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^ U - 

6 - 







M.D., D.Sc., F.G.S., ETC, 




B.A., F.G.S., ETC. 



VOL. n. 




TO Bt V.tPT. 

i'ti;:: i-ii£.V/ \(Jli\\. 



n 1911 L 






General characters of the Vertebrata — Division of the Vertebrata 
jniocUsses — Exoskeleion — Teeth — Endoskeleton^\'cncbrse — 
— Skull — Limbs — Pectoral and pelvic girdles— Use of generic 
terms, 889-910 


General characters of Pisces — Division into orders — Scales^Fins 
^ Teeth — Vertebra ^Tail — Skull — Limbs ^Distribution in 
time — Mutual relations of the orders — Conodonts, . ,91 1 -922 


The Cyclosiomi^Characters of the Elasmobranchei — Characters of 
the Ichthyotomi — Families of the Ichthyotomi — Characters of 
the Selachii — Tectospondyli and Asterospondyli — Families of the 
Selachii — Ichthyodonilites, 923-948 


Characters of the Chimeroidei — Families of the Chimeroidei — 

Characters of [he Dipnoi — Families of the Dipnoi, , , 949-957 



Characters of the Ganoidei — Suborders and families of the 
Ganoidei, 958-99' 


Characters of the Teleostei — Suborders and families of the Tele- 
ostei — Literature of Pisces 992-1017 


General characters of the Amphibia — Orders of the Amphibia — 
Characters and families of the Labyrinthodontia — The Apoda — 
Characters and families of the Caudata — Characters and fam- 
ilies of the Ecaudata — Literature of Amphibia,. , 1018-1045 


General structure of the Reptilia — Sauropsida — Classification of 
Reptiles — Geological distribution of Reptilia, . . . 1046-1052 


The Theromorphous branch of Reptiles — Characters of the Anomo- 
dontia — Relations, suborders, and families of the Anomodontia 
— The Placodontia — The Synaptosaurian branch of Reptiles and 
its characters — Relations of the Sauropterygia and Chelonia — 
Characters of the Sauropterygia— Families of the Sauroptery- 
gia— Characters of the Chelonia — Characters and families of 
the Athecate C he! onians— Characters and families of the Testu- 
dinate Chelonians 1053-1118 


The Streptostylic branch of Reptiles and its characters— Charac- 
ters of the Ichthyopterygia — Genera of the Ichthyopterygia— 
Characters of the Proterosauria — Characters of the Rhyncho- 
cephalia — Suborders and families of the Rhynchocephalia — 
Characters of the Squamata — Suborders Lacertilia — Rhipto- 
glossa — Dolichosauria — Pythonomorpha — Ophidia— Famihes 
of the Squamata 1119-1149 



Tbe ArchosauHan branch of Reptiles and its characters — Characters 
of the Dinosauria — Suborders and families of the Dinosauria — 
Characters of the Crocodilia — Suborders and families of the 
Crocodilian Characters of the Omithosauria — Suborders and 
families of the Omithosauria — Literature of Reptilia, . 1150-1207 


Characters of Aves — Distribution of Aves in time — Relations of the 
Ratita^ to the Carinatx, iJo8-i22o 


Characters of the order Saurur^e — Characters of the Ratita: — Char- 
acters of the CarinatEe — Suborders and families of the Carinatee 
— Literature of Aves, 1221-1244 


General structure of the Mammalia — Teeth of Mammalia — Skull — 
Vertebral column— Sternum— Pectoral liml) — Pelvic limb— Dis- 
tribution in time of Mammals — Mutual relations of the different 
orders — Fhylogeny of Mammals, 1245-1262 


Characters of the Prototheria — Characters of the Monotremata — 
Families of the Monotremata — Characters and affinities of the 
Multituberculata— Families of the Multiluberculata ~ Charac- 
ters of the M eta theria— Characters of the Marsupialia- Sub- 
orders and families of the Marsupialia, .... 1263-1288 


Characicrs of the Eutheri a— Characters of the Edentata— Families 
of the Edentata— Characters of the Cetacea — Suborders and 
families of the Cetacea— Characters of the Sirenia— Families of 
the Sirenia, 1289-131I 



Characters of the Ungulata — Characters of the Artiodactyla — Fam- 
ilies of the Artiodactyla — Characters of the Perissodactyla— 
Families of the Perissodactyla — Characters and famihes of the 
Toxodontia — Characters and families of the Condyiarthra — 
Characters and families of the Hyracoidea — Characters and 
families of the Amblypoda — Characters and families of the 
Proboscidea, 1312-1410 


Characters of the Rodentia — Suborders and families of the Roden- 
tia — Characters of the Camivora — Characters and families of 
the Pinnipedia — The Camivora Vera— Teeth of Camivora Vera 
— Families of Camivora Vera — Characters and families of the 
Creodonta, 1411-1453 


Characters of the Insectivora — Suborders and families of the Insect- 
ivora — Characters of the Chtroptera — Suborders and families of 
the Chiroptera — Characters of the Primates — Suborders and 
families of the Primates — Literature of Mammalia, . 1454-1474 



Classification of the Vegetable Kingdom — General succession of 
plants in time — Laurentian Graphite — Supposed Cambrian 
plants — The "Fucoids" of the older Palseoioic fomiations — 
Silurian and Devonian plants— Nematophyton (Prototaxites) — 
Sporangites — The Carboniferous and Permian plants — Struc- 
ture of coal — The plants of the Mesozoic period — Tertiary 
plants, 1477-1487 


Characters of Thallophytes — Characters and divisions of Algae — 
Diatomacese — Phycochromophycic — Fucaceae — Chlorosporese 


— Chaiacteis and fewii fomu of the Siphoneas Vertidllata: — 
Fknidese — Condlines and Nulliporcs — Coccoliths and Cocxo- 
tjdieres — Chaiacoe — Fossil Fungi and Lichens, . 1488-1499 


ClaiBCten of Connophytes — Biyophjrta — Mosses — liverworts — 
Chatactera of Pteridoi^jrtes — Characters o€ Filicacese — Ophio- 
g^ossaceie and Haiattiaceie — Fens (F>li<»s) — Rfaizocaips — 
Equisetaceae — Lycopodiaceae, ' . 1500-1513 


Chaiacteis of Pbanen^ams — Characters of Gymnospenns — Cycad- 
accx — Noeggeiathie« and Cordaiteae — Chaiacters and fossil 
fimns of Conifers^ I534-IS37 


Chuacten of Angiospenns — Chaiacteis and fossil fisims of Mono- 
cotykdoos — Ch^acters of Dicotyledons — Principal groups of 
tiie Dicotyledons and their distribution in time — Literature <d 
Palaeobotany, 1538-1560 

Appendix, 1561-1570 

Index to General Introduction and Invertebrata, . 1571-1592 

Index to Vertebrata, 1593-1616 

Index to Paleobotany, 1617-1624 


Page 901, line 13 from bottom, /or "Weinsheimer" read " Wiedi 

II 912, line 2 from tap, for "Teleotomi" rtf(w/"TeIeostomi." 
It 928, line 1 7 from top, for " HyPocladodus " read " Hybocladod. 
fi 928, line 8 from bottom,/ur " Scilliida " read " Scylliida." 
ri 944. Carcharopsis is identical with Dicrenodus (p. 928). 
II 94S, line 6 from bottom, before " no basal " add " usually." 
11 951. Myriaeatttkus '^zs. recently been made the type of a disli 

family — the Myriacanthidce (see Appendix). 
,1 955, Ceratodus has recently been recorded from the Stormb 

beds of the Karoo system of S. Africa. 
II 965. Pnigeacanthus is identified with Oracanthus (p. 947). 
II 966, line 1 1 from top, before " very minute " add " usually." 
II 966, line 5 from bottom, de/e " minute." 
II 967, line 4 from lop, for " Carboniferous " read " Devonian." 
II 969, line 9 from top, de/e " and Carboniferous," 
II 971, fig. gos, for " Heberti" read " HibbertV 
II 978. Palaoniscus occurs in the Stomibei^ beds of the Karoo i 

tem of S. Africa ; and from the underlying Beaufort beds a 

allied to Rhabdolepis has lately been described as Atherslon. 
II 979. The name " Platysomida " should be amended to " P/atj 

II 988, line 22 from bottom, dele "as in Belonorkynckus?' '. 

genus Belonorkynchus has recently been described from 

Australian Hawkesbury beds ; while Belonoslomus, or a do; 

allied form, is recorded from the Lameta Cretaceous of Indi 
II 989. Hypsicoriniis has been recently described from the Oxf 

Clay, and Eurycortmts from the Kimeridge Clay of England 
II 1004, line 10 from bottom, for " Saurinichthys " read " Taurin. 

H 1033. A species oi Bothriceps has been recently recorded from 

Karoo system of S. Africa. 



h|t 1035. The [ume Piaiyops bein>; preoccupied, may be changed to 
Piat/opouatntt. The genus should apparently l>e included in 
the Aniugauuiwidit, since the vcrtcbr* ate de»tril)cd as rbach- 

■> ioj6l C^iiotattfus also occurs in the Bunter. Th« superiority of 
lite oif CopitQtavrus over Triwatfsaurus renders it more prob- 
able that the former is the taine aa CAirtuaurut (p. 1039]. 

■• 1036. 1037. The alleged occurrence of Afast^doittaiirut ^ganttui 
and Atttopias in the Rhxlk rests on tn»ufficicnt evidence. The 
Utter genus is stated to have been recorded from the Conli- 
Kdta] inttead of the BntUh Kha;tic 

" KHi , boium line, li^le " and may not improbably be identical with 

■■ 104J, line 8 from top,>r "each" rvad "hind," and f^ "pes" 
rtad "same." AlyUt occurs in the Miocene of ^iau&an i,Ritaa 
TrouktU). Latoiaa (p. 1044), according to Cope, is not allied 
ui Ctratepkryi, bui beloa>^ to the DucogitmiMr ; it also 
occurs in the Sausan Miocene. The Palaobalrachida appear 
10 connect the Ptlohahdx u^th the AglosKn, ah represented by 
d>e DactyUthritla; or Xencpodida as they should properly be 
called. — DMlyltihr-M lieing a synonym of Xtn-tpas. 
1044. The name CyitignatAida should be replaced by Lcptodatiy- 

Ndf, Cystigiuilhus being a s)-nonym. 
loj?, line 8 from top, A^<wv " characterised " aJJ " lypicatly." 
1057, line 17 from top, bt/are " African " add " typical." 
(0^8-41. Additional characters of many of the genera are given in 

tbe Af^ndix. 
(059. line 4 from \t)p,/cr " divided " read " undivided." 
lo>9t line 7 from top,/or " single " rtad " double.'* 
1108. Oi-adia is distinct from Pal^ockelyj. See Appendix. 
1117. line 10 from bottom, the name " CycleitarbiHa " may be sub* 

siiiuied for " Emydina," since the latter is preoccupied. 
IIS& Since hhl/tyosaurus p/al/edoH differs fnKn all other species 
of hfitkyoiaunts by ttie smooth and cuinaled crowns of its 
teeth {^g. 1038), it appears, on the whole, advisable to regard it 
as the type of a distina genus, for which the name TemiKdantO' 
uuirut in proposed. 
tiyf. Rht^iotaurut prot-es to be founded upon part of a jaw of 

Paekyrhtjodus (p. Q93;. 
1176, line J from top,/*r" Kimeridgian " rtmi" Ponlandian." 
1179, line 18 from top, i»//rr " species " aiA/"of ilic former." 
1164. fig. 1 130, A is a skull of ifam's, and not £,/sida<t, as stated. 
157:, lines 16, 1 ;. from lfift,/^r"/.t»tHfi/Afrium" rfnd '^ JJmnohyut:' 








General Characters. 

