  been  made  for  the  reception  of  the 
South  African  Mustela  albinucha,  in  which  the  coloration  is  similar 
to  that  of  Ictonyx,  but  the  number  of  cheek-teeth  is  usually  reduced 
to  p  % ,  m  $,  although  there  may  be  a  second  lower  molar.  The 
auditory  bulla  is  quite  flat. 

Lyncodon.2 — This  name  has  been  proposed  for  a  small  Musteline 
1  0.  Thomas,  Ann.  Mag.  Nat.  Hist.  ser.  5,  vol.  xi.  p.  370  (1883). 
2  Gervais,  Diet.  Univ.  cCHist.  Nat.  t.  iv.  p.  685  (1849). 


MUSTEL1D&  591 

from  Patagonia,  with  j>  H ,  m  },  which  Mr.  0.  Thomas  suggests 
may  be  nothing  more  than  an  aberrant  southern  form  of  Mustela 
( 1'nfurinx)  brasiliensis.  The  auditory  bulla  is  more  inflated  than  in  the 
typical  Weasels.  This  animal  is  somewhat  larger  than  the  Stoat. 

GWo^-^-Dentition :  i  -f ,  c  -£,  p  £,  m  i;  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  Hysena.  Palate 
prolonged  somewhat  behind  the  last  molar.  Humerus  with  an  ente- 
picondylar  foramen.  Vertebrae:  C  7,  D  15,  L  5,  S  3,  C  15.  Body 


Fio.  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,  Prodromiis  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  f.  The  cheek 
series  consists  generally  of  four  premolars  and  one  molar  of  very 
uniform  characters,  with  never  more  than  two  roots,  and  Avith 
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, 
Avith  numerous  and  complex  convolutions.  There  is  a  very  short 
caecum.  The  kidneys  are  divided  into  numerous  distinct  lobules. 
There  are  no  CoAvper's  glands.  The  mammae  are  either  tAvo  or 
four,  and  abdominal  in  position.  No  clavicles.  Tail  ahvays  very 
short.  Eyes  very  large  and  exposed,  Avith  flat  cornea. 

The  animals  of  this  group  are  all  aquatic  in  their  mode  of  life, 
spending  the  greater  part  of  their  time  in  the  Avater,  sAvimming  and 
diA'ing  Avith  great  facility,  feeding  mainly  on  fish,  crustaceans,  and 
other  marine  animals,  and  progressing  on  land  Avith  difficulty. 


OTARIIDA1  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  Otariida, 
Fur-Seals  or  Sea-Bears,  which  form  a  transition  from  the  Fissiped 
Carnivora  to  the  Seals  ;  the  Trichechida>,  containing  the  Walrus ;  and 
the  Pliocidce  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 Mivart1  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  Otariidce,  and  its  absence  in  Lutra 
and  the  Phocidce,  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  OTARIID.E. 

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.* — Dentition:  i  f,  c  \,  p  ^,  m  1-y-- ;  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 

1  Proc.  Zool.  Soc.  1885,  p.  497. 

2  Peron,  Voyage  aux  Tcrres  Australes,  vol.  ii.  p.  37  note  (1816). 
38 


594 


CARNIVORA 


FIG.  271.— Skull  of  Otaria  forsteri.    (From  Gray,  Proc.  Zool.  Soc. 
1872,  p.  660.) 


side  and  not  the  other) ;  all  Avith  similar  characters,  generally 
uniradicular ;  crown  moderate,  compressed,  pointed,  with  a  single 
principal  cusp,  and  sometimes  a  cingulum,  and  more  or  less  de- 
veloped anterior 
and  posterior 
accessory  cusps. 
Vertebra?:  C  7, 
D  15,  L  5,  S  4, 
C  9-14.  Head 
rounded.  Eyes 
large.  Pinna 
of  ear  small, 
narrow,  and 
pointed.  Xeck 
long.  Skin  of 
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 
elongated,  compressed,  and  curved. 

The  Eared-Seals,  commonly  called  Sea-Bears  or  Sea-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  seek  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  Seals. 
The  best-known  species  are  0.  stelleri,  the  Northern  Sea  Lion,  the 
largest  of  the  genus,  from  the  North  Pacific,  about  10  feet  in 
length  ;  0.  jubata,  the  Patagonian  or  Southern  Sea  Lion  (Fig.  272), 
from  the  Falkland  Islands  and  Patagonia ;  0.  californiana,  from 
California,  frequently  exhibited  alive  in  menageries  in  Europe ; 
0.  -ursiiia,  the  common  Sea-Bear  or  Fur-Seal  of  the  North  Pacific,  the 
skins  of  which  are  imported  in  immense  numbers  from  the  Prybiloff 
Islands ;  0.  pusilla,  from  the  Cape  of  Good  Hope  :  0.  forsteri  and 


OTARIWsE 


595 


others,  from  the  coasts  of  Australia  and  various  islands  scattered 
over  the  southern  hemisphere.  These  have  been  grouped  by  some 
zoologists  into  many  genera,  founded  upon  very  trivial  modifica- 
tions of  teeth  and  skull.  In  a  recent  memoir  Mr.  Beddard l  con- 
cludes that  if  the  genus  be  split  up  at  all,  it  should  be  divided  into 
Otaria,  containing  only  0.  jubata  (with  its  numerous  synonyms),  and 
Arctocephalus,  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 


FIG.  272. — The  Patagoniau  Sea-Lion  (Otaria  jubata).     From  Sclater,  Proc.  Zool.  Soc.  1866,  p.  80. 

a  hook-like  process  to  the  pterygoids,  and  by  the  posterior  border 
of  the  nasals  extending  behind  the  zygoma. 

The  following  account  of  0.  ursina  in  the  Prybiloff  Islands  is 
taken,  with  slight  verbal  alteration,  from  Nordenskiold's  Voyage  of 
the  Vega :  "  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  first  to  the  place,  most  of  them 
in  the  month  of  May  or  in  the  beginning  of  June.  The  most 
violent  conflicts,  often  Avith  a  deadly  issue  for  one  of  the  parties, 
now  arise  regarding  the  space  of  about  a  hundred  square  feet 
which  each  bull -seal  considers  necessary  for  his  home.  The 

1  "On  the  structure  of  Hooker's  Sea-Lion  (ArctocepJialus  hookeri),"  Trans. 
Zool.  Soc.  vol.  xii.  p.  369  (1890). 


596  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  water's  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  scarcely  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  1 2  to  1 5  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^E. 

In  many  characters  the  single  genus  representing  this  family 
is  intermediate  between  the  Otariidce  and  Phocidce,  but  it  has  a 
completely  aberrant  dentition.  It  has  no  external  ears,  as  in  the 


TRICHECHID^E  597 

Phocidce;  but  when  on  land  the  hind  feet  are  turned  forwards  and 
used  in  progression,  though  less  completely  than  in  the  Otari></n. 
The  upper  canines  are  developed  into  immense  tusks,  which  descend 
a  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. 

Tricheclms.1 — Dentition  of  young  :  i  f- ,  c  ^,  p  and  m  -f-.  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  ^,  c  ^,  p  f ,  m  % ;  total  1 8. 
Vertebrae :  C  7,  D  14,  L  6,  S  4,  C  12.  Head  round.  Eyes  rather 
small.  Muzzle  short  and  broad,  with  on  each  side  a  group  of  long, 


Fio.  273. — Diagram  of  the  dentition  of  the  Walrus  (Trichechus  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  Otaria  ;  first  and  fifth  with  minute 
flattened  nails ;  second,  third,  and  fourth  with  large,  elongated, 
subcompressed  pointed  nails. 

TricJiechus  is  the  almost  universally  accepted  generic  name  by 
which  the  Walrus  or  Morse  2  is  known  to  zoologists,  but  some  con- 
fusion has  been  introduced  into  literature  by  the  revival  of  the 
nearly  obsolete  terms  Rosmams  by  some  authors  and  Odobcenus  by 
others.  T.  rosmarux  is  the  name  of  the  species  met  with  in  the 
Arctic  seas  ;  that  of  the  North  Pacific,  if  distinct,  is  T.  obesus.  The 
preceding  and  following  descriptions  will  apply  equally  to  both. 

1  Linn.  Stjst.  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. 


CAKNIVORA 


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.  It  is  everywhere  more  or  less  wrinkled, 


FIG.  274. — The  Walrus  (Trichechiis  roamarus). 

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  Malmgren. 

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 


TRICHECHID&  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  Saxicava  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  algae  or 
sea-weeds  have  been  found  in  its  stomach,  but  whether  swallowed 
intentionally  or  not  is  still  doubtful. 

The  commercial  products  of  the  AValrus  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 
Avell  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 


6oo  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  is 
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  PHOCID^E. 

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  4  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 p  -\, 
m  y,  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  Phoeinae. — Incisors  f.  All  the  feet  with  five  well- 
developed  claws.  The  toes  on  the  hind  feet  subequal,  the  first  and 


PHOCID& 


6oi 


fifth   not  greatly   exceeding  the   others   in   length,   and  with   the 
interdigital  membrane  not  extending  beyond  the  toes. 

H(Hichcems.1 — Dentition  :  i  f ,  c  ^,  p  £,  in  ^ ;  total  34.     Crowns 
of   molars    large,  simple,    conical,    recurved,    slightly    compressed, 


Fio.  275. — Upper  permanent  and  deciduous  dentition  of  the  Greenland  Seal  (Phoca  gramlandlca). 
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.  Vertebrae  :  C 
7,  D  15,  L  5,  S  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. 
Vertebras :  C  7,  D 
15,  L  5,  S  4,  C 
12-15.  Head 
round  and  short. 
Fore  feet  short, 
with  five  very 
strong,  subcom- 
pressed,  slightly 
curved,  rather 
sharp  claws,  sub- 
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.  groen- 
Jiiiidica,  the  Greenland  Seal;  P.  vitulina,  the  Common  Seal  (Fig. 
277);  and  P.  hispida,  the  Kinged  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). 


Fio.  276. — Skull  of  Common  Seal,  showing  form  of  teeth. 


602  CARNIVORA 


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, 
yet  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  (Phoca  grcenlandica),  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  Scotch, 
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  year  round, 
but  some  have  a  regular  season  of  migration,  moving  south  in 


PHOCIDA^ 


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- 


Fio.  277. — The  Common  Seal  (Phoca  vitulina). 

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  Gammams, 
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  Sea-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. 


6o4  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  (Haliclmms  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  Monaehinae. — Incisors  f.  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. 

Moitachus.1 — Dentition  :  i  •§ ,  c  i,  p  £,  m  ^ ;  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.  Vertebrae:  C  7,  D  15,  L  5,  S  2,  C  11.  All  the 
1  Fleming,  Philosophy  of  Zoology,  vol.  ii.  p.  187  (1822). 


PHOCID^E  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 x  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.  leptonyx,  the  Sea-Leopard,  widely  distrib- 
uted in  the  Antarctic  and  southern  temperate  seas. 

Lobodon? — 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. 

PcecilopJwca.* — Cheek-teeth  small,  with  simple,  subcompressed, 
conical  crowns,  having  a  broad  cingulum,  but  no  distinct  accessory 
cusps.  One  species,  P.  weddelli. 

Ommafophoca.5 — 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  Cystophorinse. — Incisors  -f.  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.6 — Dentition  :  i  -f,  c  -J-,  p  £,  m  J- ;  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,  Monatsb.  K.  P.  Akad.  Wissenseh.  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). 

4  New  name,  Syn.  Leptonyx,  Gray,   Charlesworth's  May.  Nat.  Hist.  vol.  i.  p. 
582  (1837)  ;  preoccupied  by  Swainson,  1821. 

5  Gray,  Zoology  of  Erebus  and  Terror,  vol.  i.  p.  7  (1844). 

6  Nilsson,  Faun.  Scandinav.  vol.  i.  p.  382  (1820). 


606  CARN1VORA 


upper  part  of  the  head.  Nails  present,  though  small,  on  the  hind 
feet.  One  species,  C.  cridata,  the  Hooded  or  Bladder-Nose  Seal  of 
the  Polar  Seas. 

Macrorhinus.1 — Dentition  as  the  last,  but  cheek-teeth  of  simpler 
character,  and  all  one-rooted.  All  the  teeth,  except  the  canines, 
very  small  relatively  to  the  size  of  the  animal.  Hind  feet  without 
nails.  Vertebrae:  C  7,  D  15,  L  5,  S  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,  M.  leoninus,  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  Plwca  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  Callophoca  is  said  to 
be  allied  to  Phoca  grcenlandica,  Platyphoca  to  Plwca  barbata,  Phocanella 
to  Phoca  foetida,  Gryphoca  to  Haliclicerus,  Palceophoca  and  Monatherium 
to  MonacJms,  and  Mesotaria  to  CystopJwm ;  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  Pinnipcdia. — 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  Mim.  Acad. 
Roy.  d.  Bclgique. 

Suborder  CREODONTA. 

The  discovery  of  fossil  remains  in  Eocene  arid  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,  Mem.  du  Museum,  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 


FIG.  278.— Anterior  portion  of  the  skull  of  Ilycenodon  leptorhynchua.    (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  palaeontological 
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  extinct  Miacidce  (p.  539), 


608  CARNIVORA 


that  it  is  on  the  whole  advisable  to  regard  them  as  representing 
a  distinct  suborder  of  Carnivora.  In  the  strong  development  of  the 
canines  (Fig.  278)  they  are  distinguished  from  the  modern  Insect- 
ivora ;  and  they  also  differ  from  the  latter  and  resemble  the  true 
Carnivores  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  obtaining  in  most  of  the  modern  Carni- 
vora. Some  of  the  most  generalised  forms  included  in  the  present 
group  approximate  so  closely  to  the  Condylarthrous  Ungulates  as 
to  indicate  that  both  groups  have  probably  had  a  common  origin. 

The  Creodonta  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  of  the 
astragalus  usually  wants  the  groove  found  in  modern  Carnivores ; 
and  the  feet  were  plantigrade.  The  curious  resemblance  of  the 
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  of  the  angle  of  the  mandible, 
sharply  distinguishes  them  from  that  order.  Space  permits  of  a 
notice  only  of  the  more  interesting  forms. 

Hycenodontidce. — This  family  is  taken  to  include  some  of  the 
more  specialised  types,  such  as  the  European  and  American 
Hycenodon  and  Oxhycena  and  the  European  Pterodon.  In  Hycenodon 
(Fig.  278)  the  dental  formula  is  i  f ,  c  ^,  p  %,  m  -f ;  the  fourth 
premolar  above  and  the  first  true  molar  below  being  formed  upon 
the  "  carnassial "  plan,  but  the  teeth  behind  these,  instead  of  being 
tuberculated  as  in  all  existing  Carnivora,  repeat  the  characters  of 
the  carnassial,  and  also  increase  in  size,  especially  in  the  lower  jaw, 
from  before  L  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  together.  Some  species, 
as  H.  leptorhynchus,  were  as  large  as  a  Wolf,  while  others  did  not 
exceed  a  Fox  in  size.  Pterodon  is  readily  distinguished  by  having 
m  -f ,  by  the  larger  size  of  the  inner  tubercles  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.  Oxhyccna  is  a  specialised  form  with  i  ^p> 
c  \,  p  -f ,  m  f ,  and  a  very  long  mandibular  symphysis. 

Proviverridce. — The  European  and  American  genus  Proviverra 
(Cynohycenodon  or  Stypolophus)  may  be  regarded  as  representing  a 
second  family.  The  dental  formula  in  this  genus  is  the  typical  i  -|, 
c  T>  P  4~>  m  f  >  tne  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  Dasyurus,  while  the  lower  molars  are  like  the  lower 
carnassial  of  Cynodidis  and  Viverm;  and  thus  indicate  how  the 
Creodonts  may  have  passed  into  the  true  Carnivores  through  the 
extinct  Miacidce. 

Ardocyonidce  and  Mesonychidce, — The  first  of  these  families  is 
represented  by  Ardocyon  primcevus,  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  Eheims.  The  dental  formula  is  i  f ,  c  ^,  p  g^  m  f .  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  Flo  279._The  three  right  upper  molars  of  Arcto- 

Smaller.        In     the     American    cyan  dueli  (a),  and  the  second  of  A.gen-aisifb);  from 

Mesonm  the  dental  formula  the  Lowest  Eocene  of  Rheiins-  **>  Protocone ;  P*. 

y  .      .  ,  paracone  ;  me,  metacone  ;  hy,  hypocone ;  ml,  meta- 

WaS  the  typical  One,  the  jaWS    conule ;  j>Z,  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.  Mesonyx 
ossifragus  was  larger  than  a  Grizzly  Bear.  Amblydonus,  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    ORDEK    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 
testes  are  in- 
guinal 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  projec- 
tion of  the  muzzle  far  beyond  the  extremity  of  the  lower  jaw  is  a 
very  general  feature.  The  humerus  generally  has  an  entepicondylar 


-  FIG.  280. — Right  lateral  aspect  of  the  anterior  portion  of  the 
cranium  of  Erinaceus  collaris.  Enlarged.  (From  Dobson,  Proc.  Zool. 
Soc.  1881,  p  403.) 


INSECTIVORA 


611 


foramen.  Certain  forms,  such  as  Talpa  and  Galeopithecm,  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  Tupaiidce,  all  are  noc- 
turnal ;  the  greater  number  are  cursorial, 
but  some  (Talpa,  Chrysochlmis,  Oryzorictes) 
are  fossorial ;  some  (Potamogale,  Necto- 
gale,  Myogale)  are  natatorial,  and  a  few 
(Tupaiidce)  arboreal ;  while  the  species 
of  the  aberrant  genus  Galeopithecm  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  FIG.  28i.  -  Upper  surface  of  the 

,  .      ,     .        ,  ,      ,         brain  of  fupaia  ferruginea.     (From 

character  ot  their  placentation  and  the  oarrod,  Proc.  zooi.  soc.  1879,  p.  304.) 
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  Galeopithecidce,  Tupaiidce, 
and  Macroscelididce.  The  number  of  dorsal  vertebra  varies  from  1 3 
in  Talpa  to  19  in  Centetes;  that  of  the  lumbar  from  3  in  Chryso- 


612  INSECTIVORA 


chloris  to  6  in  Talpa  and  Sorex ;  and  of  the  caudal  from  the  rudi- 
mentary series  of  8  in  Centetes  to  the  40  or  more  of  Microgale.  Xot 
less  variable  are  the  characters  of  the  vertebrae  themselves ;  the 
spinous  processes  often  being  very  long  in  one  and  short  in  another 
species  of  the  same  genus.  In  the  Soricidce  and  Myogale  the  neural 
arches  of  the  cervical  vertebrae  are  very  slender.  In  the  Soricidce 
and  Gymnura  the  four  anterior  vertebrae  develop  large  single  hypa- 
pophyses.  In  Galeopithecus  the  centrum  of  each  vertebra  supports 
posteriorly  a  pair  of  intercentral  ossifications ;  while  in  Erinaceus, 
Myogale,  and  Talpa  small  oval  ossicles  are  found  on  the  inferior 
surfaces  of  the  lumbar  interspaces.  In  Erinaceus,  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  vertebrae  greatly  exceeds  that  of  any  of  the  succeed- 
ing vertebrae.  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  Talpa,  having  relation  to  the  use  of  the  fore  limbs  in 
burrowing;  but  in  the  Golden  Moles  (Chrysochloris)  the  forearm 
and  manus  alone  become  specially  modified  for  this  purpose.  In 
Galeopithecus  and  Macroscelides  the  bones  of  the  forearm  (radius 
and  ulna)  are  distally  united.  The  manus  has  generally  five  digits, 
but  in  Rhynchocyon  and  in  one  species  of  Oryzorictes  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  Galeopithecus,  Tupaia, 
Centetes,  Hemicentetes,  Ericulus,  and  Solenodon  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  (Macroscelides,  Rhynchocyon},  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  of  the 
families.  Thus  in  the  Galeopithecidce,  Tupaiidce,  and  Macroscelididce 
there  is  a  long  symphysis ;  in  the  Erinaceidce,  Centetidce,  and  Potamo- 
galidce  the  symphysis  is  short;  and  in  the  Soiicidce,  Talpidce,  and 
Chrysochloiidce  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  Monograph,  referred  to  in  the  bibliography.  As  to 
the  nervous  system,  it  has  been  already  mentioned  that  the  brain 
throughout  the  order  presents  a  low  type  of  organisation ;  in  none 
of  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 
(Erinacem}  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  Galeopithecus  the  stomach  is  a  simple,  thin-walled  sac ; 
but  in  some,  as  in  Centetes  and  allied  genera,  the  pyloric  and 
oesophageal  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  Tupaia, 
as  well  as  in  the  Macroscelididce,  all  of  which  probably  feed  in 
part  on  vegetable  substances,  most  of  the  species  possess  a  caecum. 
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  (Soriddce,  Centetidce,  Chryso- 
chlwidce)  the  penis  is  capable  of  being  more  or  less  completely 
retracted  within  the  fold  of  integument  surrounding  the  anus ;  in 
some  (Galeopitheddce,  Talpidce)  it  is  pendent  in  front  of  the  anus  ; 
while  in  others  (Macroscelididce,  Erinaceidce,  Solenodontidce)  it  is 
carried  forwards  and  suspended  from  the  abdominal  wall.  In  the 
subfamily  Centetince  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  Rhynchocyon  the 
uterine  cornua  are  long  and  open  into  a  short  corpus  uteri,  which 
in  many  species  (Soriddce,  Talpidce,  Centetida\  Chrysochloridce)  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 
Erinaceus,  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  Chrysochloiis  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  IXSECTIVORA 


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  Potanwgale  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  Avithin  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 (Erinaceidce)  than  in  any  other  mammal.  In  this  family 
and  the  Centetidce  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  mammae  are 
axillary,  the  uterus  is  two-horned,  and  there  is  a  large  caecum. 

Galeopithecus J — Dentition  :  i  -| ,  c  \,  p  f ,  m  f ;  total  34.  Second 
upper  incisor  and  canine  with  two  roots.  Two  species — G.  wlans 
and  G.  phiUppinensis.  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,  Ada  Acad.  Sci.  Imp.  Pctropolis,  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,  involv- 
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 


PIG.  282. — Feet  of  Gakopithecus  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. 

Gakopithecus  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  caecum,  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  Gakopithecus  Dr.  Leche  l  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  Sdugethiergattung  Galcopithecus.     Sv.  Ale.  Handl.  vol.  xxi.  pt.  xi. 

(1886).     . 