The Vertebrata, or highest types of the entire animal kingdom, are 
distinguished as a whole from all the preceding sub-kingdoms (col- 
lectively designated as the Invertebrata) by the genera! presence of 
an internal skeleton, and more especially of a cylindrical longitudinal 
axis, termed the notochord, which is usually replaced in the adult 
bv a series of cartilaginous or bony segments collectively constitut- 
ing the vtrlebral column. This axis, or column, separates the smaller 
dorsal or neural tube of the body from the larger ventral or visceral 
'h-cmal) tube ; and the body itself, together with its appendages, is 
always symmetrical to this axis, and is never externally divided into 
>egments. Limbs may be completely wanting, but when present 
they never exceed two pairs, and are always turned away from the 
dorial or neural aspect of the body. 

That the Vertebrata have been derived from the Invertebrata at 
an extremely early epoch of the earth's history is practically certain ; 
and although we are unable to point to the direct ancestors of the 
^ub-kingdom, yet we have an inkling of this relationship exhibited 
by the presence of a notochordal structure in the earlier stages of 
the Ascidians, while there are also certain features in the organisa- 
tion of the Annelids suggestive of their being allied to the primitive 
stock whence the Vertebrates took their origin. Since, however, it 
is probable that these primitive types were soft animals, it is unlikely 
that any light will be thrown on the origin of Vertebrates by means 
of Palaeontology ; and if the problem is ever to be solved it will be 
by the aid of Embryology. 


Leaving then the origin of Vertebrates as an unsolved problem, 
in the solution of which the palaeontologist can have but little share, 
we may proceed to a brief survey of the classification and chief 
structural features of the Vertebrate sub-kingdom, so far as they con- 
cern the palseontologist. 

For this purpose the sub - kingdom may be divided into five 
classes' — viz., Pisces or Fishes, Amphibia or Amphibians, Reptilia 
or Reptiles, Aves or Birds, and Mammalia or Mammals— of which 
some of the more important features, from the point of view of the 
palaeontol(^st, will be mentioned under these respective heads. 

The first and second classes have been brigaded together by Pro- 
fessor Huxley under the name of Ichthyopsida, and the third and 
fourth as Sauropsida ; and these terms will frequently be found 
convenient. Other writers, again, from the absence or presence of 
certain structures during the course of development, group together 
the two first classes as Anamniota (Anallantoidea, or Branchiata), 
and the remaining three as Amniota (Allantoidea, or Abranchiata). 

Since the hard parts of Vertebrates are those with which alone 
the student of Palsontology usually has to deal, it will generally be 
unnecessary in this work to make any allusion to the soft parts of 
the body. With, regard, however, to these hard portions, it is ad- 
visable to give an extremely brief sketch of the more important 
elements of the Vertebrate endo- and exoskeUion for the benefit of 
those readers who are unacquainted with the elements of Compara- 
tive Osteology. It must, however, be distinctly understood by all 
who desire to practically study the history and structure of extinct 
Vertebrates, that it is absolutely essential they should have that 
thorough knowledge of the osteology of the recent members of the 
sub-kingdom which can only be gained by familiarity with actual 
specimens, accompanied by patient and laborious study of the 
numerous works on the subject. The following sketch is, indeed, 
merely sufficient to enable the reader to understand the meaning of 
the terms employed in the sequel ; and throughout the Vertebrata 
the limits of this work will necessarily permit of only some of the 
more salient features of the skeleton of the various groups being 
mentioned. It may also be observed that those groups which have 
wholly disappeared, or of which the palaeontological and evolutionary 
history is of especial interest, are more fully treated of than those 
more or less exclusively confined to the later or present epochs, and 
of which the history is fully recorded in those works to which the 
term "Natural History " is usually restricted. In the majority of 
instances, again, space does not permit of allusion to species; but 

' A sixth clasj — Leplocatdii — is formed for the receplion of (be Lancelet 
{Amphiexus), with which the palKontologUt has, unfortunately, no concern, as 
its post history is a blank. 



genera, where the species differ conwdtr- 
ters, attention is in some inKtaiice.s tltrectcil 
to Ok mere imponani specific t>'pcs. The student must not, mor«- 
flfw, expect to find that every known genus of fossil Vcrtebratci, or 
ecm entry family, is mentioned in the following chapters, of which 
tbeob^t ii to enable htm to t^in a fair general knowledge of the 
paa hitcvy, dimribution, and lending ^cruciural fcatuie^ of the bettt 
buwn groups of the various classes. 

Conmeneing with the outer skeleton or exosMrlon it inay be 
cbseiTed that, a» a rule, the palaeontologist has but little to do with 
sonctuiti developed in the tpidtrmis, or layer overlying the true 
atin n» dermis, ancc these generally perish during the process of 
peOifectton. The scales of Lizards belong, however, to this la)xr, 
and ihcii impressions are in some instances preserved : while in the 
Cbtlonu ihe t>oundancs of the horny shields, covering the bony ^hcll, 
are in tome instances the most imporfint characten. by which fossil 
ipecintm can be determined. Occasionally, again, as in the Solen- 
bcfai Ankitepuryx, the feathers of Birds have left their impret;.^ion 
10 rockj of which the material is of a fine-grained slruciure ; while 
Jtdl more rarely, as in the bony covering of some Glyplodonts, pits 
6m which hairs or bristles once grew arc prcsctved in a fossil 

Tbf (krmai cxoskektal suucturea are of considerably more ini- 

pORance fmm a palseontological point of view ; and it should be 

obnved that in nearly all the chases there appears to have been a 

fadual tcndejicy to the disappearance of the bony elements of this 

ikeleton in the higher rorms, this being especially marked in the 

otte of Fishes, In tlie latter class the scales of ;ill types, of which 

ttler menijon will bo made in the sequel, belong to the derniiR, 

and in some forms there may be a complete dermal armour formed 

of imbricating scutes, as in CaUiihtkp among the Siluroids, or, as in 

the Coffcr-dshes (Ostrac/on), consisting of calcified scutes with their 

edges in apposition. Apparently the most primitive type of tlennal 

arroouT in this class consists of small dcntirulcs supported on l)ony 

pbtes o'ld it has Ijeen found that sudi denlicules are absolutely 

hooxdogous with true teeth, of which we !>hall speak presently. 

The bony^»«iw' andjf«-*//w« of Fishes are another developmenl 

of the dermal skeleton, which will be noticed under the head of 

that class ; althoogh it may be obser\ed here that many of the 

latter arc closely allied in structure to teeth. Again, the dermal 

skeleton in many extinct .\mphibians and Reptiles takes the form 

of a more or less complete armour, cither on the ventral or the 

dorsil, and not unfrequenily on 1>oth aspects of the body, con- 

' TImm tlcrml fio-nys mnsi not U conrouinlcd with the rullal cartlln^ of 



stnicted of solid bony scutes, which may imbricate or overlap one 
another, like the tiles on a roof, or may be firmly united at their 
edges by sutural union. Such an armour may also develop enor- 
mous bony spines, often attached to the skin by an expanded base 
which represents the scutes. In the latter class, again, the bony 
shells or the Testudinate Chelonia are formed partly of dermal 
elements, blended with others belonging to the endoskeleton to 
form a continuous whole ; while in the Athecate division of the 
same order the whole of the protective armour is of dermal origin. 

The so-called membrane bones of the skull, and the clavicular 
portions of the pectoral girdle, are likewise of dermal origin, and 
therefore properly belong to the exoskeleton; but their intimate 
connection with cartilage bones renders it more convenient to con- 
sider them with the endoskeleton. 

Teeth, as belonging to the list of dermal structures, may be con- 
veniently noticed here ; and the importance of these organs to the 
palaeontologist can scarcely be overrated, since from their extreme 
density they are more often preserved in a perfect condition than 
most other parts of the skeleton, and thus frequently form the only 
safe guide to the affinities of an extinct type. Teeth are composed 
of two or more earthy constituents, varying in their structure, and 
the amount of animal matter contained in them. The 
most important element is that known as dentine (fig. 
8 1 3, d), which forms the greater portion of the body 
of most teeth, and consists structurally of extremely 
minute tubes, cells, and earthy particles. Some den- 
tine is devoid of blood-vessels ; but in other cases it 
is permeated by the latter, when it is known as vascu- 
lar, or vaso-dentim ; and both these structures may exist 
in a single tooth. In young teeth (fig. 813) the centre 
of the dentine mass is occupied by the vascular /»^ 
eavity (pc), which is open at the base ; but in the adult 
this cavity is often totally obliterated, by the formation 
of what is known as osteo-dentine, which is a kind of 
vascular dentine passing imperceptibly into the structure 
of true bone. The second primary constituent is the 
enamel (f), which, when present, immediately overlies 
the dentine of the crown, or exposed portion of the 
tooth, and is the hardest knoivn animal substance ; 
it is composed of extremely minute prismatic fibres, 
generally running at right angles to the outer surface 
of the tooth. This substance is the least constant 
element in teeth, although it is very generally present 
in those of Mammals ; among the Reptiles it is entirely wanting in 
the Ophidian Squamata, but is present in the Crocodilia. The 

FiE. 813.— Dia- 
gramma-tic verti- 
ciLl section of a 
tooili. /, Cemeni; 
rf. Dentine ; c. 
Enamel: /(.Pulp. 



eoDMituem. usually known as ttmrnt (fig. S 1 3. t). I)U1 occasion- 
tiAPWAi fftroia, is the most external of Ihc three: in same 
iniaiiccs, u in fig. 813, it is con6ned to the root, or embedded 
pOftiw of the tooth, where it consequently comes into immediate 
onaa with the deiitinc; Init in other cases it is Tound overlying 
tiKcaund of the crown, and in others as^in, where the enamel is 
vantii]^ it forms the main covering of the crown, as in the teeth 
of liie Edenute Mammals. The cement is always traversed liy 
Tucabr canals ; anil its structure ta very similar 10 that of bone, 
alAoa^ only occasiotially presenting all the peculinrities of the 

TnK teeth are generally confined to the mouth and pharj-nx, but 
Acy Mty be situated on many of the bones of the former, and their 
node of attachment varies from a simple anchylosis to the under- 
lfiii|bone, to implantation in distinct sockets. The simplest romis 
of tetlfa are small granular bodies, like the minute ossifications 
ilnaily mentioned as occurring in the skin of certain Fishes ; while 
iboKof the most complex xtructure are 10 be found in certain Fishes 
\Ikiiirodta\ in the I^byiinthodont Amphil>iaii», and among Mam- 
miit in the Edentate genus Orydrropus and many RodenU and 
L'ngiiUtes. Except in forms whcr«; all are alike, as a general 
rale the teeth at the anterior extremity of the jaws are more or 
la* nmple, while there is a gradual increase in their complexity 
towds-the opposite end, and in most cases the lower teeth arc 
funower and more elongated in an antcio-posterior direction than 
ihc upper ones. 

The importance of teeth to the palaeontologist as a means of 

deiennining the affinities of fossil fDrm.s ha.<t been already mentioned ; 

tm in many cases their evidence must be supplemented either by 

that of other remaim, or of the geological horiran whence they were 

obtained ; sir>cc otherwise serious error may result. Thua, the teeth 

of the Dinosaorian Me^n/osaiirus present a strong rc^cmbtancc to 

lonie of those of the Mammalian Mafhan^dm; while the front teeth 

of 4ome ^MToid Fish approximate to those of the Primates ; and the 

lower hittder teeth of the Krmgaroos, of llie Dinothere among the 

I Proboscidea, and of the Tapir-like animals in the PerissodactyU, arc 

&U singularly alike in form. Another fertile source of error to l)C 

' yarded against is the great difference in the form of the teeth from 

■different regions of the mouth. 

^H^Fur the mieroseopic structure of teeth and their mode of develop- 
^^Lnt, the student must refer to other works : but a large number 
P^T ihe more important types of denial structure will be found noticed 
in the coarse of the following pages. 

By far the greater number of adult X'eitebrates possess a solid 
inner, or tndciktltlon, composed of hone (into the nature of which 



t -W -0.1, 

it win be urmectfssary to cnler here) ; but in otliLTS this skdeto 
remains cnnihi^inouii throughout life, or, as in many Sharks, h£ 

solid striiclures formed by cakificntior 
in the cartihi^e, which are quite distini 
from true bone. This endosktieton ma 
be divided into on axiai pcrtion, or tlu 
or the head, tnink, and uil ; and an e^ 
pindicuiar pdrlhH supporting the Hmb* 
the relations of the two being shown i 
fig, 814. The anterior part of the axil 
skclLion is formed by the skuU (of whic 
we shall speak later) ; and this is sui 
cccdcd posteriori y by the vertebral coiumt 
typically eomposed of a niiniher of se] 
nR'tU:^, known as vertd'nr, placed in th 
rniddtf line of the dorsal side of th 
body, and continuing from the head t 
the fxtreniity of 
the tail. 'I'hi.s 
column is devel- 
oped round a rod- 
like axis known 
as the notoc/wrd. 
\n eenain of the 
lower forms the 
latter may persist 
throughout life, 
and may either /^^ t^^^}^ ^^^^^ 

simply have a ■A™;,r"''.7"'K'^5"^pS; 
fibrous or carti- (brok«i.): /«. Pm>-(Apcvh: 

laginous sheath, ;/.i-™;^^J^<bSSG! 
or ma) tiavt. cal- jf^-. Mu.aiu.tWB«. 
cifications, or car- 
tilages, in either the dorsal or ventral poi 
tion ; and thtrrc is a complete transitio 
from such a primitive type of colum 
to that of the more specialised types i 
which the vertebrae arc fully ossificc 
'Ihere is an enormous amount of variatio 
in the structure of the vertebrae of diflci 
ent groups, and even in the diflreren 
regions of the lx>dy of a single aninia 
but there is one general plan pervading them all. Thus a tyfHCi 
vertebra (fig. 815) eonsisla uf a baaal portion, or anirum, whici 
may be either disk-like or more or less elongated. Its posterio 

Labnbthmloiii AmtiKHiUn (.57/- 

mofid from lti« left tide in onJcr 
10 khow <ht •iHfoilteI'loa- <>ivatly 
nUonicd' (After FrilKb.) 


udnttrior mriaces, by which (with the intervention of cartQages) 

3 vdculatcs with the adjacrnl centra, may be nearly flat or con- 

a<c«hcn it is said lo be amphiialom (fig. 815); or its anten'or 

«bce may he more or less holtow, and the posterior convex (fig. 

Sii), vhen it is tertned/nxvAwi .- or, lastly, the reverse o\ the Xal- 

laamngODePt may obtain, when the centrum is tcimcd apistkoari- 

m; in example of the latter Mructure lieing shown in the vertebra 

at Cil»imtpoiufy/0t given to the sequel (fig. 1071^ Imiucdiately 

ipv the centrum is the 

apatitrc of the channel for 

ifae tccrption of the ftpinal 

<ai, liDown OS the Htnral 

audifxg. St 6); this canal 

bein( enclosed tatcrnlly aitd 

■paiorif by the neural 

cvla, which are surmounied 

\i\^ wnral tpim {^^ 815, 

%\tt\ 'I1ie lateral poftioru 

a/ du» arrh are termed the 

/*'^(^- 817, n\ and the 

!■* cocnccting the laiter with the spine the iamina. The arch, it 

"kttiA be iA)«a%'ed, always ossifies separately from tlie centrum, 

ted tlie hne of junction between the two when, as in many Rep- 

Fig. t>6.— AniBW (n) and jnitcrie* <c) vliwi af t. 
trDOfkiJi donal vcrlcbr* nrf ■ l-arcrtian itcinilc 
(l''.dna«»jV Ttlt jvrl klnx* !■ %t-A c (« ihacmlrvra; 
(he anitura lh« iwuni »r>al,i>l>«Kaiiil njuiiil which 
vt* uic n«unl arrlitA, «unnDUnr*i1 br >h* ntuni 
fpiac; lb* oblique botn an iht nx'popliyMi. and 
IM laMtal franirani-** ibc inuivnrw tuecwi M . 
■ Utb «M \in «ry ■hull. 

fi^t\\.-~KX>^ ■•■■ fihtlonilttt»»fltbrao(« Whale, a. DuiKramnu ik ima - 

■M* anigB e* ih* l»a~ ol iIm ilisnxK twipea af ■ MannuL Gnolly fdaead. a, Pnnja- 
) »*] m : i, Smi— > : 1. Ccnuao ; rf, Traairenc pruceu; ■, Pedld* oTanb: /, SiMual nbi 
^^:j.NMnl>c«a. (AJUrOvw.) 

they remain distinct, is known as the «<w»-o-(V«/m/ suture. 
neural arch bean a pair of processes, or facets, at cither ex- 
aity for articulation with the adjacent vertebrae; those at the 
amerior extremity (fig. 816, b) being known as fnsygapophyses, and 


looking inwards and upwards ; while those at the other extremity 
(fig. 816, c) are termed posttygapophyses, and look downwards and 
outwards.^ Besides these processes for mutual articulation, there 
are other processes on many vertebrae ; the most constant being the 
transverse processes, or diapophyses, which may be very short as in 
fig. 816, or greatly elongated as in fig. 817, a, <f. These transverse 
processes may arise either from the arch or from the centrum, and 
in the trunk frequently serve for the main attachment of the rOa 
(fig. 817, b). In fig. 816 we have an example of a vertebra with 
very short transverse processes placed mainiy on the centrum, while 
in the vertebra of Iguanodon, figured in the sequel (fig. 1058), we 
see very large transverse processes arising solely from the arch. 

The sides of the centrum in the anterior region of the body 
frequently carry other articular processes for the ribs, which may 
be termed rib or costal facets (fig. 815, cp), or parapophyses. 
Again, the inferior or haemal surface of a vertebra may bear a 
hcemal spine — the term haemal being applied to the inferior as- 
pect on account of its being directed towards the heart and large 
blood-vessels. Each vertebral centrum carries its own arch ; but 
between the true centra of the Ichthyopsida there may be inter- 
calated centra-like bodies, carrying no arches, and termed intercentra. 
Rudiments of such intercentra occur in the so-called wedge^nes or 
hamapophyses, found between the lower borders of the centra in 
certain Reptiles, and more especially between the first and second 
vertebras. Further, the Y-shaped chevron-bones, usually articulating 
between adjacent vertebral centra in the tail of many Vertebrates, 
are also morphologically intercentral elements. In Fishes the ver- 
tebral column can only be divided into trunk and caudal regions ; 
but in the majority of higher forms further divisions can be made. 
Thus the vertebrae of the neck, in which the ribs never articulate 
with the sternum, are termed cervical (fig. 818, c); the first of this 
series being designated the atlas, and the second the axis. It is 
further remarkable that in most instances the centrum of the atlas 
is not joined to its arch, but either remains as a distinct element, or 
becomes anchylosed to the front of the centrum of the axis to form 
the so-called odontoid process ; and it appears that in the latter case 
the inferior bar connecting the two lateral arches of the atlas is 
really the remnant of the first intercentrum. It is, moreover, prob- 
able that this separation of the component elements of the atlas 
should be regarded as a retention of the primitive feature obtaining 
in the vertebral column of many extinct Fishes and Amphibians, 
where, as will be noticed below, the whole of the vertebrae are thus 
disintegrated. Finally, it has been suggested that certain bony 

' These ate well seen in the vertebra of Calamospondylus above mentioaed. 



centra ; but by the disappearance of the latter they have frequent]] 
become transferred to the centra or their appendages, although ir 
other cases their original derivation is indicated by their articulatior 
at the junction of two vertebrae. Very generally ribs articulate witl 
the vertebral column by two distinct heads, which is probably th< 
original primitive type of structure. In these cases the anterior a 
lower head is termed the capitular, and articulates with the rib 
facet or parapophysis on the vertebral centrum (fig. 815, (p); whil) 
the posterior or upper head is known as the tubercular, and in thi 
dorsal vertebrae articulates with the transverse process or diapophysi: 

of the arch. In the dorsal ribs tht 
two heads may, however, coalesce 
and articulate with a facet on tht 
centrum, or with a longer or shorte 
transverse process on the arch. Fre 
quently, moreover, while the anterioi 
dorsal ribs have double heads, tht 
posterior ones have but a single heai 
articulating with the transverse pre 
cess. In the caudal region of man 
Reptiles true ribs articulate with th 
upper part of the centrum, and o 
cept as being separate, are indi: 
tinguishable from the caudal tran; 
verse processes of many Mammal: 
such as the Cetacea (fig. 817' 
Cervical ribs are present in mos 
Reptiles, and usually articulate b 
two heads to the vertebrae — th 
upper head joining a facet, generall 
placed on the arch, correspondin 
to the transverse process of the dorsal vertebrae, and the lower on 
articulating to another facet on the centrum. Very rarely in Rei 
tiles these cervical ribs may be completely anchylosed to the vei 
tebne, as in the vertebra of Calamospondylus, figured in the sequt 
(fig. 1071) ; and this suggests that at least a portion of the so-calle 
transverse processes of the cervical vertebne of Mammals, whic 
arise from the vertebne by double pedicles, really correspond t 
cervical ribs. In the Sauropterygia both heads of the cervicj 
ribs articulate with the centrum. 

When a sternum is present the distal ends of the dorsal ribs ar 
generally unossified, and are sometimes termed intermediate ril 
{ir, fig. 819); and these unossified elements unite distally *-ith th 
sternal ribs {sr), which in their turn join the sternum (fig, 817, &,b 
Further, among the Sauropsida lateral ossifications may be develope 

Fif[. Sia.— Stcmal region of a young 
OmilkoriyncAtta. f. Clavicle; /, Inlcr- 
clavicle ; >, Presternum ; mt, Mcwsier' 
num; r, tub; i>, IntermedLate lib; tr, 
SwnuJ rib. (AClet Fiower.) 







t^f Vm>— L«li i*l«nl vnw oT ihc •kuB of a« 
cmtnio DotfAiJi. Ir, lAfi mriKuli ; «. *. ■. 