616  INSECTIVORA 


line  of  descent.  The  deeply  pectinated  crowns  of  the  lower  incisors 
of  Galeopithecus  are  quite  unique  in  the  class,  and  the  only  approach 
to  the  double-rooted  canine,  except  in  Erinaceus  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 

TV-shaped  crowns. 

A.  Syruphysis  pubis  long  ;  generally  a  caecum  ;  cerebral  cavity 

comparatively  large. 

a.  Orbit  encircled  by  bone  ;    metatarsus  moderate  ;   arboreal. 

Tupaiidce. 

b.  Orbit  not  encircled  by  bone  ;  metatarsus  greatly  elongated  ; 

terrestrial     Macroscelididce. 

B.  Symphysis  pubis  short  or  none ;  no  caecum  ;  cerebral  cavity 

small ;  skull  without  postorbital  processes. 

a.  First  and  second  upper  molars  with  a  central  fifth  cusp. 
a.  Tympanic  annular,  not  forming  a  biilla.     Erinaceidce. 

b.  No  central  fifth  cusp  to  upper  molars. 

a'.  Tympanic  annular,  not  forming  a  bulla  ;  no  zygomatic 

arch.     Soriddce. 
b'.  Tympanic  forming  a  bulla ;   zygomatic  arch  developed. 

Talpidce. 

II.  Upper  molars  narrow,  with  V-shaped  crowns. 

a!  Tympanic  annular,  not  forming  a  bulla ;  zygomatic  arch 

imperfect. 

a".  No  clavicles.     Potamogalidce,. 
b".  Clavicles  well  developed. 

a'".  Skull  constricted  between  the  orbits  ;  penis  sus- 
pended.    Solenodontidce. 
b'".  Skull   not   constricted ;   penis    pendent,  retractile. 

Centetidce. 

b'.  Tympanic  forming  a  bulla  ;  zygomatic  arch  well  developed. 
Chrysochloridce. 

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  Palsearctic  and  Oriental  regions.  None  of  the 


TUPAIID& 


6i7 


existing  families  of  the  second  section  are  known  in  a  fossil  condition, 
although  it  is  suggested  that  the  extinct  Leptictidce  includes  allied 
types. 

Family  TUPAIID^;. 

Skull  with  comparatively  large  brain-case,  orbit  surrounded  by 
bone,  well-developed  zygomatic  arch,  perforated  jugal,  and  a  tympanic 


Fia.  283.—  The  Pentailed  Tree-Shrew  (Ptilocercus  lowi).    From  Gray,  Proc.  Zool.  Soc.  1848. 

J  natural  size. 

bulla.  Upper  molar  broad,  with  cusps  arranged  in  a  W.  Pubic 
symphysis  long ;  radius  and  ulna,  and  tibia  and  fibula  separate  ; 
metatarsus  only  slightly  longer  than  tarsus.  Usually  a  short  caecum. 
Habits  arboreal  and  diurnal.  Confined  to  the  Oriental  region. 

Tupaia.1 — Dentition  :  i  f ,  c  ^,  p  f ,  m  f ;  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  Raffles,  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. 

Ptilocercus.1 — 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  with  a  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 
Tupaia.  The  genus  Parasorex,  from  strata  of  similar  age,  has  the 
dental  formula  i  -f,  c  ^,  p  -f ,  m  -f,  and  is  regarded  as  connecting  the 
present  with  the  following  family. 


Family  MACROSCELIDID^E. 

Skull  with  comparatively  large  brain -case,  strong  zygomatic 
arch,  a  tympanic  bulla,  orbit  surrounded  by  bone,  im  perforate 
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  caecum.  Habits 
terrestrial,  saltatorial,  and  nocturnal.  The  family  is  confined  to 
Africa. 

Macroscelides.2 — Dentition  :  i  f,  c  ^,  p  £,  m  —3 ;  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.3  M.  tetradadylus  (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. 

fihynchocyon* — Dentition :  i  %,  c  ^,  p  •£-,  m  -| ;  total  36.  Upper 
incisor  frequently  shed  in  the  adult.  Radius  and  ulna  distinct ; 

1  Gray,  Proc.  Zool.  Soc.  1848,  p.  23.  2  Andrew  Smith,  S.  African  Qiutrt. 

Journ.  vol.  ii.  No.  1,  p.  64  (1833). 

3  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.  prcuss.  Ak.  Wiss.  1847,  p.  36. 


ERINACEID& 


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 


Fio.  234.  —  Macroscelides  tetradactylus.     x|.    (From  Peters,  Reise  iiach  Mossambique.) 

species  measures  about   8  inches  in  length  ;  and  the  long  tail  is 
covered  with  a  ringed  skin,  sparsely  haired.    Its  habits  are  fossorial. 


Family  ERINACEID^E. 

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  csecum  ;  penis  carried  forward  and  sus- 
pended from  the  wall  of  the  abdomen.  Habits  terrestrial.  Found 
in  the  Palsearctic,  Ethiopian,  and  Oriental  regions. 

Subfamily  Gymnurinae.  —  Palate  completely  ossified;  pelvis 
very  narrow  ;  fur  without  spines. 

Gymnura.1  —  Dentition  :  i  f  ,  c  \,  p  ^,  m  f  ;  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.  rafflesi  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  Hijlomys  suillus,  and  classed 
with  the  Tupaiidce.  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    Erinaeeinae. — Palate   imperfectly   ossified;    pelvis 
wide  ;  fur  with  spines. 

Erinaceus.1 — Dentition  :  i  f ,  c  $,  p  f ,  m  f ;  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.  europceus  (Fig. 
285),  each  of  these 
teeth  is  inserted  by 
two  distinct  roots 
(Fig.  280,  p.  610). 
The  first  lower  in- 
cisor is  large  and 
proclivous.  The 


Pio.  285. — Right  lateral  aspect  of  the  anterior  portion  of  the 
skull  of  the  Hedgehog  (Erinaceus  europceus).  Enlarged.  (From 
Dobson,  Proc.  Zool.  Soc.  1881,  p.  403.) 


number  of  vertebrae  is  C  7,  D  15,  L  6,  S  3,  C  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  (E.  europceus)  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,  E.  collaris  (Fig.  280),  may  be  considered  characteristic 
of  all  the  other  species,  the  only  important  differences  being  found 
1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  75  (1766). 


SORICW^E  621 

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  E.  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  E.  europceus  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,  Palceoerin- 
aceus,  appears  to  be  allied  to  Erinaceus,  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  Palceoerinaceus  with 
Erinaceus.  Of  these  Necrogymnurus,1  with  which  Cayluxotherium  is 
apparently  identical,  has  teeth  like  Gymnura,  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  Gymnura. 


Family  SORICHXE. 

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  caecum. 
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, 

1  Originally  given  incorrectly  as  Neurogymnurus. 


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 

Myoswex  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 
FIG.  286. -Left  lateral  view  of  the  difficult  group  have  been  attempted, 

cranium  and  mandible  of  Sorex  vercepacis.  -,  j  •  ji        i   o. 

In  the  cranium- i,  first  incisor  ;c,  fourth  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,  dlS- 
incisor;  p,  fourth  premolar;  m,  first  molar.  .  .  .  . 

(From  Alston,  Proc.  Zooi.  Soc.  1877.)  tmguished  by  the  apparently  trivial 

character  of  the  colour  of  the  teeth. 

Subfamily  Sorieinse. — Summits  of  the  teeth  coloured  red. 

Sorex.2 — Dentition  :  i  4,  c  £,  p  f,  m  -f ;  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 
Palaearctic  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.  3  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  73  (1766). 


SORICIDsE  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  (S,  pygmaus l}  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 


FIG.  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  Microsorex),   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  Neosorex,  are 
aquatic,  and  thus  take  the  place  of  the  Old  World  genus  Crossopus. 
These  are  S.  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.  pygmceus. 

Soriculus.1 — Dentition  :  i  4,  c  £,  p  -j— ,  m  f ;  total  30,  or  rarely 
32.  Opening  of  male  or  female  generative  organs  forming  with  the 
anal  orifice  a  shallow  cloaca.  Ear  and  tail  as  in  Sorex.  First  upper 
incisor  with  an  internal  cusp.  Habits  terrestrial. 

This  genus  is  the  only  representative  in  the  Oriental  region  of 
the  Soricince,  which  are  otherwise  confined  to  the  Palsearctic  and 
Nearctic  regions.  The  Indian  and  Burmese  species  comprise 
S.  nigrescens,  S.  cauctftfus,  and  S.  macrurus. 

Notiosorex? — Dentition  :  i  f ,  c  £,  p  ^,  m  f ;  total  28.  Tail 
moderate ;  first  upper  incisor  without  an  inner  cusp ;  other 
characters  as  in  S&riculus.  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. 

Marina.3 — Dentition  :  i  --~- ,  c  £,  p  -f- ,  m  f ;  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.  mexicana  and  micrura  extend  the 
range  of  the  genus  into  Mexico  and  Guatemala.  The  following 
account  of  the  habits  of  B.  brevicaudd  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).  3  Gray,  Proc.  Zool.  Soc. 

1837,  p.  124. 


SORICID&  625 


the  chrysalides  and  larvae  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.1 — Dentition  :  i  f,  c  £,  p  f,  m  f ;  total  30.  Opening 
of  male  or  female  generative  organs  enclosed  within  the  same  ring 
as  the  anal  orifice ;  penis  broad,  with  lateral  processes.  Ears  small, 
not  truncated.  Tail  long,  with  an  inferior  fringe  of  elongated 
hair ;  feet  also  fringed.  Habits  aquatic.  The  Palaearctic  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  larvae,  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  Croeidurinse. — Teeth  completely  white. 

Myosorex? — Dentition:  i  f ,  c  ^,  p  ^,  m  f ;  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,  Isis,  1832,  p.  275.  2  Gray,  Proc.  Zool.  Soc.  1837,  p.  124. 

40 


626  INSECTIVORA 


Shrew  from  the  Cape,  which  is  quite  unique  among  the  whole 
family  in  having  a  rudimental  seventh  pair  of  lower  teeth. 

Crocidura.1 — Dentition  :  i  f ,  c  £,  p  -^>  m  f ;  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  Myosorex.  Habits  terrestrial. 

This  Old  World  genus  includes  over  seventy  nominal  species, 
which  have  been  divided  into  four  subgenera,  C.  aranea  and  C. 
suaveolens  of  Continental  Europe,  and  C.  ccerulea  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.  ccerulea, 
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  short  distance  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. 

Diplomesodon.2 — Dentition :  i  f ,  c  -J-,  p  \,  m  -f  ;  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. 

Anurosorex.3 — Dentition :  i  % ,  c  -J-,  p  y,  m  § ;  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  A.  squamipes  occurs  in  Tibet,  while  A.  assamensis 
is  found  in  Assam.  The  latter  species  has  the  longer  tail.  The 
habits  of  both  are  probably  fossorial. 

Chimarrogale.4 — Dentition  :  i  f ,  c  -J-,  p  \,  m  -f ;  total  28.  Penis 
broad,  with  lateral  processes ;  male  or  female  generative  organs 
opening  within  the  same  integumentary  ring  as  the  anal  orifice. 

1  Wagler,  Isis,  1832,  p.  275.  2  Brandt,  in  Lehmann's  Reise,  -Zool.  Anh. 

p.  299  (1852).  3  Milne-Edwards,  Comptes  fiendus,  vol.  Ixx.  p.  341  (1870). 

4  Anderson,  Journ.  As.  Soc.  Bengal,  vol.  xlvi.  p.  262  (1877). 


SORICID^E 


627 


Tail  long,  with  an  inferior  fringe  of  elongated  hairs ;  ears  small ; 
plantar  callosities  simple  ;  toes  free.  Habits  aquatic. 

This  genus  includes  C.  himalayica  of  the  Himalaya  and  C.  platy- 
cephalus  of  Japan.  Both  have  the  feet  fringed,  and,  together  with 
the  next  genus,  may  be  regarded  as  the  eastern  analogues  of  Crosso- 
pus  among  the  red-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.1 — Dentition  :  i  f ,  c  ^,  p  ^,  m  f ;  total  28.     External 


FIG.  288. — Nectogale  elegans.    (From  Milne-Edwards,  Mammif.  Tibet.) 

ears  not  forming  a  conch,  valvular.  Plantar  callosities  forming 
adhesive  pads ;  toes  webbed.  Other  characters  as  in  Chimarrogale. 
Habits  aquatic. 

The  sole  representative  of  this  genus  is  the  Tibetan  Water- 
Shrew  (N.  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 
beds  of  the  streams  it  inhabits.  This  species  is  probably  more 
completely  aquatic  in  its  habits  than  the  allied  Chimarrogale. 

Fossil  Soricidce. — Eemains  of  existing  species  of  Sorex  or  Crasso- 
pus  occur  in  the  Norfolk  Forest  bed,  while  an  extinct  species  has 
been  found  in  the  Pleistocene  of  Sardinia.  Crocidura  occurs  in  the 
cavern-deposits  of  Madras.  Shrews  from  the  Miocene  and  Upper 

1  Milne-Edwards,  Comptes  Rendus,  vol.  Ixx.  p.  341  (1870). 


628  INSECTIVORA 


Eocene  of  Europe  have  been  referred  to  Swex  and  the  genus  Amphi- 
sorex,  which  is  a  synonym  of  Crossopus. 


Family  TALPID^E. 

Allied  to  the  Soricidce,  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  caecum ;  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  Soricidce  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 
Scaptonyx  Dr.  Dobson,  in  the  second  part  of  his  Monograph  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  : — 

MYOGALJE — Myogale. 
COND  YLUR.E —  Condylura. 


TALP^E — Talpa. 


Urotrichus. 
UROPSILI —  Uropsilus. 

Subfamily  Myogalinae. — Clavicles  and  humerus  moderately 
elongated ;  manus  without  falciform  bone. 

Myogale}  —  Dentition  :  i  f ,  c  ^,  p  £,  m  f ;  total  44.  Feet 
webbed.  Habits  aquatic.  This  genus  is  represented  by  the  two 
species  M.  moschata  (Fig.  289)  and  M.  pyrenaica,  of  which  the  former 
is  by  far  the  largest  member  of  the  family,  its  total  length  being 
about  1 6  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.  M.  pyrenaica, 

1  Cuvier,  "Tabl.  de  Classif."  in  Le^vns  d'Anat.  Compar.  vol.  i.  (1800). 


TALPIDA-: 


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  M.  moschata  occur  in  the  Norfolk  Forest  bed,  and 
were  originally  described  under  the  name  of  Palceospalax.  The 
genus  is  also  represented  in  the  Middle  and  Lower  Miocene  of  the 
Continent. 

Urotrichus.1 — Dentition :  i  f,  c  y,  p  £  or  f ,  m  f ;  total  36.    Feet 
not  webbed;  manus  broad.     Habits  fossorial.     The  Mole-Shrews, 


FIG.  289.— The  Desman  (Myogale  moschata).    J  natural  size. 

as  these  animals  are  called,  are  represented  by  U.  talpoides  of  the 
mountains  of  Japan  and  U.  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  f . 

Uropsilus.2 — Dentition :  if,  c  T,  p  f ,  m  f ;  total  34.  Manus 
narrow;  tail  naked  and  scaly.  Habits  cursorial.  The  single 
species,  U,  soricipes,  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,  Fauna  Japonica,  vol.  i.  p.  22  (1842).  -  Milne-Edwards, 

Arch,  du  Ahiseum,  vol.  vii.  Bull.  p.  92  (1872). 


630  1XSECTIVORA 


Subfamily  Talpinse. — 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.1 — Dentition  :  i  f ,  c  £,  p  f ,  m  f ;  total  36.  Extremity 
of  muzzle  simple ;  hind  feet  webbed ;  tail  short  and  nearly  naked. 
Represented  by  three  species  in  the  United  States. 

Scapanus.2 — Dentition  :  i  % ,  c  \,  p  -|,  m  -§- ;  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? — Dentition :  i  f ,  c  -£,  p  £,  m  f ;  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  :  if,  c  ^,  p  •£-,  m  f ;  total  42.  Manus 
moderately  broad,  as  in  Urotrichus.  Represented  only  by  S.ftisi- 
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.5 — Dentition  (usually)  :  if,  c  -^  p  £,  m  f ;  total  44. 
Manus  extremely  broad. 

This  genus  includes  the  true  Moles,  of  which  the  common 
English  Mole 6  (T.  europcea)  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- 

1  Cuvier,  "  Tabl.  de  Classif."  in  Le$on d'Anat.  Comp.  vol.  i.  (1800).  2  Pomel, 
Arch.  Sci.  Phys.  Nat.  vol.  ix.  p.  247  (1848).  3  Illiger,  Prodrmmis  Syst.  Mamm. 
et  Avium.  p.  125  (1811).  4  Milne-Edwards,  N.  Arch,  du  Museum,  vol.  vii. 

Bull.  .p.  92  (1872).        5  Linn.  Syst.  Nat.  12th  ed.  p.  73  (1766).       6  The  following 
account  is  taken  almost  entirely  from  Dr.  Dobson. 


TALPID&  63 1 

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 


INSECTIVORA 


the  young — generally  from  four  to 


PIG.  290.  —  Skeleton  of  Mole  x  |  (lower  jaw 
removed  to  show  base  of  skull),  c,  Calcanetim ; 
c.h.,  clavicular  articulation  of  the  humerus ;  cl., 
clavicle;  «.c,  external  condyle  of  humerus;  /., 
femur ;  fb,  fibula ;  /c,  falciform  bone  (radial  sesa- 
moid) ;  h,  humerus ;  i.c,  internal  condyle  of 
humerus ;  il,  left  ilium ;  i.p,  ramus  of  the  ilium 
and  pubis ;  is.,  ischium ;  l.d,  ridge  of  insertion  of 
latissimus  dorsi  muscle  ;  l.t,  lesser  trochanter  ;  m, 
manubrium  sterni ;  o,  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  falciforme)  in  the  manus;  sc.,  scapula;  s.h., 
scapular  articulation  of  the  humerus ;  t,  tibia ;  u, 
ulna. 


six  in  number — which  are 
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- 


TALPID&  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  vertebrae  are  usually  C  7,  D  13,  L  6,  S  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  Pala3arctic  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. :  i  % ,  c  fa  p  £, 
m  f ,  T.  wogura ;  i  f ,  c  \,  p  £,  m  f ,  T.  europcea,  cceca,  longirostris, 
micrura  ;  i  f ,  c  ^,  p  f ,  m  f ,  T.  leucura,  leptura ;  i  f ,  c  ^,  p  f ,  m  f, 
1.  moschata. 

Except  in  T.  europcea,  the  eyes  are  covered  by  a  membrane.  In 
T.  micrura  the  short  tail  is  concealed  by  the  fur.  T.  cceca  is  found 
south  of  the  Alps ;  the  remaining  species  are  Asiatic,  and  two  only 
— T.  micrura  and  T.  leucura — occur  south  of  the  Himalaya.  T. 
moschata,  of  Tibet,  is  regarded  by  some  zoologists  as  generically 
distinct  under  the  name  of  Scaptochirus. 

Remains  of  T.  europcea  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 


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  Tetracus  is  said  to  have 
affinity  both  with  Myogale  and  Erinaceus ;  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  Myogalince.  The  minute 
Amphidozotherium,  of  the  French  Phosphorites,  is  considered  to  be 
allied  to  Urotrichus. 

Family  ADAPISORICID^E 

This  extinct  family  is  represented  by  the  genera  Adapisorex  and 
Adapisoriculus,  of  the  lowest  Eocene  of  Rheims,  which  are  regarded 
as  allied  to  the  Soricidce,  but  somewhat  more  specialised.     In  the 
type  genus  the  formula  of  the  lower  teeth  is  i  2,  c  1,  p  4,  m  3 ; 
the  incisors  and  canine  being  proclivous,  and  the  molars  (of  which 
the   last   is   small   and   without   a   third   lobe)    quadritubercular. 
Adapisoricultis  is  a  smaller  form  with  differently  shaped  molars. 
Here  also  may  be  mentioned  the  genera  Orthaspidothenum  and 
Pleuraspidotherium,    from   the    above- 
mentioned  deposits,  which  are  prob- 
ably members  of  the  present  order. 
They  appear  to  have  been   animals 
somewhat  smaller  than  a  Hedgehog, 
Fio.  291.-The  last  left  upper  cheek-        •  -,     nnarlriti]hprriilar    nnnpr    rrmlars 
teeth  of  Pkuraspidotherium  oumonferi;    ^ 

from  the  Lowest  Eocene  of  Rheims.  pr,   (Fig.  291),  and  the  hinder  premolars 

protocone;  w,  metacone ;  jxi,  paracone ;  mOre  Complex  than  those  of  the 
6,  cingulum-cusp.  (From  Osborn.)  ^  .  .7  T  ,,  ,.  -, 

bnnacewce.    In  the  first-named  genus 

the  dental  formula  is  i  -f ,  c  -£-,  p  £,  m  f ;  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  POTAMOGALID^E. 

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 ; 


SOLENODONTID^E  635 

tibia  and  fibula  typically  united  distally.     No  caecum.     Confined  to 
the  Ethiopian  region. 

Potamogale.1 — Dentition :  i  f ,  c  ^,  p  -f ,  in  |  ;  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 


FIG.  292. — Potamogale  velox.     x  J.    (From  Allman,  Trails.  Zool.  Soc.  vol.  vi.  pi.  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.2 — Dentition  :  i  f,  c  ^,  p  f ,  m  | ;  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  is  not 
known  whether  a  clavicle  exists  ;  and  the  material  at  present  avail- 
able is  insufficient  to  definitely  fix  the  natural  position  of  the  genus. 


Family  SOLENODONTID^E. 

Skull  with  a  small  brain-case  constricted  between  the  orbits,  no 

1  Du  Chaillu,  Proc.  Boston  Soc.  Hist.  Nat.  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.  Vertebrae  :  C  7,  D  15,  L  4,  S  5,  C  23.  No  caecum.  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  caecal  sacs. 

Solenodon.1 — Dentition  :  i  f ,  c  -f ,  p  f ,  m  -f  ;  total  40.     This  genus, 
with  S.  paradoxus  and  S.  cubanus  (Fig.  293),  from  Hayti  and  Cuba 


Fio.  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  mammas  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  Talpidce,  the  form  of  the  crowns  of  the  molar  teeth 
connects  them  with  the  next  family. 

1  Brandt,  Mim.  Ac.  Imp.  St.  Pttersbourg,  1833,  vol.  ii.  p.  459. 


CENTETID^E 


637 


Fio.  294. — Left  lateral  view  of  the  skull  of  the  Tenrec  (Centetes 
ecaudatus).    Reduced. 