Na-uli oit'iul, alid uidili^iy canulo ; m, Ajr, tr, 
kUndifculiir. hroid, nnd htAnchial Knhc«: el. r, 
H>^n*fi>tibdUr AiiJ tint krAnchiat cUft*. (Aflcr 

I ibe rflH, termed una'nate p/xvesKs, of which Ihc ]}OBilion is shown 

I the fii«« of ihc skeleton of the Eagle given below (fig. 1 106). 

The brcul-ltone or s«mum 4fig». 817, 819) ts usually composed 

' aT 1 mtdian s«Hc-t of IwtMS or canil-nges on the i.-enttal aspect of the 

bodj, which w diviiiUc in the higher gioupa into an anterior pre- 

xHtaatm, usually consisting of 

1 «nglc ovsibcalion ; and of a, o a ff tr 

saa of mtsoiltmat eleroents, 
Mkwcd posteriorly by the 
KfMskniMm. In many &iiiir- 
opriik the SiUma) boacs have 
\im% hieial pfocesses ; and the 
oaifications in tliis c)as& may 
flDBSui of a pair of bones 
uttted hf cartUage. 

Ibe tkwtl, or anterior tcr- 
BJBitioti of the axial skeleton, 
nor diom our attention, hu: 
(he dcKription of this impor- 
ua and diflKult part must 
be of the briefest. The re- 

Kirtba of embryologists have shown that the skull is only a 

ifHCid modification of the primitive dements from which the rest 

rf ifce axial skeleton were fonncd. although it does not consist, as 

nj ooce thought, of a series of modi- 

W nmcbrx. The skull is divisible 

im a dorsal, or cranial, and a ven- 

tni Of viutral, portion ; the former, 

otfioating from a series of primitive 

tegmenU isomitet), encloses the braiii- 

nntr; while the latter, which has a 

Ksmentation of distinct and later 

is primiti**ely connected with 

function of respiration. 

The cartiest commencement of the 

j)rimiti\-c rjnilaK'"""*' cranium occurs 

in the foniiaiion of a pair of lod-like 

■cuia (hg. 820), lying at the hasc 

the brain, of which the posterior 

■hffrdal parts embrace the cx- 

ly of the noiocbord (fig. 821). 

parachordals soon unite to form 

a JtfjiVdr piatt supporting the brain 

(65. Sat); while the anterior froihordal parts unite in front to 
a space (ibid-, d) for the passage of (lie olfactory ner\'cs, 




Vif, Sii. — fpptr ""v (/ » Ial*r 
embryo, tl, CStnu or inlxcula ; f/ 
tnifl/, CKorhlid ■nd poniarbitti tm- 
ccMKi •■( d<h ; t, Eihmoxiual •cprnm ; 

Ofaclury rvnuiiex. Olhii Idlan »i ■■■ 


the united portion forming the ethmonasal septum {ibid., s). By th 
appronmation of the basilar plate to the nasai, orbital, and auditor 
capsules, these three distinct sense-regions become differentiated i 
the cranium ; and while the first and third become enclosed i 
cartilage, the lateral borders of the basilar plate, in some instance 
grow upwards to enclose the brain in a complete cartilaginot 
capsule, which in certain Sharks (fig. S22) persists throughout lift 
In the higher forms, however, the cartilage does not extend upward 
over the brain, which becomes roofed in by bone formed direct! 
from the overlying membrane. 

The visceral portion of the skull is formed by cartilages arrange 
in a series of arches in the walls of the throat (fig. 820). In th 
Ichthyopsida there may be as many as nine of these visceral arche: 
but in the other three classes they become reduced in number to 3 
most three or four, which are also functionally modified. The firs 
arch (fig. Sao, m) supports in all cases the walls of the mouth, an> 
is accordingly called the mandibular arch ; the second is termed th 
hyoid ; while the remaining ones, which persist only in the adults ( 
Fishes as supports of the gills or bronchia, are termed bronchia 
It should further be observed that these arches are separated froi 

^ «f-* pr^t^ j^j> 

WS, Vertebral columa. (After Wicdersheim.) 

one another in the embryo by a series of visceral clefts (fig. 820), c 
which the respiratory apertures of Sharks are remnants. The mar. 
dibular arch becomes divided into segments, consisting of a shoi 
proximal portion known as the quadrate, which very generally form 
the main support of the lower jaw ; and a long distal portion know- 
as MeckeFs cartilage, around which the mandible, or lower jaw, is sul 
sequently formed in those forms which develop membrane bones, bii 
which in the Sharks persists throughout life as the functional lower ja' 



Siol. The quadrate gives off an anlcrior pahtopurygoid {pala- 
^ItfMiimtt) bar. which in Sharks (fig. 813) persisU to form a kind 
of bbc op[«CT jaw. Tbe hyoid arch, which is close to ihe mandi- 
tmlar, and which in Fishes may also take a share in the support of 
tbe aandibte, ii likcvrisc scgmcmcd ; ihc most imponam eitmcnts 
is the liKer class being the kyomandibular and the rymp/uti/, which 
aitthowD in an ossified condilJon in I'lg. 823, km, sy. As already 
raoitioned, in Sharks th« primitive cartilaginous skull ia completv, 
«ad pcTsifts in this condition throughout lire ; but in the great 
raijomy of Vcrtebraies the progress of chondrificaiion is arrested, 


(W******'''^^ \, 


' ia 


ud the skalt becomes mon; or less coniplclclj' covered in with a 

«nB of ossifications dCTcloped in the membrane overlying the 

toain, while the cartilagjtious foundation itseif t.s likrwi^L- converted 

into bone. Bone!i derived from these two totally distinct sources 

nndgamate in Ihe aduli in such a manner as to afford no clue lo 

chetr dual originiL Following a modificntion of an arrangement 

idc^Ml by ProfesMir Wcinahcimcr, the more iin[)OTtinl bones of 

ihe cranium may lie enumeraied as follou's : the relative position of 

moSL of them beit^ shown in figs. 834 and 825.' The cartilage 

bocies comprise the haswaJn'laJ, dashp^rHot'if, and prtsphtnoid, 

which arc median ossificalions (not shown in the two figures) lying 

n this ordef (from the posterior extremity) on the inferior aspect 

of the craniunn. wid the first forming the floor of ihe /oramtn mag^ 

Hmm, or aperture by which the (.pinal cord enters the cranium, and 

the thrc* being collectively known a» the t^tuWnima/ axis. On 

> In ibe tcqiMl tit (igarct of the dnllt of many re])tilM. aiid more especially 
l)u»e o4 JieMj-MUtmi ntid Hlktiaurta, thnir ihc tjenenl lehltuii* oF (he Itcnn 
* ill 10 ai)t«nlS(e. 



the sides of the basioccipital we have the two exocHpUah^ forming 
the lateral boundaries of the foramen magnum, and either alone or 
in conjunction with the basioccipital, the single or double occifitai 
condyles, by which in the higher forms the cranium articulates with 
the atlas vertebra. The bony auditory capsule is composed tyjM- 
cally of the prooHe, epioHe, and opisthotic^ to which in Teleostean 
Fishes must be added the sphenoHc and pteroHc ; but, as will be 
noticed below, some of these bones may unite, when they receive a 
distinct name. The altsphenoids and orbitosphenoids are paired 
bones developed in the trabecular region ; while the single etkmmd 
and the paired turbinali occur in the nasal region. The position 
of the quadrate has been already mentioned. Among investing or 

Fig. 824. — Upper ''urface of the cranium of a Labyrinihodoni Amphibian {Kyrtvtia^ one-hftlf 
natural tiie. im, Prcmaiilla: m, Maiilla: N, Nual ; L, Lachrymal ;/, Frontal; P/, Prefrontal i 
f, Jugal ; Qy, Quadratojugal 1 c>, Opi«ihoiic ; OS, Supraoccipital ; Si/, Squamosal; Squ, 
SupratempooJ ; Pa, Parielal; Pff. P«tfronnJ ; Pto, PosiofWlal. The ijuadrate would come 
below Q7 \ Ihe large vacuities arc the orbiu; and the imall aperture in Pa the parietal fora- 
men. (After FriiKh.) 

membrane bones, which are of a more or less splint-like structure, 
we have the following paired ossifications, reckoning from before 
backwards, on the upper surface — viz., premaxilla (im), maxilla (m), 
nasal (//), lachrymal (Z), frontal (/), prefrontal {Pf), postfrontal 
{Ptf), postorbital {Pto), parietal (Pa), supratemporal {Squ), and 
squamosal {Sq). In some Dinosaurian Reptiles, as Stegosaurus, 
there appears to be a distinct bone above the orbit, which connects 
the pre- and postfrontal, and may be termed the supraorbital. The 

' Shown in thi: slniU of Ttstudo, (igui^ in the sequel, fig. 1017A. 



[jS»M M |< H i i/ (OS), vrKich, although double in the figured skull, is 
tmBj 2 single Iwne, appears to be developed infcnoily from ou- 
tSigc and superiorly from membrane ; it usually forms the upper 
bcnla of the foramen magnum, and is not unfrtiquenily produced 
poMnoriy iixo a long spine. On ihe Ulcra! aspect of llie traniura 
m fiscal Oiti Juga/ (J), and ijuaJnUoju^al {QJ), which connect 
tk qtudrste with the maxilla; while infcriorly {fig. 825) we may 
bnca median splint-like punxsphtnoid (i'i), and always n single or 

pumJ ro«wr ( V\ and the paired pter>'goids {Pi), and palatines (/*). 
TTw two bner, it may Xve observed, are developed u|>on die priini- 
Int palatoptcrygoid hor; while the parasphenviil, when present, 
uxlnlies the basicranial axis, and if largely dcvduped, as in Tclcos- 
teaa fishes and iVmphibiA (tig. 835), seems to take the place of 
the basi- attd prcqshenoid. 

In the cnuiium of which an upper x-iew is given in fig. S34 the 
iritole of the region behind the orbits is txjmpletcly rooftd over by 
hone, so that a secondary roof is thus formed aljove iht- roof of the 
nio^ smaller brain-case which lies within. In most Reptiles there 
aie, however, vacuities or fojste In this outer roof (as in fig. 836), 
aough in the Turtles and tlie Ichthyosauri (fig. 1024) this roof 
sists. In fig. 8)6 the upper-lateral vacuity it termed the sufira- 
{fosM, and is bounded below hy the saperwr ftmfvrnl (or 
VOL. 11. H 



F^ttamffsa-prefmntai) aratde rormed by the squamosal, postorbital, 
and postrrontal ; while the lower or infratemporal fossa is bounded 
superiorly by tlie last-named arcade, and below by an inferior tat- 
pom/ (oz quadrato-maxiliary) arcade^ fonntd in most Reptiles (fig- 
Sift) by the quadrate, quadratojugal, jugal, atid mjixilia. In the 
Maminalia (where it is usually termed the ayj^matic arch) ■vrc find, 
however, a single arcade foimed by the liciuaTnaKal, jugai, and max- 
ilia, and a similar arcade, hut with the apparent absence of the 
Jugal, occurs in many of the Anomodont Reptiles. This may be 
termed ihe squamoso-maxitlary arcade. In many Sauropsida, when 
a pustorbila! or poslfrontal is developed, these two arcades arc 

PIe- t(4. — (tight Iflitnl mpeci of iht ciuiium of $fkiti»it^ f^minita. 
on i£e lofi h the infnWrapcVkl (oub, baundnl t-elow hy the iivrerior. niul ahavg 1>^h* iiiti(i>>« 

Th* lowuvanlly 

bora«tor the UiicrfaBab rormed by ihe Dincui-fciiuuDour b*r. The quaiSmie U on tlM left 

usually connected behind the orbit by a process from the for- 
mer articulating with another from the juga!, and thus forming a 
bar or arch which may be termed the postoriital bar or arch. 
Similarly another bar at the posterior extremity of the supratem- 
poral fowa is foinicd by the quadmtojugal (or quadrate), squa- 
mosal (the supratemporal of fig. 894 being absent), and parietal, 
and may be termed the fostftmporal, or parietft- squamosal fiitr, or 
arch. A third fossa behind this bar is termed the posUemporai 
fmta. It will be observed lliat these numerous vacuities expose to 
view the brain-case lying within these arches or bars; and there 
seems to have been a gradual tendency to open up the completely 
roofod skull of the I^bjTinthodont Amphibia till in tlie Mamoiab 
we find, as already monlioncd, the squamoso-maxillary, or zygomatic 
arcade, and often a pot^lorhical bar, as the sole remnants of this 
primitive secondary roof. 