Family  CENTETID^E. 

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  caecum.  The 

penis  is  pendent  and  retractile  within  the  fold  of  the  integument 
surrounding  the  anus ;  the  testes  are  abdominal ;  the  mammae  are 
thoracic  and  ventral;  and  the  uterine  cornua  are  terminated  by 
the  Fallopian  tubes.  All  the  species  are  limited  to  Madagascar. 

Subfamily  Centetinse. — Tibia  and  fibula  distinct ;  testes  near 
kidneys ;  fur  with  spines. 

Centetes.1 — Dentition  r  i  f,  c  \,  p  f,  m  f ;  total  38.  Vertebrae  : 
C  7,  D  19,  L  5,  S  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 :  i  f ,  c  ^,  p  -f ,  m  -f ;  total  40.  This 
genus  is  represented  by  the  two  species  H.  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.  ecaudatus,  and  the  dorsal  spines  are  retained  in  the 
adult  state.  Vertebrae  :  C  7,  D  16,  L  5,  S  3,  C  9. 


1  Illiger,  Prodromus  Syst.  Mamm.  ct  Atrium,  p.  124  (1811). 
Proc.  Zool.  Soc.  1871,  p.  72. 


Mivart, 


638  INSECTIVORA 


JEriculus.1 — Dentition  :  i  f ,  c  ±,  p  f ,  m  f ;  total  36.  Vertebrae  : 
C  7,  D  17,  L  6,  S  4,  C  9.  The  single  species,  E.  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  Oryzorietinse. — Tibia  and  fibula  united ;  testes  near 
urethra  ;  fur  without  spines. 

Microgale.2 — Dentition  :  i  f,  c  ^,p  f,  m  f ;  total  40.     This  genus 


'  "^•"        "^""-MT'T  T*jg^_JV*^^'  ^*"^'^ 

FIG.  295.— Skull  of  Hemicentetes  semisplnosus.     x  2.    (From  Mivart,  Proc.  Zool.  Soc.  1871.) 

includes  M.  longicaudata  and  M.  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  vertebrae ;  teeth  like  those  of 
Centetes  ecaudatus,  but,  owing  to  the  comparatively  much  shorter 
muzzle,  not  separated  by  wide  spaces,  and  the  last  premolar  and 
molar  with  internal  basal  processes. 

Oryzorictes.3 — 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^E. 

Skull  conical,  not  constricted  between  the  orbits,  without  post- 
orbital  process,  but  with  well-developed  zygomatic  arch  and  tympanic 

1  I.  Geoffroy,  Ann.  Sci.  Nat.  ser.  2,  vol.  viii.  p.  60  (1837).  2  Thomas, 

Journ.  Linn.  Soc. — Zool.  vol.  xvi.  p.  319  (1882).  3  Grandidier,  Rev.  and 

Mag.  Zool.  1870,  p.  50. 


CHRYSOCHLORID&  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  Centetince ;  the 
mammae  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  Centetidce,  occupying  the 
same  relative  position  with  respect  to  that  family  that  the  Talpidce 
does  to  the  Soricidce.  Compared  with  the  Talpidce,  we  find  the 
following  differences  in  the  structural  adaptation  to  a  fossorial  life ; 
the  manubrium  sterni  is  not  anteriorly  elongated,  neither  are  the 


FIG.  296. — The  Golden  Mole  (Chrysochloris  oltusirostris). 

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.1 — Dentition  :  i  f ,  c  ^,  p  f ,  m  ~  ;  total  40  or  36 
Vertebra? :  C  7,  D  1 9,  L  3,  S  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  -f ,  with  a  basal  talon  to  the  lower  ones,  and  without  a  prominence 
in  the  temporal  fossa,  have  been  placed  in  a  separate  genus, 
Chakochloris,  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  Lacepede,  Mem.  de  I'lnstitut,  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 
Leptictidce,  of  the  Eocene  and  Miocene  of  North  America.  This 
family  includes  the  genera  Leptictis,  Mesodectes,  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,  Beise  nach  Mossambique — Sdugeth.  1852; 
Id.  "Ueber  die  Classification  der  Insectivora,"  Monatsb.  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.  Geol.  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. 


CHAPTEE    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  i  f ,  c  ^,  p  f ,  m  ^ ;  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. 

Linnaeus  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  mammae,  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  mammae 
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  vertebrae  differ 
very  slightly  in  number  and  form  throughout  the  species.  The 
general  number  of  the  dorso-lumbar  vertebrae  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  Ptcropodidce,  the  vertebrae,  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  vertebras  are  but  slightly  movable  upon  each  other,  and  in 
old  individuals  appear  to  become  partially  ankylosed  together. 
The  caudal  vertebrae  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 


Fia.  297.— Skeleton  and  flying  -membranes  of  the  Noctule  Bat  (Vesperugo  noctula).  x  J. 
c,  Clavicle;  h,  humerus ;  r,  radius;  u,  ulna  (rudimentary) ;  d',  pollex;  d-,  <P,  d\  &,  other 
digits  of  the  manus  supporting  torn,  the  wing-membrane ;  in,  m,  metacarpal  bones ;  ph*,  first 
phalanx  ;  phz,  second  phalanx  ;  ph$,  third  phalanx  ;  am,  antebrauhial  membrane ;  /,  femur ; 
t,  tibia ;  fb,  fibula  (rudimentary) ;  c,  calcar  supporting  im,  the  interfemoral  membrane ;  pel,  post- 
calcaneal  lobe. 

ischial  tuberosities.  The  relative  development  of  the  caudal 
vertebras  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  Pteropodidce  and  some 
Nycteridce  and  Emballonuridce.  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 
Phyllostomatidce. 

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- 
lophidce  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  Pteropodidce  and  some  of  the 
Phyllostomatidce,  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  Pteropodidce  the 
second  digit  is  provided  with  a  claw;  but  in  all  other  Bats  this  and 
the  remaining  digits  are  unarmed.  In  the  genus  Tricenops  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 
Avidely  separated ;  and  in  the  family  Rhinolopliidce  alone  do  these 
bones  form  a  symphysis.  The  ileo-pectineal  eminence  develops  a 
long  pectineal  process,  which  in  the  subfamily  Hipposiderince  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  Molossince  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  Nycteri*. 

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  Hipposiderince  and  in 
the  anomalous  genera  Thyroptera  and  Myxopoda,  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  Molossince 
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  vibrissae  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 
vibrissse  are  implanted. 

In  all  species  of  leaf-nosed  Bats,  and  especially  in  the  Rhino- 
lophidce,  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 


CHIROPTERA 


647 


the  RhinolophidcK,  the  only  family  of  insectivorous  Bats  wanting  the 

tragus,  the  auditory  bullse  reach  their  greatest  size,  and  the  highly 

sensitive  nasal  appendages  their  highest  development;  and  that  in  'the 

typical  group  of  the  Molossince  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  Ptero- 
podidce,  in  which  the  peculiar  vibratory 

.      «    .          j     -j  FIG.  298.—  Headof  JlfoZossttsj/faactntw. 

movements    first   noticed  in  Artibeus  (FromDobson,proc.zoo«.Soc.i876.)  a, 

also    be    Seen    when 


Antitragus  ;  6,  keel  of  the  ear-conch  ;  c, 


may 

the  animals  are  alarmed. 

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  Rhynchonyderis  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  oesophagus  is  narrow  in  all  species,  and  especially  so  in  the 
sanguivorous  Desmodont  Phyllostomatidce.  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  Phyllostomatiifn; 
where  the  left  or  cardiac  extremity  is  greatly  elongated,  forming  a 
long  narrow  caecum-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  Rhinopoma  microphyllum  and  Mega- 
derma  spasma  has  a  very  small  caecum  been  found. 

The  liver  is  characterised  by  the  great  size  of  the  left  lateral 
lobe,  which  occasionally  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  Ehinolophidce,  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- 
veloped vocal 
cords,  and 
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 
from  the  general 
type.  Thus 
the  pharynx  is 
long  and  very 
capacious ;  the 

Pio.  299. — Head  and  neck  of  Epotnophorns  franqueti  (adult  male,  aperture  OI    tne 

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  j        p 
where  they  communicate  with  the  pharynx ;  s,  thin  membranous  septum 

in  middle  line  between  the  anterior  pharyngeal  sacs  of  opposite  sides  ;  the  faUCCS,  and, 

s.m.,  aterno-mastoid  muscle  separating  the  anterior  from  the  posterior  opposite  to  it 
sac.  (Dobson,  Proc.  Zool.  Soc.  1881.) 

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  flying-membranes  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 
Emballonuridce,  and  especially  in  the  subfamily  Molossince  (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  vertebrae.  Its  small 
size  in  the  frugivorous  and  sangui- 
vorous species,  in  which  its  presence 
would  be  injurious  as  impeding 

their  motions  When  Searching  for  Fio.SOO.-Frontalsacaudnose-leafininale 
/.  !  ,11  T  i  -i  an(l  female  of  Hipjiosidenis  larvatus.  (Dobson, 

food  as  they  hang  suspended  by  Proc.  ZooL  5^.  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  Microchiropteru. 


Suborder  MEGACHIROPTERA. 

Frugivorous  Bats,  generally  of  large  size.  Crowns  of  molars 
smooth,  marked  with  a  longitudinal  groove  (cuspidate  in  Pternl- 
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.  Thomas1  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  PTEROPODID^E. 

Since  all  the  forms  are  included  in  this  family  its  characters 
may  be  taken  to  be  the  same  as  those  of  the  suborder. 

Subfamily  Pteropodinse. — Tongue  moderate;  molars  well  de- 
veloped. 

Epomoplwrus.2 — Dentition ;  i  -^-,  c  \-,  p  f ,  m  ^ ;  total  28  or 
26.  Tail  absent  or  very  short,  when  present  free  from  inter- 
femoral  membrane ;  second  digit  of  manus  clawed ;  premaxillae 
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  presence  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. 
2  Bennett,  Trans.  Zool.  Soc.  vol.  ii.  p.  38  (1835). 


P  TEROPODID&  65 1 

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.1 — Dentition  :  i  f ,  c  -f ,  p  f ,  m  -j ;  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 
another  in  dentition,  and  are  Fl°-  soi.-Head  of  Fox-Bat  (Ptewpus  penonatus). 

.    ,          , .   , .          .  ,      7     i  From  Gray.  Proc.  Zool.  Soc.  1866. 

mainly    distinguished    by    the 

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  Geofiroy,  Ann.  du  Museum,  vol.  xv.  p.  90  (1810). — Ex.  Brisson. 


652 


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, 


FIG.  302. — Female  and  young  of  Xantharpyia  collaris.    (From  Sclater, 
Proc.  Zool.  Soc.  1870,  p.   127.) 

suspended  by  their  hind  claws,  others  scrambling  along  among  the 
boughs,  and  taking  to  wing  when  disturbed." 

Xantharpyitt.1 — Dentition  as  in  Pteropus,  but  a  short  tail  present, 
and  the  fur  on  the  back  of  the  neck  similar  to  that  of  the  body. 

'  Gray,    List.    Spec.    Mamm.   Brit.   Mus.   pp.   37,    38    (1843):    Syn.   Cyno- 
nycteris. 


PTEROPODID^E  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.  cegyptiaca  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  :  i  -=•—,  c  -J-,  p  §,  m  f- ;  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.3 — Dentition  :  i  %,  c  \,  p  §,  m  f ;  total  24.  Premaxilla? 
well  developed  and  united  in 

front ;  facial  bones  much  ele-  ^~>3^^^x- 

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 

,    .-,  ••     '  .         , ,        FIG.  303.— Head  of  Harpyia  major.     (From  Dobson, 

nostrils,    as    shown    in    the  pr<*.  zooi.  soc.  is".) 

accompanying  woodcut. 

Cephalotes.* — Dentition :   i  \,  c  \,  p  f ,  m  f ;  total   28.     Pre- 

1  Jentink,  Notes  Leyd.  Afus.  vol.  i.  p.  117  (1879). — Amended.        '-  F.  Cuvier, 
Dents  des  Mammiferes,  p.  39  (1825).  3  Illiger,  Prodromus  Syst.  Mamm.  ct 

Avium,  p.  118  (1811).        4  Geoffrey,  Ann,  du  Mustum,  vol.  xvi.  p.  99  (1810). 


654  CHIROPTERA 


maxillse  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.1 — 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  Carponyetepiinse. — 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  papillae  near  the  tip. 

Notopteris.2 — Dentition :  i  %,  c  \,  p  f ,  m  |- ;  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,  N.  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. 

Eonyderis? — Dentition :  i  f ,  c  \,  p  f ,  m  f ;  total  34.  First  upper 
premolar  small,  with  a  single  root.  This  genus  is  likewise  repre- 
sented by  a  single  species  (E.  spelcea),  from  the  Farm  Caves,  Moul- 
mein,  Burma,  which  has  somewhat  the  appearance  of  Xantliarpyia  ; 
but  the  absence  of  a  claw  to  the  index  finger  and  the  characteristic 
tongue  and  teeth  at  once  distinguish  it. 

Carponyderis 4  and  Melonyderisf  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  Eonyderis.  Carponyderis  minima  is 

1  0.  Thomas,  Ann.  Mag.  Nat.  Hist.  ser.  6,  vol.  i.  p.  155  (1888).  2  Gray, 

Proc.  Zool.  Soc.  1859,  p.  36.  3  Dobson,  Journ.  As.  .Soc.  Bengal,  vol.  xlii. 

p.  204  (1873).  4  New  name  :  Syn.  Macroglossus,  F.  Cuvier,  Dents  des 

Mammiferes,  p.  40  (1825).  Preoccupied  by  Macroglossum,  Scopoli,  1777. 
5  Dobson,  Proc.  Zool.  Soc.  1877,  p.  119. 


MICROCHIROPTERA  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  marginatm  (compared  with  which  it  is 
proportionally  equally  destructive  to  fruit),  arid  extends  eastward 
through  the  Malay  Archipelago  as  far  as  New  Ireland,  where  it  is 
associated  with  Melonyderis  melanops,  distinguished  from  it  by  its 
larger  size  and  the  total  absence  of  the  tail. 

Nesonyderis.1 — Dentition  :  i  f ,  c  },  p  f ,  m  f ;  total  32.  Allied  to 
Melonyderis,  but  distinguished  by  the  absence  of  the  inner  pair  of 
lower  incisors,  and  of  a  claw  to  the  index  finger.  Tail  wanting. 
Kepresented  by  N.  woodfordi,  of  the  Solomon  Islands. 

Callinyderis.- — Dentition  :  i  f,  c  ^,  p  f ,  m  % ;  total  32.  Allied 
to  the  preceding,  but  with  a  short  tail ;  no  claw  to  index.  One 
species  from  Celebes. 

Trygenyderis.3 — Dentition  :  i  f ,  c  ^,  p  f ,  m  f ; '  total  34.  No 
external  tail ;  a  claw  on  index.  One  species  from  West  Africa. 

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  inf eriorly  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- 
lostomatidce) ;  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. 

Sedion  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  tuber culatus).  First  phalanx  of  the  middle  finger  extended 
(in  repose)  in  a  line  with  the  metacarpal  bone. 

1  0.  Thomas,  Ann.  Mag.  Nat.  Hist.  ser.  5,  vol.  xix.  p.  417  (1887). 
-  Jentink,  Notes  Leyd.  Mus.  vol.  xi.  p.  209  (1889). 

3  New  name  :  Syn.  Mcgaloglossus ;  Pagenstecher,  J.  B.  Mils.  Hamburg,  vol. 
ii.  p.  125  (1885).     Preoccupied  by  Mcyaglossa,  Rond.,  1865. 


656  CHIROPTERA 


•Family  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 
premaxillae  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  Bhinolophidce  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  Rhinolophinse. — 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. 

fihinolophus.1 — Dentition :   i  |,  c  \,  p  f , 
m  f ;  total  32.     Nose-leaf  (Fig.  304)  with  a 
central  process  behind  and  between  the  nasal 
orifices,  posterior  extremity  lanceolate,  anti- 
tragus   large.     Includes   more    than  twenty 
species.    E.  Indus,  in  which  the  forearm  has  a 
length  of  3  inches,  is  the  largest  species,  inhabit- 
ing elevated  hill  tracts  in  India  and  Malayana ; 
FIG.  304.— Head  of  Indian  R.    hipposidei^us   of   Europe,    extending    into 
Horse-shoe  Bat  (Rhinoiophm  gouth    England    and    Ireland,    forearm    1-5 

mitrattis).      (From     Dobson,  ,  -    ,  ,,  ,    „    . 

Monogr.  Asiat.  chiropt.)         inches,  is  one  of  the  smallest ;  and  Rferrum- 
equinum,   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 

1  Geoffrey,  Nouv.  Diet.  d'Hist.  Nat.  vol.  xix.  p.  383  (1803). 


RHINOLOPHID&  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  R.  tragatus,  distinguished  by  having  three 
vertical  grooves  on  the  lower  lip,  in  place  of  the  single  groove  found 
in  R.  ferrum-equinum.  Rhinolophus  is  represented  in  the  Upper 
Eocene  Phosphorites  of  Central  France  by  R.  antiquus  and  R. 
dubius ;  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.  Alastor,  from  the  same  deposits,  differs  from  Rhino- 
lophus by  the  extreme  shortness  of  the  nasal  region.  Palceonycteris, 
from  the  Lower  Miocene  of  France,  is  said  to  be  allied  to  Rhino- 
lophus, but  the  premolars  are  f ,  and  the  limb  bones  are  stated  to 
resemble  those  of  Molossus. 

Subfamily  Hipposiderinse. — Toes  e'qual,  of  two  phalanges  each  ; 
ilio-pectineal  spine  united  by  a  bony  isthmus  with  a  process  derived 
from  the  antero-inferior  surface  of  the  ilium. 

Hipposidei'us.1 — Dentition  :  i  £,  c  \,  p  -^y-,  m  f ;  total  30  or  28. 
Tail  well  developed.  This  genus,  of  which  more  than  twenty 
species  have  been  described,  differs 
from  Rhinolophus  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,  H.  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 

r  ,  .     ,        ,  i      /.  T  caratus.    (From  Dobson,  Proc.  ZooL  Soc. 

behind    the    nose -leaf,    rudimentary  1377.) 

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. 

Anthops.- — Like  Hipposidems,  but  with  the  tail  rudimentary, 
consisting  merely  of  three  or  four  vertebrae  hidden  in  the  base  of 
the  interfemoral  membrane.  Nose-leaf  very  complicated,  its  upright 
transverse  portion  emarginate  above,  and  the  projections  rounded 

1  Gray,  Proc.  Zool.  Soc.  1834,  p.  53.  The  Bats  of  this  genus  are  usually 
described  as  Phyllorhina,  but  this  use  has  been  shown  to  be  incorrect ;  see  Blan- 
ford,  Proc.  Zool.  Soc.  1887,  \\  637. 

-  0.  Thomas,  Ann.  Mag.  Nat.  Hist.  ser.  6,  vol.  i.  p.  156  (1888). 

42 


658  CHIROPTERA 


and  hollowed  behind,  and  their  substance  quite  thin.     Premolars  H. 

Represented  by  A.  ornatu*  of  the  Solomon  Islands. 

Mr.  0.  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  Codops  in  the  rudimentary  tail. 

Rhiiwnyderis l  and  Tricenops.2 — These  are  two  allied  genera  with 

well -developed  tails;  the  former 
being  represented  by  R.  aurantia 
from  Australia,  and  the  latter  by 
T.  persicus  from  Persia  and  Eastern 
Africa.  Tricenops  (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. 

Ccelops.3 — This  genus  is  known 
only  by  a  single  species,  C.  frithi, 
from  the  Bengal  Sunderbans, 

FIG.  306. — Head  of  Tricenops  persicus.     x  2.       _  j     cr  /•         r_  £      £ 

(Frorn-Do}iscm,Monogr.Asiat.chiropt.)       Java,  and    biam  (in  the  root  ot 

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  NYCTERID^E. 

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  Rhinolophidcz  by  the  presence 
of  a  distinct  tragus  to  the  ear,  and  by  the  premaxillae  being  cartila- 
ginous or  small  and  separated  from  one  another  in  front  by  a  dis- 
tinct space. 

Megaderma.* — Dentition  :  i  f ,  c  ^,  p  ^^,  m  f ;  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.  2  Dobson,  Journ.  As.  Soc.  Bengal, 

vol.  xl.  p.  455  (1871).  3  Blyth,  Journ.  As.  Soc.  Bengal,  vol.  xvii.  p.  251  (1848). 
4  Geoffrey,  Ann.  du  Museum,  vol.  xv.  p.  197  (1810). 


NYCTERID.-E 


659 


inches),  is  not  only  the  largest  species  of  the  genus  but  also  of  the 
suborder.  M.  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 


FIG.  307.— Megaderma  glgas.     x  J.    (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  M.  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  :  i  •§,  c  ^,p  J,  mf ;  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  Geoffrey,  N&uv.  Diet.  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  N.  jttvanica,  they  are  all  limited  to  the  Ethiopian  region. 


Family  VESPERTILIONID^E. 

Nostrils  opening  by  simple  crescentic  or  circular  apertures  at 
the  extremity  of  the  muzzle,  not  surrounded  by  distinct  foliaceous 
cutaneous  appendages ;  premaxillse  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  §  to  |,  rarely  (in  Antrozmis  only)  \,  premolars  f,  or  f , 
or  |,  rarely  (in  Vesperugo  noctivagans  of  North  America)  f ;  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, 
F'espertilionine,  Miniopterine,  and  Thyropterine  divisions. 

In  the  Plecotine  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-leaves ;  the 
ears  also  are  generally  very  large  and  united. 

Plecotus.1 — Dentition :  i  % ,  c  ^,  p  §,  m  f ;  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 

1  Geoffroy,  Descript.  de  VEgypte,  vol.  ii.  p.  112  (1812). 


VESPERTILIONID^E  66 1 

the  glandular  prominences  of  the  sides  of  the  muzzle,  which  meet 
in  the  centre  above  and  behind  the  nostrils.  P.  auritus  extends 
over  the  greater  part  of  the  Palsearctic  region,  occurring  in  Ireland 
in  the  west  and  the  Himalaya  in  the  east. 

Synotus.1 — Dentition  :  i  § ,  c  ^,  p  f ,  m  f ;  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 
Barbastelle  Bat  (S.  barbastellus)  and  S.  darjilingensis  from  the  Hima- 
laya. 

Otonycteris.2 — Dentition:  i  |,  c  \,  p  J,  m  -f ;  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. 