It will not be necessary to mention the various neural and vascular 
foramina of the cranium, but it must be observed that on the upper 
aspect there are paired apertures for the orbits (fig. 8j6). and either 
paired or single onca for tlic tares {ibid.); while between these aper- 
tures there may be the paired prtorhitat vacuitits, which are shown 



Ibe fiEUre of the skull of Phytesaunts among the CrocodilU. 
Macion muu ako be nude of ihe parietal Jhranun (Hg. 834), 
vfaicb is 1 vacuity occurring in the paricLils of many Reptiles and 
AiD|ili)biuu. In so-eral living forms this foramen immediately orer- 
Itei <n aborted cneduin eye embedded in the subjacent ti^isucs, and 
^littlly fuDctionle&s : but it i» not improbable llial in t)ie Labyrin- 
aod other early forms this eye &en'cd the purposes of 
As the attention of the pala-ontologist is often directed 
utAcnt, we must also ohscrt-e that the EvttoihiaH lubtt arc canals 
wnnening ibe internal cor with the pharynx or gullet Apart from 
IB minor ossifications which will be incidentally noticed in the 
I, vc oiost also call attention to the feriolu and /ympanu of 
Mianiails, since among the Cetacea these hones are of great im- 
pnancc to the palaeontologist. Both thertc bones are ronncclcd 
the internal ear, the first resulting from the coalescence of the 
epiotic, and opislhotic of the lower forms, and containing 
dx wcUia of the ear; while the btter is foniied by ossification 
Id tic tissues arouTHl the tympanic membrane, and also occurs in 
le birds. Finally, the term lympanU rin^ is a tonvenit-nt one 
Flo ^fiy to the bones surrounding the external car of Reptiles, and 
opBBllf the Chclonia. 

In iqfard to the muHdihU, or lower jaw, which wc have already 
tmA to be formed hy oiuificaiionK in the region of Meckel's car- 
tibje; each half, or ramut (fig. 827), in the Sauropsida and Am- 

ea t fy 

■ new (T ito W« r^ut of iIk msodiUt of CnaJilm. >/. SKii<I'hy>i> i 4, 
KfrbMkl; t*, Cwoiwalt w. Anj(iitu ; (■■, Sunngulir; nt, Aniinilar. Kiilucai]. 

pbbia contists of Ihe following five ouifications formed in mem- 
bnne — viz., detUary (J), ifkniat (j), a'roimd {iv), angular (««), and 
UraMgti/ar (/*), These unite with the afliatJar (ar), formed from 
Meckel's cartilage, which articulates by a glenoid eavifji »ilh the 
quadrate. In Mammals, however, there is but a single membranous 
osatficacion in each ramus, which posteriorly articulates hy a rounded 
nMu/y/f with Ihe squamosal bone of the cranium, there being iippar- 
entjy no articular osiification, and no distinct quadrate in connection 
with the mar»dible.' In the Sauropsida the quadrate articulates 

* Swt]i« biuoducioty ch«ptM on lie MamniAliatCHiipterKii.] 



directly with the periotic region; but in the greater number 
Fishes the mandible, as akeady mentioned, is connected with 
cranium by means of the hyomandibular suspensorium (fig. 823). 
Having now noticed the leading features of the axial, we nuqr 
proceed to an equally brief survey of those oT" 
the appendicular skeleton. In all the hi^icr 
Vertebrates the limbs are divided into thnt 
sections — viz., in the pectoral or fore Hmi 
the arm, fore-arm, and the hand or maimt; 
and in X\\e pelvic or hinder limb the thigh, i^ 
and t\i.efoot at pes. The first s^;ment lua t 
single bone — the humerus of the arm, and tfie 
femur of the thigh (fig. 828) ; the second hil 
two parallel bones — the radius and ulna in 
the fore-arm, and the tibia and _fibula in die 
leg (tig. 838); while the third segment con- 
tains a number of bones arranged in not moie 
than five longitudinal rows (figs. 828, 829)^ 
with the exception of the Ichthyopterygiu 
Reptiles. The bones of the manus and pet 
are again divisible into three sections — vit, 
proximally the carpus or wrist in the maniu 
(fig. 829}, and the tarsus or ancle in the pes; 
mesially the metapodium, or metacarpus of the 
manus (fig. 829), and the metatarsus of the 
pes (fig. 828); and distally the phalangeals 
(figs. 828, 829) of the digits. With the a- 
ceptton of the Ichthyopterygia, where the 
normal digits appear to have divided, in all 
known forms the number of functional digits 
does not exceed five; and these are enum- 
erated consecutively from the radial or tibial 
side, so that the pollex or thumb of the manus, 
and the hallux or great toe of the pes are always termed the first, 
and the little finger and toe the fifth digits. Except in the hallux 
there are usually not less than three phalangeals in each digit 
of the pes, but their number may be reduced in the manus. The 
bones of the metapodium correspond in number with the digits, 
and consist of a single transverse row. 

Although, as we have stated, the number of digits in the higher Ver- 
tebrates is typically five, yet there appears to be considerable evidence 
that the number was originally seven. Thus in many pentedactylate 
Mammals, and also in some Reptiles and Amphibians, there is found 
on the radial or preaxial border of the carpus or tarsus a small ossification 
which Dr Bardeleben terms the prepoller, or prehallux, and regards as 

Fi^. BiS. — Donal upeci 
of right ianomtiatc vni 
pelvic limb of the Oiiin- 
puuee ( Trv£ladjit{x\ re- 
duced, i, Innominalc ; /\ 
Femur ; t. Tibia ; i, FJbub ; 
r, Tantu : 101, Metuanus ; /, 
PhBlangeala, (After Owen.) 




ftcKpTDcatalive of u additional radial digit. S\m\\iir\y the jiist/i>nn 
btM albc oupos, which occun oa the ulnur or posiatial boidcr, and in 
Hi—nili t» usuaUy described as one uf the so-called sesamoid bonu, Is 
boted qnn b; the suae authoiity as the icprescDUlirc of a seventh 
Apt tm tbc ulnar side. 

llii(VobabIe that the nrprn and tarsiat were original!)- formed 

qn a ctxiunon t)-pe, which persist:* in a more or less unaltered 

oaidilion in certain Amphibia and Reptilla {fig. S39). In such a 

((MialiMd tji^ there 15 a di&tal row 

flf fire cupaiia (fig. 839), or Lirsnlin, 

Mkuliting with the iDctapodials. 1'his 

ii pnctded proxjnially by another row, 

CBRHitiflg of an intertiudium, flankctl 

in tV monus by a radiaU and uinart, 

-■ Ihe pes by ft /ih'aU and 

, twpectivdy articulating with 

' epipodial bones of the fore- 

idius and ulna) or leg (tibia 

' ■ mb). The middle space l»c- 

ttem iheae two transverse tows of 

bom b occtjpicd by one aniraie, or oc- 

osianally by several antralia- Modili- 

mioas from tbis type are caused by 

(fas oppression or coalesceooe of Mine 

of these elements. All the carpals 

ad tarsals in ihc Mammalia hare re- 

ccired distinct names, which will be 

noticed under the head of that class \ 

Un it may be ol»en-ed here that the 

faigber Reptiles and Birds agicc with 

liie former in having two bones in the proximal row of the tarstis 

— lii;, the etUotncMm on ihe fibular, and the astra^alHs on the tibial 


The protuberances for muscular attachment at the proximnl ex- 

^■tnity of the buinenis are termed tubfrfiilia, while those of the 

^^■uf ore known as troihanters. The latter bone in the Saur- 

^^idft may also haw an tniur troihanUr on its shaft for the at- 

' tsdunent of \\\t femom^oudal muscle ; while in the Mammalia there 

may be a tktrd trwkanter for that of the gluteus maxSmus, as is 

I thown in the femur of JiAinoetros represented in fig. 1*36. Both 

the humerus and femur have more or less distinct (ondyks at the 

. lover extremity for the articulation of the bones of the forearm and 

leg — chose of the humerus being often termed innkUa. Above the 

cmdyles or trochlea the humerus has projecting tpiojndyiti on 

eiihg ftiik; and there Is frequently a foranicn bituaicd above citbei 



Fil. *Tti — norul u|«t or lh« 
it|ht manut of a Cbt|i^U*> K<blll« 
(ClulydnX ra. RaOiia: ml. uitix: 
r, I, a, K.ul<iili, ulnuv, •!>■] imcT- 
medlum; f. Ccnuik; 1 — s,C>rpalia; 
m, MemUfpali: /, pMunul p(ia|. 
>iUte>)>i i-'v.. Ttnnttul Ju., I. bcint 
ihttnllo. (Ariel CtCfnbkut.J 



the outer or inner eptcondyle. When placed above the outer or 
radial epicondyle this foramen is tenned ectepicondylar, and wfaa 
above the inner or ulnar epicondyle entepUondylar. The proxtmd 
extremity of the ulna is often produced into an oUeranon (fig. 1300^ 
which projects behind the end of the humerus. The distal a- 
tremtty of the tibia in certain Sauropsida develops from its anterior 
aspect a ridge or process known as the cmmial crest; whUe the 
deltoid crest or ridge is a prominence situated below the head of tbe 
humerus on the radial side, to which the attention of the paheoD- 
tologist is not unfrequently directed. Finally, the patella is a so- 
called sesamoid bone developed in the tendon of a muscle passing 
over the pulley-like surface or trochlea on the anterior aspect of tlie 
distal extremity of the femur. 

Since the limbs of Fishes differ considerably from the higher type 
of structure noticed above, their consideration may be deferred till 
we come to that class. Before, however, leaving the subject of 
limbs, it should be observed that it is often convenient to allude to 
the corresponding or homologous sides of the fore and hind limbs 
by a single term. If, then, we imagine the limbs extended more or 
less nearly at right angles to the axis of the body (as on the left 
side of fig. 814), with the palm of the hand and the sole of the 
foot directed to the front or ventral aspect, the middle digit of 
each limb will be axial, when the poUex 
and radius of the pectoral, and the homo-- 
logous hallux and tibia of the pelvic 
limb, will be obviously preaxial ; while 
the fifth digit of each limb, together with 
the ulna and the fibula, will be postaxial. 
The whole of the radial and tibial sides of 
the limbs will accordingly be known as the 
preaxial, and the ulnar and fibular as the 
postaxial border. 

The remaining parts of the skeleton 
comprise "dxt. pectoral xaA pelvic girdles, by 
which the corresponding limbs are respec- 
tively connected with the trunk, and of 
which the relative positions are shown in 
fig, 818. The pectoral arch is never con- 
nected by means of ribs with the verte- 
brae, and primitively consists of three main 
bones developed in cartilage. On the 
dorsal aspect of the body we have the 
upper bone or scapula (fig, 830, j) ; while on the ventral side there 
are two parallel bones (fig. 974), the anterior of which is termed 


Fig, Bjo. — Laleril Mpecl of 
tbe carlilBf[e boiKl of the left 
>ide of ih« ptciaral girdle of a 
Liiard {Iguana^, t. Scapula :, MesofCapular process of 
do. ; cer, Coracotd ; f.cor, Pre- 
coracoida] process of do. ;, 
Me^oconicoidal proccsi of do. ; 
fyr, Foramen of do. J gl. Glenoid 


lu fmaramJ (cf)y and the posterior the foraioid (<»). At the 

miction <3i these three bones ihcrc is a caviiy for the aniculatlon 

of the bend of the humerus termed the glenoid eavity. Such 

kdK (tfijiiitirc condition of tht» girdle ; but in Reptiles the pre- 

H^ooid very rarely exists as a separate Rssific;tlion, ntthough it 

flw so among the Anomodonts, where, at least in the young, 

it fcran ft large plate, uniting belou- 10 the upper edge of the cora- 

tod, Bod entering into the formation of the glenoid cavity.' In all 

aba cise« it is, however, completely fused cither with the scapula 

n tbc coracoid. Thus in the Chclonia (fig. 1008) the prccomcoid 

mins its primitiii; forni and condition of a transverse bar, which 

i^ bowever, completely fused with the scapula. In the LizAids, 

on ihe other hand, this bone has united with the coiacoid, of which 

II fonns the prccoracoidal process {fig. 830, p.eor) — the foramen 

[V) marking the original line of reparation between the two bones. 

In Dinosaurs and many other Reptiles the prccoracoidal process 

bu diiappeared, and only the foramen remains ; while in the 

thfhytppter^-gta even this is wanting. A further reduction occurs 

in the higher Mammals, where the whole of the coracoid has dis- 

afpatltd as a distinct lionc. The coracoid is subject to threat I'ari- 

ttipa in ^lapc, and may cither simply meet its fellow by an overlap- 

Wtg or sutural jurKtion, or may aniculatc with the sternum. 

^i ciavkUj mainly developed from membrane, may be connected 

■ilfa tfae preaxial borders of the scapula and coracoid ; while a 

QCkial T-^upcd interxlai'UU (fig. 819), which is also developed 

from meml^rane, m,iy receive the inner exlremities of the two cla- 

m.\ts, and then usually overlies the uppt-r part of the Alcrnum. 

Fitully, there may also be a single or double mesinl omosUmum 

Icveloped on the ventral aspect from cartilage lying near the an- 

eriar extremity of the girdle. An illustration of the [xisiiion of this 

tone is shown in the pectoral girdle of the Frog, represented in fig. 


In the pelvic girdle we have three separate ossifications arising in 
anil^e, of which all three usually unite to form an aeetabulum 
fe ftsif' a. i) for tbe reception of the head of the femur, or thigh- 
xme. The bone commonly kiwwn as the haunch-lione forms the 
lonal dement, and is technically termed the ilium {ibid., Ji) ; it 
xwmponik to the scapula, and usually .iriiculatcs witli the sacrum 
)y short ribs ; while in the ventral half we have anieriorly the puNs 
«hirf'., P), TcpTcscnting the prccoracoid ; mid posteriorly tlie ii(hinf>t 
|iN/., /*), which corresponds to the coracoid. The two latter bones 

» ly&t piWOTicoW appeara to conwpwid with iIip '««ie \tmeA tpUaromd by 
Ptafaaor Copt, aa4 alto the one w> nMotd in the M'lnoitrmc Maninink. 

• llcre the *«t»twlum if fo.m,«d hy only iwn l>imcs. and it hii been suffgestcd 
^tX the iMBe ktleted pubis may be really an vpi)iubU. 



very generally meet those of the opposite side in a ventral syift 
physis ; and when, as is ftequently the case, the three bones ol 
either side are anchylosed together, an innominate bone results (fig. 

ii28^u'). The ischium and pubis 
of the same side may also unite 
inferiorly so as to enclose the 
obturator notci, F, which thai 
forms the obturator foramen (fig, 
ii28^m); whi]e an obturator /Pro- 
cess of the ischium may form 
a smaller foramen below the 
acetabulum, as in the pelvis of 
Camptosaurus, figured in the 
sequel (fig. 1052). As a rule, 
there is no tendency to a reduc- 
tion in the number of the pelvic 
bones in the higher fonns. It 
may be mentioned, in conclu- 
sion, that a median ossificatitn 
at the ventral symphysis of the 
pubis and ischium found in cex- 
iain Edentates has been named the /ehtstemum, and regarded, as 
the abdominal representative of the sternum. This appears, how- 
ever, to correspond with the median ossification found in the 
Ungulates (fig. 112$ bis), which is generally looked upon merely as 
an epiphysis. The pectoral and pelvic girdles of Fishes are nodrad 
in the next chapter. 

It may be well to observe here that genera being purely and simidy 
artificial divisions formed for the convenience of classification, it is quite 
unnecessary that they should be of equivalent value in different groups 
of animals. As examples of vertebrate groups in which generic temu 
are used in a wide sense, we may cite the Reptilan order Sauropterygia 
and the Mammalian family Rkinocerotida: \ while as instances where a 
more restricted application is employed, we may mention the ordei 
Chelonia, and the family Bovida, The statement we not unft«quendy 
hear that such-and-such a form must represent a distinct genus implies 
a total misconception of the import of generic terms. 

lor nolch 

uu. . J I 1 VU»9 J Sty AKUIUIU p J ■ \^U1U1B- 

Dlch: tt, i, 6, Acclmbulum, with iu va- 
; t, , Cartilagiooiu utenuoni of illam. 



Gekeral Structure, 

:inanbcra of the cloiis Pisces, commonly known as Fhhcs, form 
Ihe fim division of Professor Huxley's Ichthyopsida, and are gener- 
lifr cbAncuTuett by living in waicr; breathing by branchic, or 
^Bs, ibfooghout life ; having the heart furnished with a single i-en- 
Bide and auricle (atrium); having the limbs, when present, in the 
kna of liru; beinj; proviJed wilh unpaired median fms supported 
bf fiiMsys ; and by the skin being cither naked, or covered >vilh dcr- 
M] scales or bony scutes. There is no amnion or allantoi* devel- 
oped in the embryo^ and the reproduction is nearly always oviparous. 
CWain forms do not, however, exhibit all the above features, and 
Uie relation of the more generalised Ki»hcs to iJic Amphibia is ver>- 
inttmate. llie peculiar system of mucous canals and the lateral 
Kae are hi^y characterisitic of Fishes, althoujjh they arc not 
irmDiabljr picsctii. 

Before noticing such features as arc of especial importance to the 
palKoniologisi, it will be convenient to mention that according to 
the arraj^ement adopted in this work the class is divided into the 
foOowing six orders — viz., Cyclostomi, Elasmobranchti, Chimeroidci, 
DiptMi, Gartotdei, and Tclcostei, of which the salient features will 
be noticed in the succeeding chapter. I>r (iiinthcr has, indeed, 
proposed to bracket together the second, third, fuurth, and fifth 
ofdcrs as a subclass under the name of FaUcirhthycs, ranking as 
cqui^'alenl lo the CycloMomi and Teleostei. Professor Huxley, I)r 
Traquair, and otben have, however, shown that the Ganoidci are so 
iBttmaiely cooneried with the Teleostei, while the F.lasmobranchei, 
Qlimeroidci, and Dipnoi differ in so many ri>spccls amotig th^-ni- 
sdvea and from the foniwr, tluit such a grouping does not appear 
rantonont vn'th their tnic relationship. 

Aaotber scheme, proposed by Professor Cope and adopted by 



Fig. Sja.— C)-c1oid scale, 


Fig. 833. —Ctenoid scale, 

Mr Smith -Woodward, is to brigade the Teleostei and Ganoidei 
together in a subclass under the name of Teleotomi, with tmlinal 
divisions differing somewhat from the subordinal ones employed 
below, and to raise the Dipnoi, Chimeroidei, Elasmobranchei, and 
Cyclostomi to the rank of subclasses, with the concomitant eleva- 
tion of their respective suborders to the tank of orders. It seems, 
however, scarcely to harmonise with the divisions adopted in the 
other classes of Vertebrates to regard a group like the Chimeroidei 
as a subclass, and accordingly the view of Professor Huxley is 
provisionally followed of regarding the Elasmobranchei and Chi- 
meroidei as divisions of ordinal value. 

Although the body in all Fishes must be adapted for progressioo 
through the water, yet there is an enormous range of variation in its 

contour among the differ- 
ent groups, as we may ob- 
serve when we contrast a 
Lamprey, a Shark, a Flat* 
fish, a Ribbon-fish, and a 
Globe-fish. The dermal 
structures termed scales, 
which are so characteristic 
of Fishes, present many 
types of structure. In the Teleostei they usually form thin plates, 
frequently marked by concentric lines and not formed of true bone 

When the posterior margin is simple 
(or entire) such a scale is termed Qfdoid 
(fig. 832), but when denticulated, ctett- 
Old (fig. 833). Other examples of this 
type are shown in figs. 834, a, b. In 
many Ganoids and a few Teleosteans 
the scales are much thicker, and consist 
of a variety of true bone covered exter- 
nally with an enamel-like substance 
termed ganoine. Such scales, of which 
specimens are shown in fig. 834, e, and 
fig. 835, are termed ganoid ; they are 
arranged in oblique rows, and connected 
together by a peg-like projection, their 
shape being oblong. Scales of ganoid 
structure may, however, be much thin- 
ner, and resemble the C)xloid type in 
their contour and their imbrication. 
Lastly, the bony dermal scutes or plates, frequently armed with a 
spine, which occur in the skin of the Sharks and Rays (fig. 834, 
c, d), are strictly comparable in structure to teeth, consisting of\ b. Ctenoid uilc of Pereh 
{PtnaY, c, Dernul pltiie of Thorn. 
bacV iSaia): d. Do. of Monkfish 
(i'^Bii/iM); t. Ganoid scales of Pal^f 
OHiiciH. a ud' enlarged. 



pectoral fins may be dtvtiopcd into huge dermal spines, which in. 
some cases articulnte by a complete shackle-joint with the lusai 
bones. Similar spines arc also developed in the Elosmolininchei 


Fi|. t]T.-Uft tannJ vitw rt th« P*reh KFtrtaV *, Gill-caTCi, "rllb gJU-tlh USInd li ; A he- 
iMtlKa; n, Palri<do.: ^nncdonal 6a.% d', Sccaw] doful dix ; f, M(«f»l Im*. KeducM. 