Nydophilus.3 — Dentition  :  i  %,  c  -f,  p  £,  m  f ;  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  N.  timoriensis  of  the  Australian  region,  and  N.  microtis  of 
New  Guinea. 

Antrozous.* — Dentition  :  i  ^,  c  ^,  p  J,  m  -f ;  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,  A.  pallidus,  in- 
habits California. 

The  Vespertilionine  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 
small  size  of  the  ears,  and  the  absence  of  grooves  on  the  forehead. 

Fesperugo.5 — Dentition:  i  *~~^ ,  c  \,  p  3-3'  mfj  tota^  ^4,  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  $  in  the 
subgenera  Scotozous  and  Rhogeessa,  and  the  premolars  from  f-  to  f  (in 
the  subgenus  Lasionyderis  f ).  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,  Wirbelthiere  Europ.  p.  55  (1840).  -  Peters, 

Monatsber.  Ak.  Berlin,  1859,  p.  222.  3  Leach,  Trans.  Linn.  Soc.  vol.  xiii. 

p.  78  (1822).  4  Allen,  Proc.  Ac.  Nat.  Sci.  Philad.  1862,  p.  247. 

5  Keyserling  and  Blasius,  Wicgmann'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.  Vesperugo  probably  contains  the  greatest  number 
of  individuals  among  the  genera  of  Chiroptera,  and,  with  the  excep- 
tion of  Vespertilio,  its  species  have  also  the  widest  geographical 
range,  being  almost  cosmopolitan;  and  one  of  the  species,  the  well- 
known  Serotine  (V.  [F'esperus]  serotinus)  is  remarkable  as  the  only 
species  of  Bat  known  to  inhabit  both  the  Old  and  the  New  World ; 
one  (V.  borealis)  has  been  found  close  to  the  limits  of  the  Arctic 
circle,  and  another  (V.  magellanicus)  inhabits  the  cold  and  desolate 
shores  of  the  Straits  of  Magellan,  being  doubtless  the  Bat  referred 
to  by  Mr.  Darwin  in  the  Naturalist's  Voyage.  The  Common  Pipis- 
trelle  (V.pipistrellus),  ranging  over  the  greater  part  of  the  Palsearctic 
region,  is  the  best  known  species. 

Chalinolobus.1 — This  genus  agrees  with  Vesperugo  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  -f,  represented  by 
C.  morio  from  New  Zealand,  Tasmania,  and  Australia,  and  three 
other  species  from  Australia ;  and  Glaiiconycteris,  with  p  J,  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  ^,  c  ^,  p  £,  m  § ;  total  30.  This 
genus  comprises  a  comparatively  small  number  of  species  nearly 
allied  to  Vesperugo,  and  some*  of  which 
approach  so  closely  to  the  aberrant  types  of 
the  latter  ranged  under  the  subgenus  Scotozous, 
as  to  render  the  definition  of  the  present  genus 
almost  impossible.3  The  species  are  restricted 
to  the  tropical  and  subtropical  regions  of  the 
eastern  hemisphere,  though  widely  distributed 
within  these  limits.  The  more  typical  species 
FIG.  3os.— Head  of  Scoto-  are  distinguished  especially  by  the  single  pair 

philus  emarg\natus.  (Dobson,  .  .        J  11 

Monogr.  Asiat.  curopt.)  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, 

1  Peters,  Monatsber.  Ak.  Berlin,  1866,  p.  672. 

2  Leach,  Trans.  Linn.  Soc.  vol.  xiii.  p.  71  (1822). 

3  See  0.  Thomas,  Ann.  Mus.  Genova  (2),  vol.  ix.  pp.  84-88  (1890). 


VESPERTILIONID&  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.  S.  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  Fesperugo  (Scotozous) 
dormeri  as  to  render  it  almost  impossible  to  distinguish  Scotophilus 
and  Fesperugo.  The  figured  species  is  from  India. 

Nycticejus.1 — 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.  0.  Thomas,  that  the  discovery  of  Scotophilus  albofuscus 
renders  the  generic  distinctness  of  Nycticejus  no  longer  tenable,  and 
that  the  species  should  be  known  as  Scotophilus  humeralis. 

Atalapha* — Dentition:  i  J,  c  $,p^,m%;  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.3 — Dentition  :  i  §,  c,  T,jpf,  m  f ;  total  34.  This 
genus  includes  some  eight  species  of  small  Bats  distinguished  by 
their  prominent  tube-like  nostrils  and  hairy  interfemoral  membrane. 
H.  suillus,  from  Java  and  neighbouring  islands,  is  the  best -known 
species,  and  another  closely  allied  (H.  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  countries  in  which  they  are  found. 

Fespertilio.* — Dentition :  i  •§,  c  \,  p  f ,  ra  f ;  total  38.  Next 
to  Fesperugo,  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.  Ixxxviii.  p.  417(1819).  2  Rafinesque, 
Precis  des  Decouvtrtes  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 


PIG.  309. —Side  and 
front  views  of  the  head 
of  Cerivoula  hardwickei. 


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  Eeddish-Gray  Bat  ( V.  nattereri), 

of  very  local  occurrence ;  Daubenton's  Bat  ( V,  daubentoni) ;  and  the 

Whiskered  Bat  (V.  mystadnus). 

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.  pida,  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 
stand  as  regards  the  species  of  Vespertilio  in  a 
position  similar  to  that  occupied  by  Chalinolobus 
with  respect  to  Vesperugo. 

The  Miniopterine  division  includes  only  two 

(Dobson,  Monogr.  ASM.   crenera  an(j  js  characterised  by  the  great  eleva- 

Chiropt.)  3  ,'  ,      ,        ,        , *    ,          >  -      .   , 

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.2 — 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. 
micropus  being  shown  in  Fig.  310. 

Miniopterus.3 — Dentition  :  i  •§, 
c  T>  P  §>  *»  f ;  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),  Kcrivoula. 

2  Gray,  Mag.  Zool.  Bat.  vol.  ii.  p.  496  (1838). 

3  Bonaparte,  Fauna  Italica,  fasc.  xxi.  (1837). 


Fie.  310. — Head  of  Katahis  microput. 
(Dobson,  Proc.  Zool.  Hoc.  1880.) 


X3. 


VESPERTILIONID&  66$ 

these  the  best-known,  M.  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. 

Thyroptera.1 — Dentition  :  i  § ,  c  ^,  p  f ,  m  % ;  total  38.  In  the 
single  species  T.  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 


Fio.  311. — Suctorial  discs  in  Thyroptera  tricolor,  a,  Side  and  fc,  concave  surface,  of  thumb- 
disc  ;  c,  foot  with  disc,  and  calcar  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. 

Myxopoda.2 — The  second  genus  is  likewise  represented  only  by 
a  single  species — M.  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  Vespertilionidce. — It  is  not  improbable  that  Vesperwgo  is 
represented  in  the  Upper  Eocene  of  the  Paris  basin  by  V.  pari- 
siensis,  which  appears  to  be  allied  to  V.  serotina,  although  it  has 
been  regarded  by  some  writers  as  genetically  distinct,  under  the 
name  of  Nyctitherium.  Vesperugo  (Nyctitherium)  also  occurs  in  the 
Bridger  Eocene  of  the  United  States;  Nydilestes  from  the  same 

1  Spix,  Sim.  and  Vesp.  Bresil,  p.  61  (1823). 

2  A.  Milne-Edwards,  Bull.  Soc.  Philom.  ser.  7,  vol.  ii.  p.  1  (1878). 


666  CHIROPTERA 

deposits  being  an  allied  extinct  genus.  A  number  of  European 
Miocene  species  have  been  referred  to  F'espertilio,  but  the  term  in 
these  cases  must  be  used  in  a  somewhat  wide  sense.  Vespertiliavus, 
of  the  Phosphorites  of  Central  France,  differs  from  FespertUio  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^E. 

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  Emballonmidce  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  Vespertilionidce,  but  rarely 
extends  north  or  south  of  the  thirtieth  parallel  of  latitude. 

Subfamily  Emballonurinae. — 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  i  f ,  c  $,  p  f ,  m  f ;  total  38. 

Represented  by  two  genera,  Furipterus l  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,  Faun.  Ital.  vol.  i.  (1832-41)  :  Syn.  Furia,  F.  Cuvier,  Mim.  du 
Mustum,  vol.  xvi.  p.  150  (1828).     Preoccupied  by  Linn.  1766. 

2  Peters,  Monatsber.  Ak.  Berlin,  1877,  p.  185. 


EMBALLONURID&  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  f-. 

Emballonura.1 — Incisors  §.  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.  Fl°-  312-~ Earof£m6aZio- 

j-v  j   ..       a      T  -i         -ni    ,  •,  p    ,i        nuraraffrayana.    x2.    (Dob- 

Coleura.2— Incisors  £.     Extremity  of  the  SODj  ££  yZooL  Soc  187J , 
muzzle  broad,  forehead  concave.      Has  two 
species  from  East  Africa  and  the  Seychelles  Islands. 

Rhynchonycteris? — This  genus  is  distinguished  from  Coleura  by 
the  much-produced  extremity  of  the  muzzle.  The  single  species,  II. 
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  ^.  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.5 — Incisors  \ ;  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  Hceven),  Tijdsch.  Nat.  Ges.  1839,  p.  22. 

2  Peters,  Monatsber.  Ak.  Berlin,  1867,  p.  479.          3  Peters,  loc.  cit.  p.  477. 
4  Illiger,  Prodromus  Syst.  Mamm.  et  Avium,  p.  121  (1811). 

3  Geoffroy,  Descript.  de  VEgypte,  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  T.  melanopogon,  but  about  its  usual  position  the 
openings  of  small  pores  may  be  seen,  the  secretion  exuding  from 


FIG.  313. — Heads  of  Taphozovs  longlmanus,  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  Diclidurine  division  there  is  but  a  single  genus,  repre- 
sented by  two  species. 

Dididurus.1 — Dentition  :  i  -£,  c  ^,  p  -f,  m  f ;  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  Taphozous  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  Nodilionine  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  f,  c  ^,  p  A,  m  % ;  total  28.  The  two 
species  N.  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  to  a  Rodent  actually  led  Linnaeus  to  remove  this  species 
from  the  Bats  and  place  it  in  the  Rodents.  This  Bat  is  remark- 

1  Wied,  Isis,  1819,  p.  1629.       2  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  88  (1766). 


EMBALLONURID&  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  Rhinopoma- 
tine  division. 

Rhinopoma.1 — This  genus,   represented  by  the  single    species 
R.  microphyllum,  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  Nycteridce  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       FIG.  314.— skull  of  Rhinopoma  micro- 
the  index  finger  as  by  its  remarkably  !*»«««.    xa.  (Dobson,  Monogr.  Asiat. 

j      i       j         t.    -i  •      i-          £        Chiropt.) 

long  and  slender  tail   projecting  tar 

beyond  the  narrow  interfemoral  membrane,  inhabits  the  subterranean 
tombs  in  Egypt  and  deserted  buildings  generally  from  North-East 
Africa  to  Burma. 

Subfamily  Molossinse. — Tail  thick,  produced  far  beyond  the 
posterior  margin  of  the  interfemoral  membrane  (except  in  Mysta- 
cops) ;  legs  short  and  strong,  with  well-developed  fibula ;  upper 
incisors  strong.  This  subfamily  includes  all  the  species  of  Embal- 
lonuridce  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  varying  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,  Avhich  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  Molossine  division  is  characterised  by  the  production  of  the 
tail  beyond  the  posterior  margin  of  the  interfemoral  membrane ;  it 
includes  three  genera. 

Chiromeles.2 — Dentition  :  i  i,  c  ^,  p  4-,  ra  f ;  total  26.  Hallux 
much  larger  than  the  other  toes  and  separable  from  them,  ears 

1  Geoffrey,  Descript.  de  I'Egypte,  vol.  ii.  p.  123  (1812). 
-  Horsfield,  Zool.  Research  Java  (1824). 


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  mammae  are  placed. 

Molossus.1 — Dentition:  i  -—^^  c  ^,  p  ^-,  m  f ;  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.  M.  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, 
if  not  in  all,  species  of  Bats).  In  the 

FIG.  315. — Head  of  Molossus  glaucinus.  '       j  . 

(Dobson,  Proc.  Zooi.  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* — Dentition  :  i  -^,  c  ^,  p  ^-,  m  f ;  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  Molossus,  in  many  of  the 
species  (as  in  the  woodcut  of  the  head  of  N,  macrotis,  Fig. 
316)  showing  vertical  wrinkles.  N.  tceniotis,  one  of  the  largest 
species,  alone  extends  into  Europe,  and  has  been  taken  as 
far  north  as  Switzerland.  N.  johorensis,  from  the  Malay  Penin- 
sula, is  remarkable  from  the  extraordinary  form  of  its  ears. 
N.  brasiliensis  is  nearly  as  common  as  Molossus  obscurus  in  tropical 
America,  and  extends  farther  north  (California)  and  south  than 
that  species. 

1  Geoffroy,  Ann.  du  Mus&um,  vol.  vi.  p.  154  (1805). 

2  Geoffroy,  Descript.  de  VEgyptc,  vol.  ii.  p.  114  (1812). 


EMBALLONURID^E 


671 


In  the  Mystacopine  division  the  tail  perforates  the  interfemoral 
membrane  and  appears  upon  the  upper  surface. 

Mystacops.1 — This  genus  includes  only  M.  tuberculatus  of  New 
Zealand,  where,  together  with  Chalinolobus  tumwio,  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 
quadrupedal  of  Bats.  Other  peculiarities  FlG-  316;-^ead 

»,  ri-i  111       macrotis.    (Dobson,  Proc.  Zool.  Soc. 

of  structure  are  found  in  the  remarkable  187«5.) 

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, 


FIG.  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  Emballonuridce. — In  the  cavern-deposits  of  Madras  remains 
of  the  existing  Taphozous  saccolcemus  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.  Mystatina ;  Gray,  Voyage  of  the  "Sulphur"  "Mamm." 
p.  23  (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  PHYLLOSTOMATID.E. 

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 ;  premaxillae  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  Ehinolophidce,  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  Pteropodidce,  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  Vampirine  and  Glossopliagine  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  Chilonyeteriinse. — 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 : 
*  f»  c  \'P  t>  w  t ;  total  34. 
The  crown  of  the  head  is 
moderately  elevated  above 

FIG.  318.— Head  of  Mormops  blainvillei.    (Dobson,          ,       ,      .   ,  ,.  ,   ,,       i       - 

Cat.  Chiropt.  Brit.  Mvs.)  the  *aclal  .lln.e>  and  the.  baS1' 

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,  Trans.  Linn.Soc.  vol.  xiii.  p.  76  (1820-22).— Amended. 


PHYLLOSTOMATID^E  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  Phyllostomatinse. — 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  Fampirine  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 ;  interf emoral  membrane  well  developed ;  tail  generally 
distinct,  rarely  absent ;  inner  margin  of  the  lips  not  fringed.  The 
dentition  is  :  i  -~^-,  c  ^,  p  -j^j-j  vn  f ;  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,2  and  Dolichophyllum.3 — 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.  aurita,  in  which  the  nose-leaf  is  much  elongated,  and  the 
ear-conch  and  tragus  are  unusually  large. 

Vampyrusf  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  Fampyrus,Lophostoma,  Micronycteris? 

1  Tomes,  Proc.  Zool.  Soc.  1863,  p.  81.  2  New  name  :  Syn.  Macrotus  ; 

Gray,  Proc.  Zool.  Soc.  1843,  p.  21.  Preoccupied  by  Macrotis,  Dej.  1833. 
3  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.  5  Gray,  Proc.  Zool.  Soc.  1866,  p.  113.  Syn.  Schizostoma  ; 
Gervais,  1855.  Preoccupied  by  Broun,  1835. 

43 


674  CHIROPTERA 

Trachyops,  Phylloderma,  Phyllostoma,  Anthorhina,1  Mimon,  Hemiderma  2 
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  Vampyrus  spectrum,  is  a  well- 
known  species  in  South  America ;  P. 
elongatum  (Fig.  319)  differs  in  its  smaller 
size  and  much  larger  nose-leaf.  Hemi- 
derma brevicauda  is  a  small  species, 
FiG.w.-He*d  of  Phyllostoma  don-  which  forms  a  connecting  link  between 

gatum.    (From  Dobson,  Proc.  Zool.  Soc.      .  .  .     ,  . .    .   .    c 

1866.)  this  and  the  next  division.  Ehinophijlla 

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  papillae ;  lower  lip  with  a  wide  groove  above,  and 
in  front  margined  by  small  warts ;  nose-leaf  small ;  tail  short  or 
absent.  Dentition :  i  i,  c  f ,  p  -|  ^-,  m  -~  ;  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,3  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  Glossopliaga  and  Phyllonycteris  the  upper 
incisors  form,  as  in  the  Fampyrine  division,  a  continuous  row  between 
the  canines ;  in  Monophylla  and  Leptonycteris 4  they  are  separated 
into  pairs  by  a  narrow  interval  in  front;  while  in  Lonchoglossa, 
Glossonycteris,  and  Chceronyderis  they  are  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 

1  New  name:  Syn.  Tylostoma;  .Gervais,  1855.  Preoccupied  by  Sharpe,  1849. 
2  Gervais,  Castlenau's  Exped.-Zool.  p.  43  (1855) :  Syn.  Carollia,  Gray,  1838. 
Preoccupied  by  Carolia,  Cantraine,  1837.  3  The  references  to  the  genera  of 
this  and  the  following  division  will  be  found  in  Dobson's  Catalogue.  *  New 

name :  Syn.  Ischnoglossa,  Saussure,  1860.     Preoccupied  by  Kraatz,  1856. 


PHYLLOSTOMATJD^E  675 

they  are  deciduous  even  in  youth.  The  zygomatic  arch  is  wanting 
in  Phyllonyderis,  Glossonycteris,  and  Chceronyderis. 

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          ^I0'  ^~®' — Head  °f  Chteronycteris  mexicana,  shoving 
,  j         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  corollas  of  certain  flowers.  This 
type  of  tongue  is  shown  in  the  woodcut  of  the  head  of  Chceronyderis 
(Fig.  320);  and  it  is  paralleled  among  the  Megachiroptera  by  the 
Carponycteriine  Pteropodidce. 

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 
margin  of  the  lips  fringed  with  conical  papillae.  Dentition : 
*  ~2^T>  P  f »  m  !=I  >  the  number  of  the  molars  being  either  f ,  f , 
or  f  in  different  species ;  premolars  and  molars  very  broad  (except 
in  Sturnira),  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,  A.  perspicillatus.  Waterton  believed  that 
A.  planirostris,  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  Pteropodida  more  closely  than 
those  of  any  other  Bats ;  and  the  presence 
of  tufts  of  long  differently  coloured  hairs 
over  glands  in  the  sides  of  the  neck  shows 

FIG.  321.— Head  of  Centuno  senex.  ,° 

(Dobson,  cat.  chiwpt.  Srit.  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 :  i  ^,  c  ^,  p  -f, 
m  J  ^  ;  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,  in testiniform.  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 


PHYLLOSTOMATID^E  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.  rufiis)  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  2^,  with 
a  remarkably  long  and  strong  thumb ; 
it   is    destitute    of    a   tail,   and    has    a 
peculiar  physiognomy,  well  represented      FK».  322.— Head  of  vampire  sat- 
in Fig.  322.     The  body  is  covered  with 

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  AViecl,  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- 

Fio.  323.— Dentition  of  Desmodiis  rufus.  a,  Front  tremities  of  the  large  Upper 
view  of  upper  teeth ;  b,  left  lateral  view  of  upper  and  incisors.  The  exceedingly 
lower  teeth.  &  ' 

narrow  oesophagus  opens  at 

right  angles  into  the  slender,  intestine-like  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  oesophagus,  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. 

Diphylla.1 — A  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  Phyllostomatidce. — Remains  of  Vampyrus  spectrum,  as  Avell 
as  of  several  species  of  Phyllostoma  or  closely  allied  types,  are  found 
1  Spix,  Sim.  et  Vesp.  Brazil,  p.  68  (1823). 


PHYLLOSTOMA  TID^E  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  0.  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,  Eecherches  Anatomiques  sur  les  Mam- 
miferes  de  VOrdre  des  Chiropteres,  Paris,  1881 ;  W.  T.  Blanford,  "Notes  on  Indian 
Chiroptera,"  Journ.  As.  Soc.  Bengal,  vol.  Iviii.  (1888).  See  also  papers  by  Jentink, 
Bocage,  and  others. 


CHAPTER   XIV 

THE    OKDER    PEIMATES 

THIS  order  in  the  system  of  Linnaeus  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.1 

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 
f ,  and  that  of  the  molars,  with  the  exception  of  the  Hapalidce, 
being  -f .  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  1'embryologie  des 
Lemuriens  et  sur  les  affinites  zoologiques  de  ces  animaux,"  in  the  Ann.  des 
Sciences  Nat.  October  1871;  and  P.  Gervais,  "Encephale  des  Lemures,"  in 
Journ.  de  Zoologie,  torn.  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.  1873,  p.  484. 


PRIM  A  TES 


68 1 


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  Semn&pithecince  of  the 
Cercopithecidce ;  and  there  is  always  a 
caecum,  which  is  generally  of  large  size. 
The  placenta  may  be  either  non-deciduous, 
or  discoidal  and  deciduous.  There  are 
always  two  mammae  in  the  pectoral 
region,  except  in  Chiromys;  and  the  Flo.  324._Aiimentary  canal  of 

testCS  descend  into  a  Scrotum.  GcOago,  the  greater  part  of  the  small 

The  Lemuroidea  are  decidedly  low  in  int«8tine  being  omitted,  d,  duo- 

. ,  i         .  .,-          ,1-1  denum ;  i.  ileum ;  cm,  caecum ;  r, 

the  scale  of  organisation,  their  placenta-  rectum. 
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  Anaptonwphus 
they  are  trituberculate. 


682  PRIM  A  TES 


Suborder  LEMUROIDEA. 

The  Latin  term  Lemur  was  applied  by  Linnaeus  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  Prosimice  by 
some  systematic  writers.  When  the  name  was  bestowed  by 
Linnaeus  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 


LEMURIDSE  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  mammae.  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  Kodents,  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  LEMURID^E. 

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  PRIM  A  TES 

second  of  the  hind  foot)  with  flat  nails,  and  corresponding  form  of 
ungual  phalanges. 

Subfamily  Indrisinae. — The  dentition  of  the  adult  consists  of 
thirty  teeth,  usually  expressed  by  the  formula  i  f ,  c  $,  p  f ,  m  f ; 
but,  as  indicated  above,  they  may  be  i  f ,  c  ^,  p  f-,  m  f .  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.  Mammae  two,  pectoral.  Caecum  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. 