(fig. S38); Iiut here they are ainiply inserted into the 6csh, or are 
atuched only by cartilage, sw that iheir basal end is rounded off. 
Siniilar spines (fig. 83S, i) may aJso occur behind llie head. Such 








riK- S)!.— Spltci and Iceth of EUnnohrinchd. i, Nuchil ipinc fX Flnruc^ttuit I *, Pin- 
^ \'>o. al Cltm»fty<Uiu. UiiloiiKctaiu. i-t wcrcduoa). 

Spines when found fossil are frequently known as uhlhyodenUhs, 
and form important object* to the pnla.'otnologist. Their structure 
is identical with that of the sonialled placoid scales and of teeth. 




being 1 central pulp-cavity, surrounded by a layer of dentine 
'ermtieaaat, which U corered with enAincl. 

He leeifa of Fishes present a g;reater amount of variation than is 

fo«Mlin sny other class. They niay be entirely absent, or may be 

(nsQil on all the bones of the muuth, and also on the hyoids and 

bnndMl arches, while they may be attached merely to the nn-m- 

bme of the mouth-cavity. Very frequently they arc attached by 

ladiylisiit to the underlying bone or cartilage (lig. ^36), but they 

n^ he implanted in di&tinct sockets or alveoli. The dentine is usu- 

ill; dininguished from that of the teeth of higher Vertebrates by its 

ptata mculnrity. The coating of enamel is 

ItBoalljf TCry thin ; but it is more developed 

A tile cutting-tcdh of Sargvs. Occasionally 

(Am^mfiu) radiating prolongations of the pulp- 

ovity may penitrale the dentine from the centre 

B 4e pcripher)-, tlius producing a structure like 

i*ai of the teeth of the Labyrinthodont Am- 

piiibtUM. There i* generally a constant renewal 

al the teeth of Fishes during the whole of life ; 

boi occasionally one set persists. 

Tuminf; to the endoskeleton, and commenc- 
ing vith the vertebral column, we find that the 
imebne can only be divided into a trunk and 
a cuMbI series, and that there is a gradual pro- 
pasioa in lespect of osfiification from the lowest 
to the highest forms. Thus, in the Cyclustonii, 
thr noiochonl persists throughout life, and is 
SeoeraJly unscgmcntcd, although rudimcntal 
neural arches and spines are developed in /V/- 
mmysom. The vertebral column of the Carli- 
liginou!! Ganoids is very similar to that of the 
bOer ; but in Bony Ganoids, Elosmobranchci, 
tnd Teleostei paired cartilages, arising both 
ilxive and below the notochord, gradually sur- 
round it. and thus form ^Irongly nniphicceloua 
vertebral centra. In the Sharks these centra 
exisi without arches, but in the other orders 
there are wdWc^'eloped neural arches; and in the trunk region 
^ben arc also lateral basa/ pnxtitti, which in the tail unite infcriorly 
to Ibfin a hamal arch for the caudal artery-, .ind develop a hx*mal 
qiine <fig. 839). There is a great tendency for the neural arches 
to remain open superiorly; and the only Fish in wliicli the venebrK 
arc not amphicoelous is the Ganoid l^pidostnis, in which they are 
opisthoccetous. Oiily in the Chimeroidci and certain ELtsmo- 
ane there definite articulations between the vertebral 


Fie- Sjt).— AalerloT »■ 

of ■Tflconl'aii Fhb tti, 
Ntural .filne: •■•. N»u- 
ml ureh; «, Aiiitular pro- 
r«itH ; ha, HmtniX nrrh ; 
k; Hsoial tpinc {Afm 


column and tlie cranium ; the poi^terior aspect of the basioccipl- 
lal in other groups forming a cup like that of a vertebral ccotnun. 
The mode in which the vwlcbraJ column tcrniinates posteriorly it 
of considerable importance in classification, The most primitiTt 




Fig. 84a.— A, f»l]t*Unu. uid Di OittBUfiU (Caoaidili n ■liea' diphTccnal caudal ifc 

a, Pcciont; t, Ventnl; c, arul: if, Darul dn. RnliKed. 

type occurs in tht Cyclostomi, Dipnoi, and many Ganoids, where 
the nntochord continues to the extremity of the body, and ts sjiii- 
iiietrically surrounded by the caudal fin, as in fig. 8+0 ; this type is 
known as dijfkyarm/. In the other, or A-Arroarcfl/ type, thenotochord 


ft, Swocd-fiih : D, SldrBfrtn, w ihnw Ruukwl (hotnonru]} and typk*) 

is bfnt upwards, owing to the greater deielopment of the lower as 
compared to the upper half of the tail. I'his feature may be observ> 
able externally, as in the Sturgeons (fig. S41, b) and Sharks ; or may 
be maaked, as in the majority of Tclcostcan Fishes {fig. 841, a), by 
the s)'in metrical arrangement of the fin-rays. The skeleton (fig. 



Plf. Sta.— Tail «f FtoaiHlvr. (flftri f\Ttaii ) 9. 
VtrttlR*) calnnui- ■, Turntd-up tnd of lb* im*- 
dbsid : ^a Hxpunu booa. 

1) dio»5, howcrer, the apward bend of the nolochord, although 
Ikauftrf nuch less marked ia the adult than in the young. 'Hiis 
vaAtA. bctefo cer cal type is sometimes described as homocerat/. 
Hk coilaced tuemal ^Mnes found in this type of tail arc known as 
kjftrat bones ((ig. 84s, 4), 


«bile the OHificd extremity 
cf ihe DOtocboiil u termed 

The skulls of Ftsbca pre- 

mt nnaitons in icgud to 

Adr d^rec of ossification, 

aa)ogoi» to those obtaining 

dtte vertebral column. The 

fntnl uniciure of the pri- 

nitnecwtikginoiii skull has 

lea alreatly irxlicated in 

CbqKtr xhr. (fig. 823), but 

»• most here glaiKe briefly at certain bones developed in the more 

y^B*^ forais which are peculiar to the class. Thus, taking as 

10 oample the idiull of a TdcoMcan Fish (fig. 84^), where die 

pimttivc cranium is 

CDoccated by the dc- 

ttlofaait of inventing 
bMle^ we find two 
peculiar ossifications in 
ihe aoditory region 
howl) as the pcerotic 
(JW, Pie), which is 
ooealiivm\ to repre- 
sent the squamosal and 
oputiwcic of higher 
Venebrates, arxl the 
tphenotic {i^d., S/A). 
A laige panisphcnoid 
is itlwayE present infe- 
rioriy (fig. 844). The 
intervention of the hy- 
otnajidibular and sym- 
plectic {Ct^ 833 and 

rale and the squamosal '*•'■ A«»io«tar; J?t, D«uiy; ^, ite. (AIwi WnJuihsim.) 

region has been already 

mentioned in Chapter xW. as peculiar to Fishes (although ii Is by no 

universal in the class) ; and ve must also mention that, in 

ition to the rwnnal palatine and pterygoid twncs developed 



Fia. If).— ■''^ iMKnil nrm ef the ttull <rf % Traul (Salm^Y 
A'A ^ri*""^; /^t, PmimEc; J>t, SphtnoiK; <'i. Sum- 
ot< ■ * I'lrietal; A Fnnul . i^etk. F.lh'ri'iiiJ ; (m, 

Ai' L9<y(xtvq; jVy, Niim.1 i /'«<», IV*ma"iilji; 

ii. ■ • . 'x, Jujio'; l*f'. Me(opnryi;«ii1 ;, 'W/, 

t, Sjvvkcik 1 Qm, Qublnue : /V, U/, Sfj-, )'ib-, >»icr-, and 



round the cnnilaginous iwlatopterygoid bar, there occur in Teleo&ti 
the mfso- and metaflerygoid (fig. 84 j, Mt, Mfp). \\\ the same 
the orbilat region likewise develops a series of membrane-t 
round the eye, forming the sttborhitais or orbital ring {ihiJ., 
while the ^■"'^i'''"''' «r operculum (of which the first trace is founc 
the Chimseroids in a fold from the hyomandihular overlapping the 1 
first gi!!-slit) is formed by the preojvrttiiar, aptratlar, suboperaclar, , 
and iafraoptradar {ibid.^ Pr, Op, Sop, Jop), which are broad, scale- ' 
like membrane Ijonci. In the branc/uostegal memirant, whicb 
unites with Ihc gill-cover in closing the branchial chamber, there ■ 
is developed a number of iiraachiosfegal rays {fig, 843, fisS, and 
fig- 844, Ao) ; but these may be partly or entirely replaced bjr , 

^In- B14.— R!chl ude of the cianiuin ur»]Kr uf « feich, losether viib ibc hj-otd 
chuTsnheK and Ihi aalvlc Dtrdlo. i*. Ki^'I^Aynl : t». Kplhml : rk, Centoh>'al : M. 
^1 GloMohytil 1 Hi. ITrutiyil ; h«, BrnnUiiinlcuiiI ray> ; ri^, CcrtlelirBiKlil*! ; tt, "f" 
Xr, On* oT iha "^tll-rsVin " of itic (itvi IiriDLliLiI anh; /a. Pamphmpid : t». Suptwodptal ; 
M, PbuUHporal; kI. Ku|>ru:Uiia.ul>f ; ■/. dnvitolari fcl, 'I'be tsopatco oTtlic poMcbifaa- 
lar ; ac, Sopuli : t», CorsMid. 

jugniar plaits, occupying the space between the rami of ihc man- 
dible. The hyoid arch (fig. 844) is attached to Ihe inner side of 
the hyomandibuliir by a stylokyal {i{>id., sfi), articulating infcnorty 
with the tpihyai (tK), and the latter with Ihe large etrafohyal (ek)', 
the two latter carrying the branchiostegals. The inferior pett of 
this arch is formed by the batihya/ {bh), from which the ghsiohyal 
{j:h) extends forwards ijito the tongue, and pOEttcriorly articulates 
with the first of the basibranthiais, mentioned below. The urokyal 



I in wrticaily-cooi pressed median bone, extending backwards 

dw huibya^. Bcliind the hyotd aicli occur tlic brixruhial 

. the ftrsl of whtcli consists of a tnedinn i-w/Mi****;'.!/, and 

ilnenlljr from below opwards of a Hypohranehiai. <€rati^ran(hiat 

(fijt 844, r*), tfihramhiai (M), and pharyHgobranehml. The lallcr 

toco ill the second and thud arches are tailed sujvrior pharynj^aif, 

Jly cany teeth. Finally, the f^'/l-rakfrs (fig. 844, ^-r) arc 

ke bones attached to the inner margins uf the branchial arches. 

b the mandible there is usually (lig. 843} 

1 1 dtnl»7 snd articular piece ; but an angu- 

\tt,»A more nuely a splenial or coionoid. 

Bay tUo be prcMRt. 