Indiis.1 — Upper  incisors  subequal  in  size.  Upper  canine  larger 
than  the  first  premolar.  Muzzle  moderately  long.  Ears  exserted. 
Carpus  without  an  os  centrale.  Tail  rudimentary.  Vertebrae : 
C  7,  D  12,  L  9,  S  4,  C  9. 

The  only  well-established  species  is  the  Indris  (7.  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.2 — 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.  Vertebrae :  C  7,  D  12,  L  8,  S  3,  C  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.  verreauxii,  and  P. 
coronatus  (Fig.  326).  Some  of  these  are  to  be  found  in  almost 

1  Geoffrey,  Mag.  Encyclop.  2d  ann.  vol.  i.  p.  46  (1796),  "Indri." 
2  Bennett,  Proc.  Zool.  Soc.  1832,  p.  20. 


LEMURIDsE 


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 


FIG.  825.— Indris  (/jtdris  brevicaudatd).    From  Milne-Edwards  and  Grandidier, 
Mammiftres  de  Madagascar,  pi.  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 


FIG.  326. — Propithecus  coronatm.    (From  Milne-Edwards  and  Grandidier,  Mammifkres  de 
Madagascar,  pi.  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.  Tail  long.  Vertebrae:  C  7, 
D  11,  L  9,  S  3,  C  23. 

One  species,  A.  laniger,  the  Woolly  Lemur,  or  Avahis,  consider- 
ably smaller  than  any  of  the  last  genus.     It  differs  from  them  in 

1  Jourdan,  Mem.  de  I'lnstitut,  vol.  ii.  p.  231  (1834). 


LEMURID& 


68? 


its  habits,  being  quite  nocturnal,  and  not  associating  in  small  troops, 
but  being  ahvays  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  Indrisince.  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  f ,  c  ^,  p  f ,  m  f . 
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  caecum  much  less  devel- 
oped. Tarsus  but  slightly  elongated,  the  calcaneum  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.1 — 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.  Mammae  two,  pectoral.  Vertebrae :  C  7,  D  12,  L  7  (or  D 
13,  L  6),  S  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 
trees  on  all  four 
feet,  but  also 
jumping  with 

marvellous  agility.     They  are  gregarious,  living  in   small  troops, 

are  diurnal  in  their  habits,  but  most  active  towards  evening,  when 

1  Linn.  Syst.  Nat.  12th  ed.  vol.  i.  p.  44  (1766). 


Fio.  327. — Skull  of  Ring-tailed  Lemur  (Lemur  catta).     x  J.    we, 
Upper  canine  ;  Ic,  lower  canine  ;  pm,  premolars ;  TO,  molars. 


688 


PRIM  A  TES 


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 


FIG.  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  Indrisince  has  as  yet  lived  long  enough  in  captivity  to  be 
brought  alive  to  Europe,  various  species  of  Lemurince  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, 


LEMURIDA:  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  l  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  L.  macaco  the  male  is  black 
and  the  female  red.  L.  mongoz,  L.  collaris,  and  L.  albifrons  are 
other  well-known  species. 

Hapalemur? — 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.  Mammae  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.  simus,  but  it  is  doubtful  if  it  is 
more  than  a  variety. 

Lepidokmur.3 — Upper  incisors  absent  or  rudimentary.  Muzzle 
more  elongated  than  in  the  last.  No  distinct  os  centrale  in  the 
carpus.  L.  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  premaxillse,  containing  a  pair  of  small 
styliform  incisors,  has  been  described  by  Peters  *  under  the  name 
of  Myxocebiis  caniceps. 

Subfamily  Galaginse. — Dentition  as  in  Lemurina\  from  which 
the  members  of  this  subfamily  are  distinguished  by  the  elongation 
of  the  tarsus,  caused  by  a  peculiar  modification  of  the  calcaneum 
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? — Last  upper  premolar  very  much  smaller  than  the 
first  molar,  with  only  one  external  cusp.  The  animals  included 

1  Proc.  Zool.  Soc.  1879,  p.  132.  2  Gray,  Proc.  Zool.  Soc.  1870,  p.  829. 

3  I.  Geoffrey,  Cat.  Mus.  Hist.  Nat.  Paris,  p.   75  (1851).      Amended  from 
L'p'dcmur.  4  Monatsb.  Ak.  Berlin,  1874,  p.  690. 

5  Geoffrey,  Ann.  duMusium,  vol.  xix.  p.  171  (1812). 

44 


690  PRIM  A  TES 


under  this  name  appear  to  form  a  transition  between  the  true 
Lemurs  and  the  Galagos.  The  genus  was  originally  established  by 
Geoffrey  St.  Hilaire  in  1812  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  Microcebus.  They  are  all  small,  some  being  less  than 
a  rat  in  size,  long-tailed,  and  nocturnal  in  their  habits.  One  of  the 
largest,  C.  furcifer,  is  of  a  reddish-gray  colour,  and  distinguished 
by  a  dark  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.  gliroides,  C.  milii,  etc.)  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  fat-tailed  Sheep  of  the  Cape,  but  which  by  the  time 
they  emerge  from  their  torpor  has  acquired  its  normal  proportions. 
The  smallest  species,  to  which  many  names  have  been  given 
(C.  pusillus,  rufus,  smithi,  etc.),  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.1 — Last  upper  premolar  with  two  large  external  cusps, 
and  nearly  equalling  the  first  molar  in  size.  Calcaneum  about  one- 
third  the  length  of  the  tibia,  and  the  navicular  much  longer  than 
the  cuboid.  Vertebrae :  C  7,  D  13,  L  6,  S  3,  C  22-26.  Tail  long, 
and  generally  bushy.  Ears  large,  rounded,  naked,  and  capable  of 
being  folded  at  the  will  of  the  animal.  Mammae  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  eyes  and  ears,  soft  woolly  fur,  and  long  tails. 
There  are  several  species,  of  which  G.  crassicaudatus,  from  Mozam- 
bique, is  the  largest.  A  similar  species,  or  perhaps  variety,  from 
1  Geoffroy,  Mag.  Encydop.  2d  ami.  vol.  i.  p.  49  (1796). 


LEMURID&  69 1 

Angola  is  G.  montieri.  G.  garnetti,  alleni,  maholi,  demidoffi,  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  Lorisinoe. — Dental  formula  as  in  Lemurinee.  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.  Mammse  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  Lemuridce,  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), 
Stenopsoi.  Illiger  (1811),  but  they  were  in  1812  divided  by  Geoffroy 
into  two  genera,  Nycticebus  and  Loris,  a  division  which  has  been 
accepted  by  most  modern  zoologists. 

Nycticebus.1 — 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  maxillae.  Body 
and  limbs  stout.  No  external  tail.  Vertebras  :  C  7,  D  17,  L  6,  S  3, 
C  12.  The  species  are  N.  tardigradus,  the  common  Slow  Lemur  or 
Loris,  of  the  Malay  Countries,  Sumatra,  and  Borneo ;  N.  javanicus, 
of  Java ;  and  N.  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  Musium,  vol.  xix.  pp.  162,  163  (1812). 


692 


PRIM  A  TES 


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  Avith  one  of  their  hands.  The 
female  produces  but  one  young  one  at  a  time.  L.  tardigradus  was 
placed  by  Linnaeus  at  the  head  of  the  list  of  species  of  his  genus 
Lemur,  and  its  habits  doubtless  suggested  the  generic  name  which 


FIG.  329.— The  Gray  Loris  (Nycticebus  cinereus).    From  A.  Milne-Edwards,  N.  Archives 
du  Museum,  vol.  iii.  pi.  3. 

was  transferred  by  Geoffroy  to  the  less  nocturnal  and  spectre-like 
Madagascar  members  of  the  group.1 

.Z/om.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  premaxillae  produced  forwards  considerably 
beyond  the  anterior  limits  of  the  maxillae,  and  supporting  a  pointed 
nose.  Body  and  limbs  slender.  No  external  tail.  Vertebrae :  C  7, 
D  14,  L  9,  S  3,  C  6.  This  genus  is  represented  only  by  the  Slender 
Loris  (L.  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.  Sliroeder  van  der  Kolk  and 
W.  Vrolik,   "  Recherches  d'Anatomie  comparee  sur  le  genre  Stenops  d'llliger,"  in 
Bijdragen  tot  de  Dierkunde,  Part  I,  Amsterdam,  1848-54. 

2  Geoffroy,  Mag.  Encydop.  2d  aim.  vol.  i.  p.  48  (1796). 


LEMURID.  /. 


693 


FIG.  330.—  The  Slender  Lori*  (Loris  gracilis).     From  Blanford, 
Mammalia  of  British  India,  p.  4". 


long  thin  body,  long,  angularly  bent,  slender  limbs,  and  no  tail. 

Its  habits,  according  to  Mr.  W.  T.  Blanford,1  are  "very  similar 

to    those    of 

Nycticebus   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. 

Perodidicus.2 — A  short  tail,  about  a  third  of  the  length  of  the 
trunk.  Two  or  three  of  the  anterior  dorsal  vertebrae  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  Bosnian, 
who  met  with  it  in  his  voyage  to  Guinea.  It  was,  however,  lost 
sight  of  until  1825,  Avhen  it  was  re-discovered  in  Sierra  Leone,  and 
fully  described  by  Bennett  in  1830  under  the  name  of  Perodicticus 
geoffroyi.  Bennett's  generic  name  has  been  retained,  but  the  specific 
name  bestowed  by  Gmelin,  adopted  from  Bosnian,  has  been  restored. 
It  is  also  found  in  the  Gaboon.  It  is  strictly  nocturnal,  and  slower 
in  its  movements  even  than  Nycticebus  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  Ardocelnis  of  Gray.  It  is  found 

1  Mammalia  of  British  India,  p.  48  (1888). 
2  Bennett,  Proc.  Zool.  Soc.  1839,  p.  109. 


694  PRIM  A  TES 


at  Old  Calabar,  and  is  very  rare,  only  a  few  individuals  having  as 
yet  been  met  with.     Vertebrae :  C  7,  D  15,  L  7,  S  3,  C  9.1 


Family 

Dentition :  i  j,  c  ^,  p  f ,  in  f ;  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  Avith  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. 
Vertebrse  :  C  7,  D  13,  L  6,  S  3,  C  27. 

Tarsius* — The  family  contains  the  single  genus  Tarsius,  of 
which  but  one  species  is  known,  T.  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  at  a  time.3 


Family  CHIROMYIDJS. 

Dentition  of  adult  :  i  \,  c  $,  p  £,  in  f  ;  total  1  8.  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  f  ,  c  %,  m  f  ,  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~ 
kundige  Onderzock  van  den  Potto  van  Bosman,  1859)  for  P.  caldbarensis,  Huxley, 
Proc.  Zool.  Soc.  1864,  p.  314.  "  Storr,  Prodromus  Meth.  Mamm.  (1780). 

3  H.  Burmeister,  Beitriige  ~ur  niihrercn  Kcnntnissder  gattung  Tarsius,  1846. 


CHIROMYID& 


695 


developed  and  distinct  from  the  tibia.  All  the  digits  of  lx)th  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,  S  3,  C  27. 

Chiromys.1 — This  family,  like  the  last,  is  formed  for  the  recep- 
tion of  a  single  genus,  Chiromys,2  containing  one  species,  C.  mada- 
gascarienxis,  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  its  large  opposable 
hallux,  with  a  flat 
nail,  but  all  the 
other  toes  have 
pointed  compressed 
claws,  like  that  of 
the  second  toe  in 
the  Lemurince  and 

,i  -i         i  ,1.    i      FIG.  331.— Skull  of  Ave-aye  (C/uVowi/s  mailagascariensis).     X* 

the  second  and  third  )Ius  g^ Coll  s^rgeons: 

in  the  Tarsiidce.  Tail 

long  and  bushy.  General  colour  dark  brown,  the  outer  fur  being 
long  and  rather  loose,  with  a  woolly  undercoat.  Mammae  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 

1  Cuvier,  "Table  de  Class."  in  Ltsons  d'Anat.  Comp.  vol.  i.  (1800). 

'-  It  was  first  named  Daubeitionia  by  Geoffrey ;  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  PRIM  A  TES 

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.1 

E'i'md  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 
Ccenopithecus  lemuroides,  regarding  the  animal  to  which  the  specimen 
belonged  as  partaking  of  the  characters  both  of  the  Lemurs  and  the 
American  Monkeys.  Most  other  palaeontologists  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  CcenopithecMs  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  Adapts  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,  A,  which  was  originally  described  as  Necrolemur 
antiquus,  but  appears  to  be  generically  identical  with  Microchcerus 

1  R.  Owen,  "On  the  Aye-aye,"  in  Trans.  Zool.  Soc.  1862,  vol.  v.  p.  33  ; 
W.  Peters,  "Ueber  die  Slitigethier - Gattung  Chiromys,"  in  Abhand.  Konigl. 
Akad.  dcr  Wissenscliaftcn,  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  f- ,  c  |,  p  f,  m  $-, 
or  the  same  as  in  the  existing  Tarsim  ;  but  it  is  not  improbable  that 
in  some  instances  the  first  lower  premolar  may  have  been  developed. 
The  upper  molars  of  M.  erinaceus  differ  from  those  of  M.  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  Avhich  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.  nltmriJtti  of  the    (C^^Fl^^n^Slv  $      '' 
same  deposits  conies 

nearer  in  dimensions  to  FIG.  332.-The  last  nve  right  uP1^r  cheek-teeth  of  3/tcro- 
...  .  _  chcerus  anlvptus  (A)  &nd  Mtcrochcerus  ennaceus(B).  Twice 

M.  enmceUS.       Ihe  Upper    naturai  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  Anaptomorphus,  from  the  North  American  Eocene, 
has  a  skull  of  about  the  same  size  as  that  of  the  smallest  species  of 
Microchcems,  but  the  dental  formula  is  i  -H-,  c  \,  p  f ,  m  f ,  and  the 
upper  molars  are  of  the  tritubercular  type. 

The  well-known  Adapis  (Aphelotherium  or  Palceolemur),  of  the 
Upper  Eocene  of  France  and  England,  differs  from  all  existing 
Lemuroids  in  possessing  four  premolars a ;  the  dental  formula  being 
*  f »  c  T>  P  T»  m  f •  The  fourth  upper  premolar  has  two  outer  cusps, 
and  the  upper  molars  (Fig.  333)  resemble  those  of  Lepidulemur  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.  pansiensis  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  A.  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 
FIG.  m-The  left  upper  cheek-teeth  zoologicai  type  intermediate  between 

of  Adapis  magna,  from  the  Upper  Eocene      ,        ,°  11  i       i 

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  Ccenopithecus  lemuroides 
is  inseparable  from  Adapis  parisiensis,  but  the  postero- internal 
column  of  the  upper  molars  is  said  to  be  larger.  The  genera 
TomitJierium  and  Nothardus,  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  £ ,  c  ^,  p  £, 
m  f.  The  quadrituberculate  upper  molars  have  well -developed 
accessory  intermediate  columns  (protoconule  and  metaconule),  and 
thus  resemble  those  of  Microcliwrus ;  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  Eocene,  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  loAver  molars  are  quin-       FIG.  334.— The  right  upper 
quetubercular,  and   not    unlike   those    of  cheek-teeth  of  Pk*u<dapis  remen- 

i.-.  /,  1,11-  i  r     ^i        sis;  from  the  Lowest  Eocene  of 

Microsyops;    the    dental    formula    of    the  Rheim8.   X|.  3», 3, 4, premolars ; 

lower    jaw   is    i    2,  C  I,  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  j,  c  ^,  p  f ,  m  f .     In  Plesiadapis  the  lower 


ANTHROPOIDEA  699 

and  the  first  upper  incisor  are  enlarged,  the  upper  molars 
(Fig.  334)  tri  tubercular,  and  the  lower  quadritubercular.  Indrodon, 
of  the  lowest  Eocene  of  the  United  States,  resembles  Plesiadapix  in 
its  tritubercular  upper  molars,  and  appears  to  have  a  nearly  similar 
dental  formula.  Mixodectes,  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  Lcmuridce"  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,"  iu 
Phil.  Trans,  vol.  clxvi.  pp.  569-587  ;  F.  Pollen  and  D.  C.  Van  Dam,  Recherckes 
sur  la  Faune  de  Madagascar,  2me  parte,  "  Mammiferes,"  1868.  For  the  fossil 
types  see  M.  Schlosser,  "Die  Affen.,  Lemuren,  etc.,  des  Europaischen  Tertiars," 
in  Beitr.  Pal.  (Estr-Ungar,  1888. 


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  Hapalidce).  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  mammae.  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- 
lidce,  Cebidce,  Cercopithecidce,  Simiidce,  and  Hominidce,  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  PRIM  A  TES 


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 
A  teles  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  Cereopithecinas  both 
pairs  are  short  and  subequal.  Only  in  the  Hapalidce  and  some  of 
the  Cebidce  are  the  legs  proportionately  as  long  as  in  Man. 

The  tail  is  as  much  as  three  times  the  length  of  the  body  in 
Ateles ;  while  in  the  Simiidce  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  Chrysothrix,  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  roxellance  (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  Cebidce  they  are  decidedly  compressed  laterally,  while 
in  the  HapaUda  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  Cercopithecidce  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  diana  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  cedipus  and  in  some  species 
of  Semnopithecus ;  while  in  the  Bonnet  Monkey  (Macacus  sinicus)  it 
radiates  in  all  directions  from  a  central  point  on  the  vertex.  Long 
hair  clothes  the  shoulders  in  Cynocephalus  Juimadryas  and  Hapale 
humeralifer;  and  the  end  of  the  tail  has  a  tuft  in  two  species  of 
Cynocephalus  and  in  Macacus  sinicus.  Many  of  the  African  Colobi 
and  some  species  of  the  Howlers  have  very  long  hair  on  the  flanks ; 
and  in  PitJiecia  this  development  of  hair  extends  to  the  greater  part 
of  the  body  and  the  tail,  P.  satanas  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  Simiidce,  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  Eriodes  and  Macacus  tibetanus  (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  Cercopithecidce  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  PRIM  A  TES 


approximation  to  man,  and  the  same  is  true  with  regard  to  the 
occipital  production,  which  is  most  developed  in  Chrysothrix.  Most 
of  the  Simiidce  exhibit,  however,  a  distinct  convexity  of  the  occiput, 
and  thereby  differ  markedly  from  the  Cercopithedda,  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  Chrysothrix.  There  is  an  external  bony 
meatus  auditorius  in  Man,  the  Simiidce,  and  the  Cetcopithecidce,  but 
none  in  the  Cebidce  and  Hapalidce. 

The  premaxillse  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  Mycetes.  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  Simiidce,  are  generally  replaced  in  the 
Cercopithecidce  by  a  coarse  diploe,  but  they  are  present  in  the 
Cebidce  and  Hapalidce,  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  Simiidce  and  most 
of  the  Cercopithecidce.  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  i  f ,  c  ^,  p  f ,  m  f,  total  32  ;  but  in  the 
Cebidce  the  cheek-teeth  are  p  f ,  m  f ,  and  in  the  Hapalidce  p  f ,  m  f . 
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  Simiidce  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  vertebrae  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 
Cebidce  generally  have  thirteen ;  and  the  same  number  obtains  in 
the  Chimpanzee  and  Gorilla,  while  the  Orang  resembles  Man  in 
having  but  twelve.  The  lumbar  vertebrae  show  a  range  in  number 
of  from  four  to  seven.  In  the  Simiidce  there  are  four  or  five  of 


FIG.  335.— Skeleton  of  the  Black-handed  Spider  Monkey  (Ateles  geo/royi).    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  Ateles. 
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  Simiidce  and  Man,  its  greatest 
absolute  length  occurring  in  the  Gorilla,  and  the  relative  greatest 
length  being  found  in  Hylobates.  The  Simiidce  never  have  less  than 
five,  and  may  have  six  sacral  vertebrae ;  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  vertebrae. 
Except  in  the  Simiidce  and  Macacus  inuus,  the  number  of  caudal 
vertebrae  in  the  Apes  always  exceeds  four,  but  they  may  be 


704  PRIM  A  TES 


reduced  to  five  in  the  Mandrill  (Cynocepliahis  maimon).  In  Macacus 
and  Uacaria  the  shortness  of  the  tail  is  attained  by  the  small 
size  of  the  vertebrae  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  Ateles 
(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 
vertebrae.  Chevron  bones  are  present  in  all  forms  having  a  distinct 
tail ;  and,  together  with  other  processes  for  muscular  attachment, 
attain  their  greatest  development  in  A  teles. 

The  vertebral  processes  known  as  metapophyses  and  anapophyses, 
which  are  generally  inconspicuous  in  Man,  and  are  but  small  in  the 
Simiidce,  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  Cebidce,  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  Simiidce  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  Simiidce  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  Mycetes,  being  remarkably  small  in  Simla. 


ANTHROPOIDEA  705 

The  Cebidce  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  Simiidce  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  Simiidce.  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,1  "  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  Cercopithecidce  the  posterior  cornu  is  relatively 
shorter  than  in  man,  it  again  becomes  elongated  in  the  Cebidce,  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  syndadyhis. 
In  the  latter  the  cerebellum  is  slightly  uncovered,  but  it  is  so  con- 
siderably in  the  former.  In  Chrysothrix  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  Cebidce  it  is  much  larger  relatively  than  it  is  in 
Man,  and  is  absolutely  larger  than  the  hippocampus  major.  Of  all 
i  Article  APE,  Encyclopccdia  Sritannica,  ninth  edition. 
45 


;o6  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  Simla  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  Ateks, 
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  Cebidce  than  in  the  Simiidce  and  Cerco- 
pithecidce.  In  all  Apes  below  the  Simiidce  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  Cebidce." 

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  Cebidce.  The  peculiar  sacculation  of  the  stomach  in  the  sub- 
family Semnopithecince  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  valvulae  conniventes,  and  it  is  only  in  Man  and 
the  Simiidce  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  Simiidce  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.  Hylobtites  has  a 
liver  (Fig.  352)  which  perhaps  is  nearer  to  the  human  than  that  of 
any  of  the  other  Simiidce.  In  the  Cercopithecidce  the  liver  differs 


ANTHROPOIDEA 


707 


from  that  of  the  Simiidce  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  Cebidce  (Fig.  336) 
and  Hapalidce,  in  ad- 
dition to  the  deeply 
cleft  lateral  fissures,  is 
characterised  by  the 
great  size  and  quad- 
rangular form  of  the 
caudate  lobe  (c),  which 
attains  its  maximum 
development  in  A  teles. 
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- 

, .  mi  i  FIG.  336.— Under  surface  of  the  liver  of  the  Black-handed 

OH.          iney     may        e  gpjder  Monkey  (Aides  mdaiiochir).    u,  Umbilical  fissure ; 

dilatations  of   the  laryn-  vc,  vena  cava ;  U,  left  lateral  lobe  ;  fc,  left  central  lobe ;  re, 

geal     ventricle        as       in  "8^*  central  l°De  ;  rl,  right  lateral  lobe  ;  s,  Spigelian  lobe  ; 

7J. .  ,  c,  caudate  lobe  ;  a,  gall-bladder. 

Simla,    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  Cercopithecidce. 
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  axillae,  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-like  structure. 
The  posterior  cronua  of  the  hyoid  (thyro-hyals)  are  never  entirely 
absent,  but  the  anterior  or  lesser  cornua  may  be  so,  as  in  Mycetes. 
The  anterior  cornua  never  exceed  the  posterior  cornua  in  length ; 
but  they  may  be  (e.g.  in  Cercopithecus)  more  largely  developed 
relatively  than  in  Man,  and  may  even  be  jointed,  as  in  Lagothnx. 

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 


;o8  PRIMATES 

Man.  In  Hylobates  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  Simiidce  or  Cercopithecidce,  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  Macacus  inuus  is  found  as 
far  north  as  Gibraltar,  M.  tibetanus  and  Semnopithecus  roxellance 
inhabit  western  Tibet,  while  in  Japan  we  have  M.  speciosus.  In  the 
New  World  one  species  of  Ateles  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  (Semnopithecus)  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 
Simiidce  and  Cercopithecidce  are  exclusively  confined  to  the  Old  World, 
the  Cebidce  and  Hapalidce  are  equally  restricted  to  the  New,  and  we 
may  accordingly  proceed  to  notice  a  few  points  in  relation  to  generic 
distribution.  Of  the  larger  Simiidce  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 
(Anthropopithecus)  and  not  improbably  of  an  Orang  (Simia)  in  Northern 
India.  The  Gibbons  (Hylobates)  are  now  exclusively  Oriental. 
Europe  has  only  Macacus  inuus  of  Gibraltar,  also  found  in  Africa 
north  of  the  Sahara,  and  therefore  strictly  Palsearctic  in  distribu- 
tion. The  Ethiopian  region  includes  in  the  Cercopithecidce  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 


HAPALID^E  709 

of  the  Oriental  region ;  but,  as  already  mentioned,  outlying  species 
extend  into  various  parts  of  the  Palsearctic  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  Chrysothrix,  Nyctipithecus,  Cebus,  A  teles,  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,  Chrysothrix,  and  Callithrix  extend  farthest; 
but  they  do  not  probably  all  extend  to  the  farthest  limit  yet  known, 
namely  30°  S.  The  species  found  farthest  south  are  Mycetes  caraya, 
Cebus  fatuellus,  and  Callithrix  personatus. 

Family  HAPALID.E. 

Dentition  :  if,  c  ^,  p  f ,  m  f ;  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  Cebidce.  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 


7io 


PRIM  A  TES 


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.1 — Lower  canine  not  longer  than  the  incisors.    A  number 
of  species  have  been  described,  among  which  may  be  mentioned 


FIG.  337.— The  Golden  Marmoset  (Midas  ckrysoleucas).    From  Proc.  Zool.  Soc.  1868,  pi.  24. 

H.  jacchus,  H.  albicollis,  H.  aurita,  and  H.  humeralifer.  Remains  of 
species  of  this  genus  have  been  found  in  the  cavern-deposits  of 
Brazil. 

Midas.2 — Lower  canine  considerably  longer  than  the  incisors.  No 
less  than  twenty-four  species  of  this  genus  have  been  named,  among 
which  the  Silky  Marmoset  (M.  rosalia)  of  Columbia,  the  Pinche 
Monkey  (M.  cedipus)  of  South -Eastern  Brazil,  and  the  Golden 
Marmoset  (M.  chrysoleucas,  Fig.  337)  are  well-known  types. 

1  Illiger,  Prodromus  Syst.  Mamm.  et  Avium,  p.  71  (1811). 
2  Geoffrey,  Ann.  du  Museum,  vol.  xix.  p.  120  (1812). 


CEBID& 


Family  CEBID^E. 


Dentition:    i   f ,  c  £,  p  f ,   m  f ;    total   36.     Tail  frequently 
prehensile  ;  digits  with  nails  ;  other  characters  as  in  the  Hapalida. 

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  Hapalidce,  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 
Cercopithecidce  of  the  Old 
World.  The  genera  are 
usually  arranged  in  five 
subfamilies. 

Subfamily  Myeetinse. — 
Lower  incisors  vertical ; 
hyoid  bones  enormously 
inflated  ;  tail  long  and  pre- 
hensile, naked  beneath  at 
the  end ;  pollex  well  de- 
veloped. 

Mycetes.1 — The  sole  re- 
presentatives of  this  sub- 
family are  the  well-known 
Howling  Monkeys,  all  of 
which  are  included  in  the 

genus  Mycetes.  They  are  FIG.  S3S.— Side  view  of  skull  and  hyoid  bone  of  the 
Of  more  bulky  build,  and  Red  Howling  Monkey  (Mycetut  seniculus).  From  De 

have  more  produced  muzzles  B 

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  Illiger,  Prodromus  Syst.  Mamm.  et  Avium,  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  (M.  seniculus) 
and  the  Ursine  Howler  (M.  ur sinus)  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.1 — 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.  hirsute,  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 
Cebidce  by  their  short  Baboon-like  tail.  The  Bald  Uakari  ( U.  calm) 
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 
caecum  of  U.  calva,  according  to  Mr.  F.  E.  Beddard,  measures 

1  Geoffroy,  Ann.  du  Museum,  vol.  xix.  p.  115  (1812). 

2  Gray,  Proc.  Zool.  Soc.  1849,  p.  9.     Amended  from  Ouakaria :  Syn.  Brachy- 
urus;  Spix,  Sim.  et  Vesp.  JSrasil,\p.  11  (1823).     Preoccupied  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  caecum  is  also 
found  in  Callithnx  and  PitJwcia. 

Subfamily  Nyetipitheeinse. — Lower  incisors  vertical ;  hyoid 
normal ;  tail  long,  non-prehensile ;  pollex  well  developed. 

Three  genera  are  included  in  this  subfamily,  the  species  being 
partly  insectivorous. 

Callithrix.1 — Head  small,  depressed,  and  not  elongated;  nares 


Fio.  339.— The  Moloch  Teetee  (CallitTirix  moloch).    From  Archives  du  Mustum,  vol.  iv.  pi.  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  Nyctipitliecus ;  while  the 
1  Geoffrey,  Ann.  du  Mustum,  vol.  xix.  p.  112  (1812). 


714  PRIMATES 


small  canines  and  the  bushy  tail  readily  mark  their  distinction  from 
Chrysothrix.  Remains  of  Callithrix  have  been  found  in  the  Brazilian 
caves. 

Chrysothrix.1 — 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 


PIG.  340. — The  Lemurine  Douroucouli  (Nyctipithecus  lemurinus).    From  Archives  du  Museum, 

vol.  iv.  pi.  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.2 — 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. 

1  Kaup,  Thierreich,  vol.  i.  p.  51  (1835). 
2  Spix,  Sim.  et  Vesp.  Brasil,  p.  25  (1823). 


CEBID&  7 1 5 

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  (N.  trivirgatus) 
continue  to  the  crown,  and  in  the  other  (N.  felinus)  meet  on  the 
top  of  the  forehead.  N.  trivirgatus  was  first  described  by  Humboldt, 
who  discovered  it  on  the  banks  of  the  Cassiquiare,  near  the  head- 
quarters of  the  Rio  Negro." 

Subfamily  Cebinse. — 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. 

Ateles.1 — 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  A.  melanochir  (Fig.  341),  A.  paniscus  of  Guiana, 
A.  geoffroyi  of  Central  America,  A,  ater  of  Eastern  Peru,  and 
A.  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  Geoffrey,  Ann.  du  Museum,  vol.  vii.  p.  260  (1806). 
2  I.  Geoffroy,  Diet.  Class,  vol.  xv.  p.  443  (1829). 


7i6 


PRIM  A TES 


nails.  The  species  are  E.  arachnoides,  E.  hemidactylus,  and  E. 
hypoxanthus. 

Lagothrix.1 — 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  Erwdes  in 
their  fur  and  compressed  nails,  but  differ  in  the  more  widely 


FIG.  341.— The  Black-handed  Spider  Monkey  (Aides  mdanochir). 
From  Proc.  Zool.  Soc.  1871,  pi.  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  Geoffrey,  Ami.  du  Museum,  vol.  xix.  p.  106  (1812). 


CEBID^E 


71? 


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 


Fio.  342.— Humboldt's  Lagothrix  (LagotKrix  humboldti).    From  Proc.  Zool.  Soc.  1863,  pi.  31. 

account  of  the  excellence  of  its  flesh  as  food."  Five  species  are 
usually  recognised,  viz.  L,  canus,  L.  humboldti,  L.  castelnaui,  L. 
tschiidii,  and  L.  geoffroyi. 

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.  Rlgne  Animal,  p.  44  (1777). 


7i8 


PRIM  A  TES 


At  least  twenty  species  have  been  described  (C.  fatuellus,  C.  lunatus, 
C.  capucinus,  C.  albifrons,  C.  hypoleucus,  etc.),  but  it  is  probable  that 
some  of  these  are  not  entitled  to  stand,  since  there  is  a  large 


FIG.  343.— The  White-cheeked  Sapajou  (Cebus  lunatus).    From  Proc.  Zool.  Soc.  1865,  pi.  45. 

amount  of  individual  variation.     Fossil  remains  of  species  of  Cebus 
have  been  described  from  the  Pleistocene  cavern-deposits  of  Brazil. 


Family  CERCOPITHECID^E. 

Dentition:  i  f-,  c  \,  p  -I,  m  f ;  total  32.  Crowns  of  molars  elon- 
gated an tero- 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.  Sternum  narrow. 
Caecum  without  vermiform  appendage. 

This  family  includes  all  the  Old  World  Apes,  with  the  excep- 
tion of  the  Simiidce,  and  may  be  divided  into  the  subfamilies  Cerco- 
pithecince  and  Semnopithecina'. 

Subfamily  Cereopitheeinae. — Pelvic  and  pectoral  limbs  approxi- 
mately equal;  tail  variable;  cheek -pouches  present;  stomach 
simple. 


CERCOPITHECID&  7 1 9 

This  subfamily  comprises  the  African  Baboons,  the  common 
Indian  Monkeys  constituting  the  genus  Macacus,  together  with  the 
African  Cercopitkecus  and  Cercocebus  and  a  few  allied  types. 

Cynocephalus.1 — Muzzle  much  elongated  (Fig.  344),  with  the 
nostrils  terminal ;  ischial  callosities  very  large ;  tail  more  or  less 
short ;  muzzle  swollen  by  enlargement  of  the  maxillae.  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, 


FIG.  344. — Skeleton  of  the  Chacma  Baboon  (Cyiwceplialus  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." 2  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  Lacepede,  "  Nouv.  tabl.  nieth."  (1799)  in  Mim.  de  TInstitut,  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, 
etc.     Also  a  large  overgrown  Ape  and  Baboon,  so  called.'     'Drill '  is  used  in  the 
same  sense  in  Charl ton'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  PRIM  A  TES 

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.  It  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 
etre  plus  hideux  que  le  Mandrill." 


CERCOPITHECID&  72I 

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 


Fio.  345. — The  Yellow  Baboon  (Cynocepkaltis  babuin).    From  Archives  du  Museum, 
vol.  ii.  pi.  34. 

La  Mtnagme  du  Museum  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.  leucophwus),  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.  babuin),  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  PRIM  A  TES 


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.  fakoneri. 

Theropithecus.1 — Distinguished  from  Cynoceplialus  by  the  nostrils 
not  being  terminal,  but  situated  as  in  Macacus.  This  genus  is 
represented  by  the  Abyssinian  Gelada  (T.  gelada)  and  the  allied 
T.  obscurus. 

Cynopithecus.- — The  Black  Ape  of  Celebes  (C.  niger)  forms  a 


FIG.  346. — The  Tibetan  Macaque  (Macacus  tibetamis).    From  Milne-Edwards,  Recherclies  des 
Mammifercs,  pi.  ?A, 

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  maxillae,  as 
in  some  of  the  species  of  Cynocephalus.  The  muzzle  is  also  more 
produced  than  in  Macacus. 

Macacus.3 — 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  (M.  inuus)  of  Northern 
Africa  and  Gibraltar,  the  Macaques  are  now  exclusively  Asiatic, 
one  species  (Fig.  346)  occurring  in  Tibet,  and  another  (M. 

1  I.  Geoffrey,  Arch,  du  Musium,  vol.  ii.  p.  576  (1841). 

-  I.  Geoffroy,  Voyage  de  Bdangcr,  p.  66  (1834). 
3  Lacepede,  Mtm.  dc  Vlnstitut,  vol.  iii.  p.  450  (1801).     Amended. 


CERCOPJTHECW.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  Inuux 
is  impracticable.  In  M.  inuus  the  tail  is  wanting  ;  in  M.  UManus 
(Fig.  346)  and  M.  nemestrinus  of  Tenasserim  it  is  short ;  in  the 
common  Bengal  Monkey  (.17.  rhesux)  it  is  about  one-half  the  length 
of  the  head  and  body,  while  in  M.  cyrumolgw  and  its  allies  it  is 
still  longer.  In  the  Indian  Lion -tailed  Monkey  (M.  silemis)  it  is 
tufted  at  the  end. 

The  following  summary  of  the  habits  of  the  Macaques  is  taken 
from  Mr.  W.  T.  Blanford's  M«iiiin<ik  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  Semnopitheci.  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  M.  rhesus  is  found  in  the  Himalaya  at  an 
elevation  of  over  8000  feet. 

Fossil  remains  of  Macacus  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  Aulaxinuus.  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.1 — An  African  genus  agreeing  with  Macacus  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    AVhite- eyelid    Monkeys,    and     include 

1  Geoffrey,  Ann.  du  Museum,  vol.  six.  p.  97  (1812). 


724 


PRIMATES 


C.    collari*,    C.  fulvj'titoxus,    ('.   aihioji*,  and    C.  albigena  ;  all    being 

from  West  Africa. 

CercopitJwcus.1 — Muzzle  more  or   less  short ;   ischial   callosities 

moderate ;  tail  long ;  no  talon  to  third  lower  molar.     Build  more 

slender  than  in  Macacus.     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  (6*. 
cattitrichus),  the  Grivet 
(C.  griseo-viridis),  the 
Vervet  (C.  lalandi),  the 
Pluto  Monkey  (C.pluto, 
Fig.  347),  the  Patas 
(C.  ruler),  the  Diana 
Monkey  (C.  diana\  and 
the  Mona  Monkey  (C. 
mona}  are  well-known 
types. 

Subfamily  Semno- 
pitheeinse. 2  -  -  Pelvic 
limbs  longer  than  the 
pectoral;  tail  very 
long;  no  cheek- 
pouches;  stomach  sac- 
culated.  Build  slender. 
This  subfamily  is 
represented  by  three 
genera,  of  which  one  is 

Fio.347.-The  Pluto  Monkey  (Cercopithecus  pluto).    From       African    and    **?*"? 
Gray,  Proc.  Zool.  Soc.  1848,  p.  57.  Asiatic.         Mr.    W.    T. 

Blanford,  in  his  Mam- 
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  Colobince. 


CERCOPITHECIDsE 


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 
two  longitudinal  muscular  bands, 
corresponding  in  situation  to  the 
greater  and  lesser  curvatures  of 
an  ordinary  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 
O3sophageal  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  lai'ge  sac, 
slightly  constricted  on  its  under 
surface  by  the  prolongation  of 
the  inferior  longitudinal  band,  F'«- 348—uterai  view  of  the  skull  and  palatal 

1     . .  .   '    aspect  of  the  cranium  of  Semnopithecus  nemceus. 

or    that    corresponding   to    the  (From  De  Biainviii«>.) 
great  curvature.  The  oesophagus 

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  cesophageal 
entrance. 

Nasalis.1 — Skull  resembling  that  of  the  Cercopitheclna  in  that 

the  lower  border  of  the  nasal  bones  extends  considerably  below  the 

lower  border  of  the  orbits,  whereas  in  the  other  SenmupUhecma  the 

aperture  of  the  nares  extends  upwards  between  the  orbits.     Nose 

1  Geoffrey,  Ann.  du  Mustum,  vol.  xix.  p.  90  (1812). 


726 


PRIM  A  TES 


produced  into   a  large  proboscis.     Other   characters  as   in  Semiw- 
pithecus. 

This  genus  includes  only  the  Proboscis  Monkey  (X.  lurrtifus)  of 
Borneo,  remarkable  for  the  great  prolongation  of  the  nose  in  the 
adult.  In  young  animals  the  nose  is  relatively  much  shortei*,  and 


FIG.  340. — Scmnopithccus  rojcellance.    (From  Milne-Edwards,  KechercTies  des 
Mammifkres,  pi.  36.) 

bent  upwards  after  the  manner  of  that  of  Semnopithecus  roxettance 
(Fig.  349). 

Semnopitliecus.1 — 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.  roxellance2  (Fig.  349), 

1  F.  Cuvier,  Hist.  Nat.  des  Mammiferes  (1821),  "Semno-pitheque." 
2  Separated  generically  by  some  writers  as  Rhinopitliccus. 


CERCOPITHECID.  A  727 

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  Nasalis  lan-nfii*.  The  genus  is  represented 
in  India  and  Burma  by  no  less  than  fourteen  species,  of  which  the 
common  Indian  Langur,  or  Hanuman  Monkey  (S.  entellus)  and 
the  larger  Himalayan  Langur  (S.  schistaceus)  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  which  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  S.  schistaceus  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  Semnopithecus  occurs  in  the 
Pleistocene  and  Pliocene  of  India,  and  it  has  also  been  recorded 
from  the  Pliocene  of  France  and  Italy. 

Colobus.1 — This  African  genus  differs  from  Semnopithecus  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  beautiful  mantle  of  long  silky 
hair  which  hangs  down  from  each  side  of  the  body,  and  for  their 
tufted  tails.  In  ('.  i/n<'ir:>(.  from  Abyssinia  these  are  white,  and  the 
rest  of  the  body  and  limbs  black.  Others  (as  ('.  .•o/fnini*)  are  entirely 
black.  The  skins  of  the  long-haired  species  are  largely  imported 
into  Europe  for  the  manufacture  of  ladies'  muffs,  etc. 

Extinct  Genera. — Certain  types  of  Apes  from  the  European 
Tertiaries  indicate  genera  referable  to  the  Cercopithtcida,  but 
distinct  from  any  of  those  now  living.  Of  these  Mesopithecus,2  from 
the  Lower  Pliocene  Pikermi  beds  of  Attica,  is  known  by  almost 
complete  skeletons,  and  resembles  M<n-tini.<  in  the  shortness  and 
stoutness  of  the  limbs,  but  agrees  with  Semnopithecus  in  the  characters 
of  the  skull  and  teeth.  An  allied  Monkey  from  the  Lower  Pliocene 
of  Perpignan,  in  France,  differs  from  MesopiihffiUS  pent ''I  id  by  its 
superior  size,  proportionately  more  produced  muzzle,  and  larger 

1  Illiger,  Prodi-omits  Syst.  Mamm.  et  Avium,  p.  69  (1811). 
-  Wagner,  GWeftrte  Anzcigen,  vol.  viii.  No.  38,  p.  310  (1839). 


728  PRIMATES 


hind  talon  to  the  last  lower  molar;  it  has  been  described  under 
the  name  of  Dolichopithecus.1 

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  Cercopithecidce  and  Simiidce.  According 
to  Dr.  Ristori,3  it  resembles  the  former,  especially  Ci/nocephalus  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  Simiidce,  of  which  it  may  have  been  an  ancestral  type. 

Family  SIMIID^E. 

"  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.  Caecum  with  vermiform 
appendage.  Centrale  of  carpus  sometimes  absent.  Other  characters 
as  in  Cercopitheciclce. 

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. 

Hylobates.* — 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  (H.  syndadylus) 
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  Cercopithecidce.  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  Siamang  (H.  syndadylus), 
which  attains  a  height  of  3  feet,  and  has  been  generically 
separated  by  some  writers  as  Siamanga.  It  is  remarkable  as 

1  Deperet,  Comptcs  Rendus,  vol.  cix.  p.  982  (1889) ;  see  also  AKm.  Soc.  Gtol. 
France,  "Palseontologie,"  vol.  i.  (1890).  2  Gervais,  Cmnptes  Rciidus,  vol. 

Ixxiv.  p.  1217  (1872).  3  Scimmie  Fossili  Italiane,  Boll.  Comm.  Geol.  1890. 

4  Illiger,  Prodromus  Syst.  Mamm.  ct  Avium,  p.  67  (1811). 


SIMIID^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 
(//.  hoolock),  ranging 
from  the  South  of 
Assam  through 
Sylhet  and  Cachar  to 
the  Irawadi  Valley 
near  Bhamo,  the 
White-handed  Gib- 
bon (H.  lar,  Fig. 
350),  which  is  found 
in  Tenasserim  and 
throughout  Malay- 
ana,  the  Dun-coloured  Gibbon  (//.  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 
bamboos  or  branches  that  bend  beneath  their  weight,  and  allow 
them  to  drop  until  they  can  seize  the  ends  of  other  bamboos  or 


Fio.  350.— The  White-handed  Gibbon  (Hylobates  lar).    From 
Blanford,  Mammals  of  British  India,  p.  8. 


730 


riU  MATES 


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 


- 


FIG.  351. — The  Dun-coloured  Gibbon  (llylobatts  entelloides).    From  Arehiv.  du  Museum, 
vol.  ii.  pi.  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'  eggs,  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 


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  ri. 

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  shoAv  the  general 
absence  of  lateral 
fissures  and  the 
small  size  of  the  caudate  lobe  (c)  characteristic  of  the  liver  of  all  the 
^imiidce,  except  Gwillti  (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  Piiopithecus,  indicates  an  extinct  Gibbon  which  does 
not  appear  to  be  generically  separable  from  Hylobates. 

Simla.1 — 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  vertebra?,  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,2  of  which 
there  is  probably  only  a  single  species  (S.  satyrus).     These  animals 
1  Linn.  Syst.  Nat.  12th  cd.  vol.  i.  p.  34  (1766). 
-  A  Malay  word,  signifying  "  Man  of  the  Woods." 


FIG.  862.— Under  suilaec  of  tin-  liver  of  llylolates lar.  v,  Umbili- 
cal fissure ;  p,  jxjrtal  ttssure ;  vc,  vena  cava  ;  I,  left  lobe ;  r,  right 
lobe  ;  »,  Spigelian  lobe  ;  c,  caudate  lobe  ;  g,  gall-bladder. 


732 


PRIM  A  TES 


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 
Gardens  at  Calcutta.  The 
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  Simla  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  vertebrae  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 
AnthropopithecMS  in  the  number  of  ribs,  it  differs  in  the  more 


FIG.  353. — Side  view  of  the  skull  of  adult  Orang  (Simia 
satyrus).    From  Trans.  Zool.  Soc.  vol.  i.  pi.  53. 


SIMIID^E 


733 


important  characters  of  that  of  the  whole  series  of  trunk-vertebrae.1 
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 


Fio.  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  vertebra1,  and  has  thus  given  rise  to  the  statement  that  the  number  of 
vertebra  in  the  Orang  is  the  same  as  in  Man. 


734 


PR  IMA  TES 


and  the  molars  are  characterised  by  the  complex  structure  of  their 
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.1 — 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  ; 
no  ischiatic  callosities.  Male  much 
larger  than  female,  and  Avith  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  5^  feet.  The  first  intro- 
duction of  this  animal  to  the  notice 
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 
1  I.  Geoffrey,  Comptes  Mendus,  vol.  xxxiv.  p.  84  (1852). 


FIG.  355.— Skeleton  of  the  Gorilla. 
(From  De  Blainville.) 


SIMIIDsE 


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 


FIG.  3515.— Tho  Gorilla  (Gorilla  savagei).     From  Trans.  Zool.  Hoc.  vol.  iv.  pi.  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  vertebrae  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  PRIM  A  TES 


number  of  the  dorso-lumbar  vertebrae,  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. 

Anthropopithecus.1 — 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  (A.  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  denned  A.  calms  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,  Lemons  Orales  (1839).  The  Chimpanzees  have  been  very 
generally  described  under  the  name  of  Troglodytes,  but  since  this  name  is 
preoccupied  for  a  genus  of  birds,  it  is  incumbent  to  follow  the  strict  rule,  and 
adopt  the  name  Anthropopitliccus,  although  both  the  present  witers  have 
elsewhere  expressed  the  opposite  opinion. 


SIMIID^E 


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 


Fio.  357. — The  Chimpanzee  (Anthropopithecus  troglodytes).    From  Mr.  Wolfs  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  PRIM  A  TES 

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  Palceopithecus,  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  A.  sivalensis  makes  a  nearer  approxima- 
tion to  the  human  type  than  is  the  case  with  its  living  congeners. 

Dryopithecus.1 — 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,2  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  Cercopithecidce.  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  JRendus,  vol.  xliii.  p.  219  (1856). 
2  Mim.  Soc.  Geol.  France,  "  Palaeontologie, "  vol.  i.  Mem.  No.  1  (1890). 


HOMINID^E  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  HOMINIDYE. 

In  the  Systema  Naturce  of  Linna3us  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  Rkgne  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  his  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,1  is  to  revert  towards  the  Linnaean 
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  Vertcbratcd  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  d' Anthropologie. 


740  PRIM  A  TES 

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  Hominidce.  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  Hominidce  and  Simiidce  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 
Linnaeus,  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& 


— (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 
palaeontological  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.1 

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  PRIM  A  TES 

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  Linnaeus  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  Linnaeus — the  European, 
Asiatic,  African,  and  American.  These  Were  expanded  into  five 
by  Blumenbach  by  the  addition  of  the  Malay,1  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. 


HOMINIDJE  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 


HOMINID&  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  PRIM  A  TES 

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^E  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- Malay  an  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. 

a.  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^E  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  Caesar.  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 beingal ways  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.1  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. 


HOMINID^E  753 


Labrador,  and  Nova  Scotia,  or  the  occasional  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  both  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,  EUments  d'Anthropologie  Gentrale,  1885  ;  A.  de 
Quatrefages,  Histoire  Gdnerale  des  Races  Humaines  (1.  Questions  Gentrales,  1887  ; 
2.  Classification  des  Races  Humaines,  1889) ;  Quatrefages  and  Hamy,  Crania 
Ethnica  (1873-1879) ;  D.  G.  Brinton,  Races  and  Peoples,  1890. 


INDEX 


AARD-WOLF,  540 
Aard-Vark,  211 
Absorbent  system,  63 
Acanthoglossus,  125 
Acanthomys,  476 
Aceratheriiim,  411 
Achcenodon,  292 
Achyrodon,  114 
Acrobates,  155 
Acrotheriiim,  440 
Adapts,  697 
Adapisorex,  634 
Adapisoricidce,  634 
Adapisoriculus,  634 
Addax,  345 
^idewote,  339 
Adinotherium,  440 
jElurictis,  524 
jElurodon,  562 
^Eluroidea,  501 
jEluroptts,  560 
Mlurus,  562 
dSpyceros,  341 
JEpyprymniis,  164 
Agabelus,  260 
Agouti,  488 
Agrioducrus,  293 
Ai,  182 
Air-sacs,  68 
Alactaya,  480 
Albinism,  10 
Alcelaphus,  334 
^Zces,  326 
Allantois,  77 
AUodon,  111 
^ttqps,  413 
Allotheria,  109 
Alpaca,  303 
^•1  mblotherium,  114 
Amblypoda,  436 
Amorphochilus,  666 
Amphictis,  539 
Amphicyon,  555 


Amphidozotherium,  634 

Archizonurus,  157 

Amphilestes,  114 

Arctictis,  534 

Amphiperatherium,  135 

Arctocebiis,  693 

Amphisvrex,  628 

Arctocephalus,  595 

Amphitherium,  114 

Arctocyon,  609 

Amphitragulus,  330 

Arctocyonidce,  609 

Aviynodon,  412 

Arctogale,  533 

Anaptomorphus,  697 

Arctoidea,  556 

Anchilophus,  376 

Arctomyinse,  454 

Anchippodus,  441 

Arctomys,  454 

Anchitherium,  376 

Arctonyx,  574 

Ancylopoda,  413 

Arctotherium,  561 

Ancylotherium,  413 

Argali,  355 

>4woa,  361 

Armadillo,  195 

Anomaluridai,  449 

Artibeus,  676 

Anomalurus,  449 

Artiodactyla,  275 

Anoplotheriidce,  293 

^lr!>icofe,  466 

Anoplotkerium,  294 

Arvicolinae,  465 

Anteater,  191 

Ass,  383 

Scaly,  205 

Atalapha,  663 

Antebrachium,  47 

^Ke^e*,  715 

Antechinomys,  139 

/KAerwra,  487 

Antelopes,  334 

Auchenia,  298 

Anthops,  657 

^litZocorfzw,  483 

Anthorhina,  674 

Aulaxinuus,  723 

AnthracotheriidcB,  292 

Aurochs,  367 

Anthracotherium,  292 

Australasian  region,  102 

Anthropoidea,  699 

Avahis,  686 

Anthropopithecus,  736 

Axis,  320 

Antilocapra,  333 

Aye-aye,  695 

Antilocapridce,  333 

Antilope,  340 

Babirusa,  287 

Antlers,  308 

Baboon,  719 

.4«<roz0MS,  661 

Bachilherium,  307 

AnuroswKK,  626 

Badger,  575 

Aoudad,  356 

American,  576 

Apar,  199 

Sand,  575 

Ape,  699 

fialccna,  236 

Aphdoj)s,  411 

Balcenidce,  234 

Aphdotherium,  697 

Balainodon,  251 

Archcelurus,  524 

Balamoidea,  234 

Archaeoceti,  246 

Bcdasnoptera,  242 

Archceomys,  484 

tialamotus,  240 

756 


INDEX 


Bandicoot,  141 

Calotragus,  339 

Chamois,  349 

Banteng,  365 

Camel,  296 

C'hampsodelphis,  259 

Bassaricyon,  566 

Camelidce,  295 

Cheeta,  523 

Bassaris,  566 

Camelus,  296 

Chevrotain,  305 

Bats,  641 

Uanidce,  544 

Water,  306 

Bathyergus,  478 

Cants,  546 

Chilonycteris,  672 

Bdeogale,  537 

Copra,  352 

Chimarrogale,  626 

Bear,  558 

Capreolus,  327 

Chimpanzee,  736 

Beaver,  458 

Capromys,  482 

Chinchilla,  487 

Beisa,  343 

Capybara,  491 

Chinchittidae,  487 

Beluga,  262 

Caracal,  518 

ChirogaUus,  689 

Berardins,  256 

Cardiatherium,  491 

Chiromdes,  669 

Bettongia,  163 

Cardiomys,  491 

Chiromyidce,  694 

Bharal,  356 

Cariacus,  329 

Chiromys,  695 

Bibos,  360 

Caribou,  324 

Chironectes,  134 

Bighorn,  355 

Carnivora,  496 

Chiroptera,  641 

Binturong,  534 

Carottia,  674 

Chiru,  341 

Bison,  362 

Carponycteris,  654 

Chiruromys,  476 

Black-Fish,  269 

Carpus,  48 

Chlamydoj)horina;,  196 

Bladder,  69 

Carterodon,  484 

CMamydophorus,  196 

Blarina,  624 

Cos<w,  457 

Chlamydotlierium,  201 

Blastomeryx,  330 

Castoridce,  457 

Chceronycteris,  674 

Blaubok,  343 

Castoroididce,  488 

Chceropotamida:,  292 

Blessbok,  335 

Castoroides,  488 

Chceropotamus,  292 

Blood,  63 

Cat,  517 

Chceropsis,  280 

Bolodon,  111 

Cawa,  489 

Chceropus,  143 

Boneia,  653 

Caviidce,  489 

Cholcepus,  182 

Bontebok,  334 

Cavy,  490 

Chorion,  77 

Bosch-Vark,  286 

Cayluxotherium,  621 

Chrysochloridce,  638 

Boselaphus,  345 

Cebidce,  711 

Chrysochloris,  639 

Bothriolabis,  291 

Cebochcerus,  292 

Chrysothrix,  714 

Bottlenose,  253,  270 

(7efo*s,  717 

Cimoliomys,  113 

Bovidce,  334 

Cement,  15 

Circulation,  63 

Brachium,  47 

Centetes,  637 

Civet,  526 

Brachyphylla,  675 

Centetidce,  637 

Palm,  532 

Brachytarsomys,  465 

Centurio,  676 

Classification,  84,  88 

Brachyurus,  712 

Cephalogale,  562 

Claviglis,  460 

Bradypodidae,  179 

Cephalophus,  338 

Claws,  12 

Brady  pus,  181 

Cephalorhynchus,  266 

Coati,  566 

Brain,  69 

Cephalotes,  653  . 

Ctofow,  339 

Bramatherium,  333 

Cercocebus,  723 

Cododon,  184 

Brocket,  330 

Cercoleptes,  567 

Galops,  658 

Brontotherium,  413 

Cer  corny  s,  483 

Cbfiria,  250 

Bruta,  176 

CercopitJiecidce,  71  8 

Coleura,  667 

Bubalus,  361 

Cercopithecus,  724 

Colobus,  727 

Budorcas,  351 

Cerivoula,  664 

Colour,  8 

Buffalo,  361 

Cervalces,  327 

Comphotherium,  621 

Bush-dog,  553 

Cervicapra,  340 

Condylarthra,  438 

Cervidce,  313 

Condylura,  630 

CACHALOT,  249 

Cervince,  316 

Conepatus,  574 

Cadurcotheriiim,  412 

Cervulus,  316 

Connochcctes,  336 

Caecum,  59 

Cervus,  319 

Contracama,  491 

Ccdogenys,  489 

Cetacea,  225 

Coryphodon,  437 

Ccenopithecus,  696 

•  Cetotheriuin,  245 

Coryphodontidte,  438 

Caenotheriidce,  294 

Chcenohyus,  291 

Cotylophora,  307 

Ccenotherium,  294 

Chcctomys,  486 

Cotylopidce,  293 

Callinycieris,  655 

Chalcochloris,  639 

Cotylops,  293 

Callithnx,  713 

Chalicomys,  458 

Coypu,  482 

Callophoca,  606 

Ckalicotheriidce,  413 

Cranium,  35 

Calomys,  463 

Chalicotherium,  413 

Crassitherium,  223 

Caloprymnus,  164 

Chalinolobus,  662 

Creodonta,  606 

INDEX 


757 


Cricetodipus,  479 

Desmodus,  677 

Echinoyale,  634 

Cricetodon,  464 

Desmotylus,  223 

Echinomys,  483 

Cricetomys,  477 

Diaphragm,  67 

Echinothrix,  477 

Cricetus,  463 

Dicer  at  fieri  um,  411 

Edentata,  176 

Criotherium,  349 

JJicfwbunus,  294 

tfffodientia,  178 

Crocidura,  626 

Dichodon,  294 

Eland,  348 

Crossarchus,  537 

Dichodontidce,  294 

Elaphodus,  318 

Crossopus,  625 

Dididurus,  668 

Elasmognathits,  371 

Crusta  Petrosa,  15 

Dicolpomys,  484 

Elasmotherium,  411 

Cryptophractus,  201 

Dicotyles,  289 

Eleotragus,  340 

Cryptopithecus,  699 

Dicotylidce,  289 

Elephant,  424 

Cryptoprocta,  525 

Didelphia,  128 

Elepkantidce,  423 

Ctenacodon,  112 

Diddphyidce,  133 

Elephas,  424 

Ctenodactylus,  481 

Didelphys,  134 

Eleutherocercus,  203 

Ctenomys,  482 

Ltidymictis,  539 

Eliomys,  459 

Cunictdus,  470 

Digestive  system,  53 

Eliurus,  465 

Cuscus,  149 

Dinictis,  523 

Elk,  326 

Cyclopidius,  293 

Dinoceras,  437 

EUobius,  472 

Cydoturus,  193 

fHnocyon,  556 

Elolherium,  292 

Cyncelurus,  523 

DinomyidcE,  489 

Emballonura,  667 

Cynictis,  537 

IHnomys,  489 

EmhallonuridtK,  666 

Cynocepfudus,  719 

Dinotheriidfe,  435 

Enamel,  15 

C'ynodictis,  555 

Dinotherium,  435 

Enhydra,  570 

Cynogcde,  534 

Dinoziphius,  251 

Enhydriodon,  570 

Cynohycenodon,  608 

Diobroticus,  458 

Enhydrocyon,  562 

Cynoidea,  544 

J)ioplotheriumt  223 

Entomophaga,  178 

Cynomys,  455 

Jtiphylla,  678 

Eohippus,  374 

Cynonycteris,  652 

Diphyodont,  20 

Eomys,  464 

Cynopithecus,  722 

Diplarthra,  275 

Eonycteris,  654 

Cynopterus,  653 

Diplomesodon,  626 

Eotherium,  224 

G'yow,  551 

Dipodidce,  479 

Epiblema,  488 

Cystophora,  605 

Dipodomys,  479 

Epiglottis,  67 

Dipodops,  479 

Epihippus,  374 

Dacrytherium,  294 

JJiprotodon,  171 

Epomophorus,  650 

Dactylomys,  483 

Diprotodontia,  144 

Eporeodon,  293 

Dactylopsila,  152 

Diprotodontidce,  171 

Equidcc,  376 

Damalis,  334 

7>ipws,  480 

JS^wiw,  381 

Daphcenus,  555 

Distcechurus,  155 

Erethizon,  484 

Dasymys,  462 

Dcedicurus,  203 

Ericulus,  638 

Dasypodidce,  194 

Dog,  551 

Erinaceidcc,  619 

UasypodiiuK,  197 

Dolichophyllum,  673 

Erinaceus,  620 

Dasypotherium,  201 

Dolichopithecus,  728 

Eriodes,  715 

Dasyprocta,  488 

Dolichotis,  490 

Ermine,  590 

Dasyproctidcc,  488 

Dolphin,  270 

Eschatius,  303 

Dasypus,  197 

Dorcatherium,  306 

Ethiopian  region,  98 

Dasyuridce,  136 

Dorcopsis,  166 

Eucastor,  458 

Dasyurus,  138 

Dormouse,  459 

Eucelus,  251 

Daubentonia,  695 

Douroucoli,  714 

Eupetaurus,  454 

Deer,  317,  319 

Dremotheriuin,  330 

Eupleres,  538 

Delphinapterus,  262 

Dromatherium,  113 

Euryceros,  346 

Delphinidos,  260 

Itromicia,  154 

Muryurus,  203 

Delphinoidea,  247 

Dryolestes,  114 

Eusmilus,  524 

Delphinus,  271 

Dnjopilhecus,  738 

Eutatus,  201 

Dendrohyrax,  418 

Duck-bill,  120 

Eutheria,  173 

Dendrolagus,  165 

Ductless  glands,  65 

Emtomys,  467 

Dendromys,  463 

Dugong,  221 

Eye,  72 

Dental  system,  13 

Duikerbok,  338 

Dentine,  14 

Duplicidentata,  491 

FALLOW  DEER,  323 

JJeomys,  473 

Felidac,  502 

Dermoptera,  614 

Echidna,  125 

FeZw,  502 

Desman,  629 

Echidnidce,  124 

Felsinotheriuin,  223 

758 


INDEX 


Fennec,  553 

Grisonia,  579 

Hippotiyris,  384 

Fennecus,  553 

Ground  Sloth,  184 

Hippotragus,  343 

Feresia,  270 

Gryphoca,  606 

Holochilus,  464 

Fiber,  470 

Grypotherium,  189 

Holomeniscus,  303 

Flying  Fox,  651 

Guanaco,  301 

Homalodontotherium,  412, 

Lemur,  615 

Guib,  347 

414 

Squirrel,  453 

Guinea-Pig,  490 

Hominidce,  740 

Foot,  52 

&wto,  591 

#omo,  739 

FOSSO,  527 

Gymnobelideus,  154 

Homodont,  22 

Foussa,  525 

Gymnoptychus,  454 

Hoofs,  12 

Fox,  552 

Gymnura,  619 

Hoolock,  729 

Fox-Bat,  651 

Hoplocetus,  251 

Furia,  666 

Habrocoma,  482 

Hoplophoneus,  524 

Furipterus,  666 

Habrothrix,  464 

Hoplophorus,  202 

Hair,  7 

Horns,  310 

Galago,  690 

Halichcerus,  601 

Horse,  382 

Galeopithecidce,  614 

Halicore,  220 

Hunting  dog,  553 

Galeopithecus,  614 

Halicoridce,  220 

Hyaena,  540 

Galera,  579 

Halitheriidce,  222 

Hycenarctus,  561 

Galictis,  579 

Halitherium,  222 

Hycenidce,  540 

Galidea,  538 

Hallomys,  465 

Hycenocyon,  562 

Galidictis,  538 

Hamster,  463 

Hycenodon,  608 

Gaur,  365 

Hapale,  710 

Hycenodontidce,  608 

Gayal,  365 

ffapalemur,  689 

Hydaspitherium,  333 

#aze#a,  341 

Hapalidce,  709 

Hydrocfuerus,  490 

Gelocus,  294 

Hapalotis,  476 

Hydromyinae,  461 

Gemsbok,  343 

Haploceros,  351 

Hydr&mys,  461 

Genet,  528 

Haplodon,  457 

Hydropotes,  328 

Genetta,  528 

Haplodontidce,  457 

Hylobates,  728 

Geogale,  635 

Hare,  492 

Hylomys,  619 

Geographical  distribution, 

Harpyia,  653 

Hyoid,  39 

93 

Harpyiocephalus,  663 

Hyomoschus,  306 

Geological      distribution, 

Harte-beest,  335 

Hyopotamus,  292 

107 

Hearing,  73 

Hyopsodus,  698 

Geomyidce,  478 

Heart,  63 

Hyotherium,  291 

Geomys,  478 

Hedgehog,  620 

Hypertragulus,  307 

Georychus,  478 

Helicophora,  340 

Hypogeomys,  465 

Gerbillinae,  462 

Hdictis,  578 

Hypsiprymnodon,  162 

Gerbillus,  462 

HeliophoUus,  478 

Hypsiprymnodontince,'i62 

Gibbon,  728 

HeUadotherium,  333 

Hypsiprymnopsis,  111 

Gira/a,  331 

Helogale,  537 

Hypsiprymnus,  163 

Giraffidce,  330 

Hemiauchenia,  303 

Hyrachyus,  373 

Glands,  12 

Hemicentetes,  637 

Hyracidce,  415 

Glauconycleris,  662 

Hemiderma,  674 

Hyracodon,  412 

Globicephalus,  268 

Hemigale,  533 

Hyracodontotherium,  439 

Glossonycteris,  674 

Hemigalidea,  538 

Hyracoidea,  415 

Glossophaga,  674 

Hemitragus,  354 

Hyracotherium,  373 

Glutton,  591 

Herpestes,  535 

Hyrax,  417 

Glyptodon,  203 

Herpetocetus,  245 

Hystricidce,  484 

Gly2>todontidce,  202 

Herpetotherium,  135 

Hystricomorpha,  480 

Gnu,  336 

Ifeterocephalus,  478 

Hystrix,  486 

Golunda,  476 

Heterocetus,  245 

Goat,  352 

Heterodont,  23 

IBEX,  353 

Gopher,  478 

Heterohyrax,  418 

Ichneumon,  535 

Goral,  351 

Heteromys,  479 

Icticyon,  553 

Gorilla,  734 

Hipparion,  380 

Ictitherium,  539 

Grampus,  267 

Hippodactylus,  381 

Ictonyx,  579 

Grampus,  270 

Hippohyus,  291 

/rfops,  640 

Graphiurus,  459 

Hippopotamidce,  278 

/ndris,  684 

Greenland  Whale,  236 

Hippopotamus,  278 

Indrodon,  699 

Grimmid,  338 

Hipposiderus,  657 

/wio,  259 

INDEX 


759 


Insectivora,  610 

Liotmnus,  113 

Mastodon,  431 

Intestine,  59 

Listriodon,  291 

Megachiroptera,  650 

Iitni/H,  723 

Liver,  60 

Megaderma,  658 

Ischnoglossa,  674 

Llama,  299,  302 

Megaloglossus,  665 

Isectolophus,  374 

Lobodon,  605 

Megamys,  488 

Issiodoromys,  491 

Loncfieres,  483 

Megaptera,  241 

Ivory,  14 

Lonchoglossa,  674 

Megatheriidce,  183 

Ixacanthus,  259 

Lonchorhina,  673 

Megatherium,  185 

Lophiodon,  373 

Melanism,  9 

JACKAL,  550 

Lophiodontidce,  373 

jV/efes,  575 

Jaguar,  521 

Lophiomeryx,  294 

Mellivora,  576 

Jerboa,  480 

Lophiomyidce,  460 

Melonycteris,  654 

Lophiomys,  460 

Melursus,  560 

KANGAROO,  159 

Lophiotherium,  374 

Menacodon,  115 

Kerivoula  =  Cerivoula, 

Lophocetus,  259 

Meniscoessus,  113 

Kidney,  69 

Lophostoma,  673 

Meniscomys,  454 

Killer,  267 

Loricata,  179 

Meniscotherimn,  439 

Kinkajou,  567 

Loris,  692 

Menodus,  413 

Koala,  156 

Loxolophodon,  437 

Mephitis,  572 

Koalemus,  157 

Lungs,  68 

Merychippus,  380 

Kdbus=C6bust 

Zw<ra,  567 

Merycochcerus,  293 

Koyia  =  Cogia, 

Lycalopex,  552 

Mesodectes,  640 

Kudu,  348 

Lycaon,  553 

Mesohippus,  376 

Kusimanse,  538 

Lymphatics,  65 

Mesomys,  483 

Lyncodon,  590 

MesonychidcK,  609 

Lagenorhynchus,  270 

Lynx,  518 

Mesonyx,  609 

Lagidium,  488 

Mesopithecus,  727 

Lagomyidce,  491 

Macacus,  722 

Mesoplodon,  254 

Lagomys,  491 

Machcerodus,  524 

Mesotaria,  606 

Lagorchestes,  166 

Macrauchenia,  414 

Mesotherium,  440 

Lagostomus,  488 

MacraucheniidoB,  414 

Mesozoic  mammals,  108 

Lagostrophus,  165 

Macroglossus,  654 

Metacarpus,  49 

Lagothrix,  716 

Macrophyllum,  673 

Metamynodon,  412 

Lambdotheriidcc,  413 

Macropodidce,  158 

Metatheria,  128 

Lambdotherium,  413 

Macropodinoe,  164 

Metriotherium,  294 

Langur,  727 

Macropus,  167 

Miaddae,  539 

Lantanotlierium,  618 

Macrorhinus,  606 

Miacis,  539 

Larynx,  67 

Macroscelides,  618 

Microcavia,  491 

Lasionycteris,  661 

Macroscelididce,  618 

Microcebus,  690 

Zatox,  570 

Macrotherium,  413 

Microchoerus,  696 

Leg,  51 

Macrotus,  673 

Microchiroptera,  655 

Lemming,  467 

Malacomys,  462 

Microconodon,  113 

Lemur,  687 

Mammary  glands,  75 

Microgale,  638 

Lemuridce,  683 

Mammoth,  428 

Microlestes,  111 

Lemuroidea,  682 

Man,  739 

Micromeryx,  330 

Leopard,  514 

Manatee,  215 

Micronycteris,  673 

Lepidolemur,  689 

Manatidce,  215 

Microsorex,  624 

Leporidce,  492 

Manatus,  215 

Microsyops,  698 

Leptictidce,  640 

Mandrill,  719 

Microtus,  466 

Leptictis,  640 

Manidce,  204 

J/i<fci*,  710 

Leptobos,  367 

J/awt's,  204 

Milk-teeth,  20 

Leptomeryx,  307 

Manus,  48 

Mimon,  674 

Leptonycteris,  674 

Maral,  322 

Miniopterus,  664 

Leptonyx,  605 

Markhoor,  354 

Mink,  586 

Leptotragulus,  304 

Marmoset,  709 

Miohippus,  376 

/ieptts,  492 

Marmot,  454 

Miosiren,  223 

Lestodon,  189 

Prairie,  456 

Mixodectes,  699 

Leucocyon,  553 

Marsupialia,  128 

Mole,  630 

Limnosyops,  413 

Marten,  580 

Golden,  639 

Linsang,  530 

Maries,  580 

Star-nosed,  630 

Lion,  504 

Mastacomys,  476 

Mole-Rat,  477 

76o 


INDEX 


Molossus,  670 

Narwhal,  261 

Orang,  731 

Monachus,  604 

Nasalis,  725 

Orca,  267 

Monatherium,  606 

Nasua,  566 

OrceWa,  267 

Monkey,  699 

Natalus,  664 

Oreas,  348 

Monodelphia,  173 

Nearctic  region,  102 

Oreodon,  293 

Monodon,  260 

Necrogymnwnts,  621 

Oreopithecus,  728 

Monophylla,  674 

Necrolemur,  696 

Oreotrayus,  339 

Monophyodont,  20 

Necromantis,  679 

Oriental  region,  100 

Moose,  326 

Nectogale,  627 

Ornithodelphia,  117 

Morenia,  484 

Nectomys,  464 

Ornithorhynchidce,  119 

Mormops,  672 

Nemorhcedus,  350 

Ornitkorhynchiis,  119 

MoropuSj  413 

JVeobaloma,  241 

Orohippus,  374 

Morotherium,  413 

Neofiber,  472 

(h-otherium,  374 

Morse,  597 

Neomeris,  266 

Orthaspidotherium,  634 

Moschirme,  314 

Neoplagiaulax,  113 

Ortlwmys,  484 

Moschus,  314 

Xeosorex,  624 

Orycteropodidce,  208 

Moufflon,  356 

Neotoma,  464 

Orycteropus,  208 

Mouse,  475 

Neotragus,  338 

Orya;,  343 

Mouth,  54 

Neotropical  region,  103 

Oryzomys,  463 

Mulita,  201 

Nerves,  71                            . 

Oryzorictes,  638 

Multituberculata,  109 

Nesocerodon,  491 

Otena,  593 

Mungoose,  535 

Nesocia,  475 

Otariidce,  593 

Muntjac,  316 

Nesodon,  439 

Otocyon,  554 

Muridce,  461 

Nesomys,  465 

Otmnys,  462 

Mus,  473 

Nesonycteris,  655 

Otonycteris,  661 

Muscardinus,  460 

Nesotragus,  339 

Otqpterus,  673 

Musk  Deer,  314 

Neurotrichus,  629 

Otter,  568 

Ox,  358 

Nilghai,  345 

Sea,  571 

Rat,  470,  626 

Nimravus,  524 

Ounce,  517 

Musquash,  470 

Noctilio,  668 

Ovaries,  75 

Mustela,  579 

Nostrils,  66 

Owiftos,  357 

Miistelidce,  567 

Notharctus,  698 

Oviduct,  75 

Mycetes,  711 

Nothropus,  183 

(Ms,  354 

Mydaus,  575 

Nothrotherium,  184 

Oxen,  360 

Mylodon,  189 

Notiosorex,  624 

Oxhycena,  608 

My  odes,  467 

Notopteris,  654 

Oxymycterus,  464 

Myogale,  628 

Nototheriidce,  172 

Myolagus,  492 

Nototherium,  171 

PACA,  489 

Myomorpha,  459 

Nyctereutes,  552 

Pachyacanthus,  224 

Myopotamus,  482 

Xycteridce,  658 

Pachydermata,  87 

Myoscalops,  478 

Nycteris,  659 

Pachynolophus,  374 

Afyosorex,  625 

Nycticebus,  691 

Pachyuromys,  462 

Myoxidce,  459 

Nycticejus,  662 

Paciculus,  465 

Myoxus,  459 

Nyctilestes  665 

Palsearctic  region,  97 

Myrmecobiince,  140 

Nyctinomus,  670 

Palceocastor,  458 

Myrmecdbius,  140 

Nyctipithecus,  714 

Palceocetus,  245 

Myrmecophaga,  190 

Nyctitherium,  665 

Paloeoerinaceus,  621 

Myrmecophagidce,  190 

Nyctophilus,  661 

Palceolemur,  697 

Mysarachne,  634 

Palceomanis,  208 

Mystacina,  671 

OCELOT,  521 

Palceomeryx,  330 

Mystacoceti,  234 

Ochetodon,  464 

Palceonycteris,  657 

Mystacops,  671 

Octodon,  481 

Palceophoca,  606 

Mystromys,  462 

Octodontidce,  480 

PalcBopontoporia,  259 

Myxocebus,  689 

Odobcenus,  597 

Palceoprionodon,  539 

Myxopoda,  665 

Odontoceti,  247 

Palceoreas,  348 

Ogmorhinus,  605 

Palceoryx,  344 

NAILS,  12 

Ommatophoca,  605 

Palceospalax,  629 

Nakong,  346 

Onotragus,  339 

Palceosyops,  413 

Nandinia,  534 

Onychogale,  166 

Paiceotapimis,  373 

Nanotragus,  339 

Onychomys,  463 

Palasotheriidce,  375 

Nares,  66 

Opossum,  133 

Palteutherium,  375 

INDEX 


761 


Palceotragoceros,  349 

Phylloderma,  674 

Porpoise,  263 

Palauchenia,  303 

Phyllonycteris,  674 

Potamarckus,  488 

Palhycena,  544 

Phyllophaga,  178 

Potamochaerus,  286 

Palla,  341 

Phyllorhina,  657 

Potamogale,  635 

Palm-Civet,  532 

Phyllostoma,  674 

Potamogalidce,  634 

Paloplotherium,  375 

Phyllostomatidce,  672 

Potamophilus,  534 

Palorchestes,  170 

Physeter,  248 

Potamotherium,  570 

Panda,  562 

Physeteridce,  247 

Potoroinoe,  162 

Pangolin,  205 

Physeterina,  248 

Potoroo,  163 

Panochthus,  203 

Physeterula,  251 

Potorous,  163 

Panther,  514 

Physetodon,  251 

Pouched-Rat,  478 

Pantholops,  341 

Physodon,  251 

Praopus,  201 

Paradox-urus,  532 

Pica,  492 

Prehallux,  49 

Paramys,  457 

Pichiciago,  196 

Prepollex,  49 

Parusorex,  618 

Pig,  282 

Primates,  680 

Peccary,  289 

Pilosa,  179 

Priodon,  198 

Pecora,  307 

Pinnipedia,  592 

Prionodon,  530 

Pectinator,  481 

Pithanotomys,  484 

Priscodelphinus,  259 

Pedetes,  480 

Pithechirus,  477 

Procelurus,  523 

Pediotrar/us,  339 

Pitkecia,  712 

Proboscidea,  418 

PeZea,  339 

Placenta,  75 

Probubalus,  361 

Pellegrinia,  484 

Plagiaulacidce,  113 

Procamelus,  304 

Pelvis,  50 

Plagiaidax,  111 

Procapra,  341 

Pelycodus,  699 

Plagiodon,  483 

Procavia,  417 

Peragale,  143 

Platacanthomyince,  461 

Procoptodon,  170 

Peralestes,  115 

Platacanthonnys,  462 

Procyon,  564 

Peramelidce,  141 

Platanista,  258 

Procyonidce,  562 

Perameles,  142 

Platanistidce,  257 

Prodelphinus,  271 

Peratheriiim,  135 

Platycercomys,  480 

Prodremotherium,  307 

Periptychus,  439 

Platygonus,  291 

Proechidna,  126 

Perissodactyla,  368 

Platyonyx,  188 

Prohalicare,  223 

Perodicticus,  693 

Platyphoca,  606 

Prohycena,  562 

Perognathus,  479 

Platypus,  120 

Prolagostomus,  488 

Pes,  52 

Plecotus,  660 

Promegatherium,  189 

Petauroides,  152 

Plesiadapis,  698 

Promylodon,  190 

Petaurus,  153 

Plesiarct&mys,  457 

Prongbuck,  333 

Petrodromus,  618 

Plesictis,  590 

Prophoca,  606 

Petrogale,  167 

Plesiocetus,  245 

Propithecus,  684 

Petrwnys,  482 

Plesiometacarpalia,  316 

Prorastomatidce,  224 

Phacochcerus,  288 

Plesiosorex,  634 

Prorastomus,  224 

Phalanger,  149 

Plesispermophihis,  457 

Protechinomys,  484 

Phalangeridce,  147 

Pleuraspidotherium,  634 

Proteleidce,  539 

Phalangerince,  149 

Pleurolichus,  479 

Proteles,  539 

Phalanges,  49 

Plexochosrus,  491 

Proterotheriidce,  414 

Phalangista,  149 

Pliauchenia,  304 

Proterotherium,  414 

Phascolarctince,  155 

Pliolagostomus,  488 

Protoadapis,  698 

Phascolarctus,  156 

Pliolophus,  374 

Protokippus,  380 

Phascologale,  139 

Pliopithecus,  731 

Protolabis,  304 

Phascolomyidce,  144 

Poebrotherium,  304 

Protoreodon,  293 

Phascolomys,  145 

Pcecilogale,  590 

Prototheria,  117 

Phascolonus,  146 

Pcecilophoca,  605 

Protoxodon,  440 

Phascolotherium,  114 

Poephagus,  360 

Protragelaphus,  349 

Pkenacodus,  439 

Pogonodon,  524 

Protragoceros,  349 

Phenacomys,  466 

Poiana,  531 

Proviverra,  608 

Phlseomime,  462 

Polecat,  587 

Promverrida;,  608 

Phlceomys,  462 

Polymastodon.,  113 

Pros,  317 

Phloramys,  484 

Polyprotodontia,  133 

Pseudaslurus,  523 

PAoca,  601 

Pimtistes,  259 

Pseudalopex,  552 

Phocasna,  263 

Pontoporia,  259 

Pseudochirns,  151 

Phocanella,  606 

Porcupine,  486 

Pseudois,  355 

Phocidce,  600 

Tree,  485 

Pseudorca,  268 

762 


INDEX 


Pseudorhinolophus,  657 

SABLE,  584 

Sloth,  Ground,  184 

Pseudosciurus,  454 

Saccomys,  479 

Smell,  72 

Psittacotherium,  442 

Saccopteryx,  667 

Sminthopsis,  139 

Pteralopex,  654 

Saccostomus,  477 

Sminthus,  479 

Pterodon,  608 

Sacrum,  43 

Solenodon,  636 

Pteromys,  453 

•Sugra,  341 

Solenodontidce,  635 

Pteropodidce,  650 

Saki,  712 

&>rae,  622 

Pteropus,  651 

Salivary  glands,  55 

Soricidce,  621 

Ptilocercus,  618 

Sambur,  320 

Soriculus,  624 

Ptilodus,  113 

Samotherium,  333 

Sotalia,  272 

Pudua,  330 

Sapajou,  717 

Souslik,  456 

Puma,  520 

Sarcophilus,  137 

Spalatidce,  477 

Putorius,  585 

Scaldicetus,  251 

Spalacopus,  482 

Scales,  11 

Spalacoth'erium,  115 

QUAGGA,  384 

Scalops,  630 
Scapanus,  630 

Spalax,  477 
Spaniotherium,  294 

Scapteromys,  464 

Spemwphilus,  456 

RABBIT,  494 

Scaptochirus,  633 

Sperm  Whale,  249 

Bandicoot,  143 

Scaptonyx,  630 

Spider  Monkey,  715 

Racoon,  565 

Scelidotherium,  188 

Spttogale,  574 

Rangifer,  324 

Schizodelphis,  259 

Spiny  Anteater,  124 

Rasse,  527 

Schizodon,  482 

Spleen,  65 

Rat,  474 

Schizostoma,  673 

Squalodon,  257 

Ratel,  576 

Sciuravus,  457 

Squalodontidce,  257 

Rat-Kangaroo,  163 

Sciuridce,  450 

Squamata,  179 

Red  Deer,  322 

Sciurodon,  454 

Squirrel,  451 

Rehbok,  339 

Sciuroides,  454 

Stegodon,  427 

Reitbok,  349 

Sciuromorpha,  448 

Steneofiber,  458 

Reproductive  organs,  74 

Sciuropterus,  453 

•Stfewo,  271 

Respiratory  system,  '  63 

Sciurus,  450 

Stenoderma,  676 

Rhabdosteus,  259 

Scopophorus,  339 

Stenoplesictis,  539 

Rhachianectes,  241 

Scotophilus,  662 

Stenops,  691 

Rhinoceros,  402 

Scotozous,  661 

Stenorhynchus,  605 

Rhinocerotidce,  402 

Sea-Leopard,  605 

Stereognathus,  110 

Rhinogale,  537 

Sea-otter,  571 

Sternum,  44 

Rhinolophidce,  656 

Seal,  600 

Sthenurus,  170 

Rhinolophus,  656 

Eared,  594 

Stoat,  590 

Rhinonycteris,  658 

Selenacodon,  113 

Stomach,  57 

Rhinophylla,  674 

Semnopithecus,  726 

Strepsiceros,  347 

Rhinopithecus,  726 

Sense  organs,  69 

Sturnira,  676 

Rhinopoma,  669 

Serow,  351 

Stylacodon,  114 

Rhipidomys,  463 

Sheep,  354 

Stylinodon,  442 

Rhithrodon,  464 

Shoulder-girdle,  46 

Styloceros,  317 

Rhithrosciurus,  452 

Shrew,  622 

Stylodon,  114 

Rhizomys,  477 

Tree,  617 

Stypolophus,  608 

Rhizoprion,  257 

Water,  625 

Subungulata,  414 

Rhogeesa,  661 

Siamang,  728 

Swwfo:,  281 

Rhynchocyon,  618 

Siamanga,  728 

Suina,  278 

Rhynchonycteris,  667 

Sight,  72 

Suricata,  538 

Rhytina,  221 

Sigmodon,  464 

SMS,  281 

Rhytinidce,  221 

Simla,  731 

Syllophodus,  484 

Ribs,  44 

Simiidce,  728 

Symtwodon,  413 

River-Hog,  286 

Simocyon,  562 

Synaptomys,  467 

Rock-  Wallaby,  167 

Simplicidentata,  448 

Synetheres,  485 

Rodentia,  443 

Siphneus,  472 

Synotus,  661 

Roe,  327 

Sirenia,  212 

Systemodon,  374 

Rorqual,  242 

Sivatherium,  322 

Rosmarus,  597 

Skeleton,  33 

TAKIN,  351 

Ruminants,  307 

Skull,  34 

TaZpa,  630 

Rupicapra,  349 

Skunk,  572 

Talpidce,  628 

Rytiodus,  223 

Sloth,  180 

rajnawdwrt,  192 

INDEX 


763 


Tamias,  452 

Toxodon,  439 

Urus,  367 

Taphozous,  667 

Toxodontia,  439 

Uses  of  mammals,  4 

Tapir,  371 

Touch,  72 

Uterus,  75 

Tapiridas,  370 

Trachea,  67 

Tapinilus,  294 

Trachyops,  674 

VAMPVRE,  676 

Tapirus,  370 

Trachytfterium,  224 

Vampyrus,  673 

Tardigrada,  178 

Tmgelaphus,  346 

Vertebrae,  39 

Tarsiidce,  694 

Tragoceros,  349 

Vesperimus,  463 

Tarsipedince,  148 

Tragops,  341 

Vespertiliavus,  666 

Tarsipes,  148 

Tragulidce,  305 

Vespertilio,  663 

Tarsius,  694 

Tragulina,  305 

Vespertilionidce,  660 

Taste,  72 

Tmgulus,  305 

Vesperugo,  661 

Tatouay,  198 

Trechomys,  484 

Vicugna,  300 

Tatusia,  200 

Triacanthodon,  113 

Viscacha,  488 

Tatusiince,  200 

Tricenops,  658 

Vishnutherium,  332 

Taxidea,  576 

Triehechidce,  596 

Viverro,  526 

Tayra,  579 

Trichechus,  597 

Viverricula,  527 

Teetee,  713 

Trichosurus,  150 

Viverridce,  525 

Teeth,  13 

Trichys,  487 

Vole,  465 

Tegument,  7 

TncZw,  162 

Vulpes,  552 

Teledu,  575 

Triconodon,  113 

Vulpine  Phalanger,  150 

Telemetacarpalia,  323 

Trilodon,  484 

Temnocyon,  555 

Trituberculism,  30 

WALLABY,  169 

Tenrec,  637 

Tritylodon,  111 

Walrus,  597 

Terphone,  338 

Trochictis,  570 

Wapiti,  322 

Tertiary  inarnmals,  115- 

Troglodytes,  736 

Wart-Hog,  288 

Tetraceros,  338 

Trogontherium,  458 

Weasel,  589 

Tetraconodon,  292 

Trygenycteris,  655 

Whale,  225 

Tetracus,  634 

Tubulidentata,  179 

White  Whale,  262 

Tetrastylus,  488 

Tupaia,  617 

Wolf;  548 

Tlierulomyidae,  484 

TupaiidcB,  617 

Woivereue,  591 

Theridomys,  484 

Tursiops,  271 

Wombat,  145 

Theropithecus,  722 

Tylppoda,  295 

Thigh,  51 

Tylostoma,  674 

Xantharpyia,  652 

Thomomys,  478 

TypMomys,  477 

Xenurus,  198 

Thoracophorus,  203 

Typotherium,  440 

Xeromys,  461 

Thylacine,  137 

JTerMS,  452 

Thylacinus,  136 

Uacaria,  712 

Xiphodon,  294 

Thylacoleo,  157 

Uakari,  712 

Thymus  gland,  66 

Uintatheriidce,  437 

YAK-    ^fid 

Thyroid  body,  60 
Thyroptera,  665 

Uintatherium,  436 
Umbilical  vesicle,  77 

1  AK,   OO^r 

Yapock,  134 

Yolk'Sftc   7 

Tiger,  511 

Unau,  183 

Tillodontia,  441 

Ungulata,  273 

Tillotfterium,  441 

Urinary  organs,  69 

^opws,  480 

Tinoceras,  437 

Urocyon,  553 

Zebra,  385 

Titanomys,  492 

Uromys,  476 

Zeuglodon,  246 

Titanotheriidce,  413 

UropsUus,  629 

Zeuglodontidce,  246 

Titanotherium,  413 

Urotrichus,  629 

Ziphiince,  251 

Tolypeutes,  199 

Ursida?,  557 

Ziphius,  254 

Tomitherium,  698 

Ursus,  557 

Zoological  regions,  96 

THE   END 


Printed  by  R.  &  R.  CLARK,  Edinburgh 


QL  701  .P64  SMI 

FLOWER,  WILLIAM  HENRY, 
1831-1899. 

AN  INTRODUCTION  TO  THE 
STUDY  OF  MAMMALS, 

AKH-2714  (AB)