In [be appendicubr skeleton, we iind the 

ptOml girdle of Ganoidit ainl 1'clcoateans 

csuiuiKg infcriorly of llie primary cartiia- 

fetm clement^ corrcspoodtng to the sca)i- 

nil ml coracoid (5g. 844, ir, co), and 

■pnicfly Add Ulcrally of a secondary chain 

cf booet developed from membrane, and 

tiwdiling supenorly with the ptcrouc 

of the skull. The hones of this 

chain are named Iram above 

fioiftemporal <fij{. 844, //), w- 

(W), elavuuiar {fly, and a 

ir of two pieces (/W) ; while 

may be also an infriidavU -'»r be- 

loi the clavicular. In Elasmnbianchci 

ooly the eanilaginous primitive girdle is 

dndoped; while in Dipnoi the girdk- 

i td very peculiar stnicturc, and some- 

riiat tntennediate between thai of Elas- 

nobtanchei and Tdeosiei. The pelvic 

ptdic is generally wanting ; bul in the 

D^moi there is a median cartilaginous 

pJate, with anterior and posterior paired 

prucesKS, of Khicli the former are iliac. 

and the taitCT give attachment to the hind 

baJbt. Elasmobranchii generally show a 

degenerate pcU-ts of this type. 

Tbc pectoral and (Jclvic limbs, or fins, arc 
so simdar in structure thai ihcy may be eonsidcrC'd togeiher, 
altboagh the development of the latter is less specialised ihan 
of the former. No representatives of the arm and forearm 
the higher Vertebrates can be detected in Fishes, the b.isal ami 
bones or cartilages articulating directly with Uie pectoral 
vol- II. c 

Mb. 8«j.— SkckUiiiuiUn Wl 

lytClorilllBibnfCrfnt'i^"*- "• 
Jutcd. (A/ur GCalll".) 


girdle. In the Dipnoi, and especially in Cerdtodus, the pectoo^F 
(fig. 845) and pelvic fins are supported by a cartilaginous, medu^ 
segmented axis, bearing jointed radii on the dorsal and ventni 
borders — these radial cartilages being terminated by homy demit 
fin-rays ; and the dorsal radii (left side of figure) being more nunw^ 
ous than the ventral. This type of fin, which also occurs in the- 
Ichthyotomous Elasmobranchei, is known as the archipterygam. 
From this slightly unsymmetrical type of fin that of existing Sd~ 
achian Elasmobranchs (fig. 846) may be derived by the gradual b^k 
pression of the ventral series of rays, and the development of tie 
dorsal, which has now become lateral Basally the jointed radial cv- 
tilages articulate proximally with the pro-, mm^ 
and metapterygium, which in their turn are tt- 
tached to the pectoral arch, and the latter oT 
which corresponds to the basal axial caitilige 
of the fin of Ceratodus (fig. 845). In 4e 
pelvic fin of the Selachians the mesopteryginn 
is absent, and the propterygium more or lev 
rudimentary. This type of (in is known as the 
khtkyapterygium. The fins of Ganoids and 
Teleosteans may be derived from the Selachian 
type ; but the primary cartilaginous skeleton is 
Fig. 846.-skdeion of educed, and a secondary one developed by the 
itw left ,p«ior»i limb of introduction of membrane bones. 
>, Pro-; fv, Mew: »/, ficforc leaving the structure of Fishes, men- 

Weu^erypum. Much ^^^^ ^^^ y^ ^^^^ ^f ^^^ otoHtfU, which SIC 

small, rounded, elliptical bodies, usually with one 
convex and one concave side, tying in the tympanic sac, and com- 
posed of both calcic carbonate and phosphate. These bodies have 
been carefully studied by Dr Koken, and several genera identified 
by their evidence in a fossil state. 

As regards their distribution in time. Fishes being the lowest 
class of the Vertebrata, it would naturally be supposed that they 
were the earliest representatives ; and this appears to have been the 
case. The earliest known fishes in Britain belong to the Ganoid 
group Placodermata, and occur in the Lower Ludlow group of the 
Silurian ; while the Elasmobranchei were represented in the topmost 
group of the same series. In the Devonian and Carboniferous 
periods Fishes become abundant ; but all the forms from these 
horizons, and up to the Cretaceous, belong to the Elasmobranchei, 
Chimseroidei, Dipnoi, and Ganoidei — the specialised Teleostei not 
making their appearance, so far as we know with certainty at [wesait, 
till the Cretaceous. The Ganoids of the suborder Amioidea ap- 
proximate, however, so closely to the Teleostei, that it has been a 
question whether some of the members of the Jurassic family L^to- 



•iiicb ore usually placed in the former, should not be 

[tmuferred lo the latter group ; and we ma^ ihu^ t-onfidetiUy expect 

Liofifldacuic|ilete tninsiiion between the two, Alihough many of 

jibe Cretie«oa$ Tdeonei ore more or less closely allied to existing 

jljpesv il a iKii until the Eocene thai we find a fish-rauna conipar- 

I able Co that oT the present day; and we m.iy note thnt the resem- 

; bhaac oT the /ishes of the Roccnc to those now living is in marked 

tsMist to what obtiiins in Maminals, whcrtf the m:ijurity of Eocene 

lEnea are extinct. The persistence of i>onic genera of Fishes 

Ibraugbout long geological epochs is indeed a noteworthy cir- 

omtanee, and is nowhere more marked than in the case of 

CmasAa, which has lived on continuously from the Triassic period 

I^^Eutofic, and also from that of certain North American beds, 

< idifefc tie luually referred to the Permian. 

In rq;aid to the orijjin of the various orders of Fishes, it is pretty 
I oidcnt that the Telcostci were derived from the Ganoidei, »nd thai 
rthtUipooi were closely related to one branch of the latter. The 

MMMrf ««7T^. Fran ik. sIlxM ot RuHla; ■, Frool lU Cub»niF«»w e* Nonh 

pfayk^enetic relationship of ihe Ganoids to the Glnsmobinnchcans 
b, however, still unsettled, although recent rese-irches tend to show 
a dose connection Ijetwcen the more piimitivi; and least specialised 
glKKipK of the two orders. Palseontolog)- has not, indeed, yet taught 
IS froinwhat group of animals these primitive Ganoids and Ehsmo- 
bnnchs were thcm«Ives dcnvcd. It has, however, been sugyested 
Professor Cope that the Placodcnnoid Ganoids were closely re- 



kted to the Ascidian Invertebrates ; and if this suggestion 
prove well founded, it would seem to indicate that the grou 
tioned is closely allied to the real ancestors of the class. P 
however, these ancestors are to be sought in another directio 
it has been thought that minute tooth-like bodies found i 
ranging from the Upper Cambrian to the Carboniferous, and 
as Conodonts (fig. 847), are really the teeth of Fishes. It n 
sidered probable at one time that these curious fossils, whicl 
exceed two millimetres in length, might be teeth of extinct m 
of the Cyclostomi ; but their internal structure is so differer 
the teeth of the existing forms of that order, that if they be 
Fishes at all, they must apparently indicate an extinct divisi 
great variety of forms of these Conodonts have been describ 
have received distinct generic and specific names. It is the 
of some authorities whose judgment is entitled to great coj 
tion that these fossils should be regarded as the jaws of An 
or Trilobites ; but the question as to their real nature n 
regarded as still undecided. 

' Vide supra, vol. i. p. 480. 



CLASS PISCES— eonfinued. 

Orders CrcLOsrour and ELASHOHRANcmsi. 

OtBu 1, CvciosTOMi. — The Cyclostomi, which include the 
Hig^Uies ^.Vj-.r/«^ and Bdrliottoma) and Lampreys {Petnmiyzott) 
'•ong It present unknown in a fossil state, require no funher notice 
in the ^toeni work. 

Omm tl. EtASMOBtuscHEi. — This order, which is also known 

■nder the name of Chondropterj-gii, includes a pccuh'ar extinct 

potp temied the Ichthyotomi, together with the modern Sharks, 

PsK^bbcs. Saw-lishea, and Ra>'3, collectively consiituling the Sc- 

hthii, and all of which ate typically of marine habits. For paL-eon- 

Wogkal purposes this order inay be cKaracterlsed as follows : The 

ildeton is invariably cartilaginous, and membmne bones are, with 

Hnt possible exceptions, absent in the skull ; the vertebral column 

i^ horevCT, generally divided into distinct segments, of which the 

CBUn n*ay be marked by a calcification differing in structure from 

We bone. In the skull, which may be t-ilticr movably or immov- 

Aly conncrtcd with the vertebral column, the palatopterygnid bar 

md hyomandibular tiuspensorium are never fused with the cranium. 

l^'bcn an exoskeleton is developed, it coiuists of small dermal gran- 

iies, of which the structure is the same a;; that of teeth. In all 

oistiQg forms the optic nerves simply crOM one another, wiihotit 

icy interlacing of their component fibres ; the bulbits artcriosiis of 

the heart has three series of valves ; the inw«ine is furrished with 

a spira] valrc ; and the ova are of large size and few m numlrcr. 

A few words may be said explanatory of some of the alwve-nien- 
tioned and other features in existing forms before proceeding to the 
vpatmxAc put. In alt fovms, as already meitiinned, there is a 
Kpsme ftapcnsorial arrangement articulating with the cranium, to 
which ihe tnandible is attached ; this structure being termed hymty- 
Ustuily there b a hyomandibuLar sust>cn3onuui inicrvcmng 



es — - 

between the cranium and the palatopterygoid bar ; but in NotUam^^ 
(fig. 86i) the hyomandibular element takes no share in the suppotf^ 
of the mandible, and the palatopterygoid articulates directly vih 

the cranium ; this probably being 
the primitive type. The ^lls (fi( 
848) are attached to the skin ^ 
their margins ; while they usuiBf <- 
communicate with the exterior l^. 
five apertures, or clefts, which mq 
be very rarely increased to six or ' 
seven. The mouth is very gens- . 
ally situated on the inferior aspai 
of the body (fig. 849), and is for- 
nished with numerous teeth carried 
on the palatopterygoid bar and 
Meckel's cartilage (fig. 861). The* 
teeth may be either sharp and sqin- 
ate, or articulated together so u 10 
form a more or less pavement-ltke 
structure J and in the former oat 
there is a continuous succession of 
new teeth developed from bdiind 
as the old ones are worn out Both 
Fig.848.— Dia^mmof anetidcoTthcgiiii median and paired fins are present; 
^'ii;ll;r.S^dT^?ol,"^r.i.''^r.r." the position of the peMc pair being 
;rdi1'*.Xt'hL.'^uVh^™A?r.lS™'ii always abdominal. In all existing 
pu^iii*; I, Sepu twiwcen pouchei; ir, forms the skeleton of the limbs 
""' forms an ichthyopterygium (fig. 

846) ; but in the Carboniferous and Permian Ichthyotomi there 
is either a uniserial or biserial archipterygium, like that of the 
Dipnoi. The posterior termination of the vertebral column is gener- 
ally heterocercal, with the upper lobe of the caudal fin greatly eltxi- 

ba 4 

Pig. S^.—Spiny Dog-fiih lAcmUu'at). KedDced. 

gated (fig. 849). The spines frequently borne by the doraal fins 
and in the nuchal region, constituting the so-called ichthyodonilites, 
have their bases simply embedded in the flesh, and are consequendy 
immovable. There is no swim-bladder. The term spiracUs is ap- 


'napotures connected with re^piron'on found on ihc upper 
of the bead; while ibc interea/ary cartiia^s arc ovoid or 
iped struaures occurring between die neural arches of 

[Soiiifa-Woodward ob>crve<i tlut a gradual adrance in the de- 
Fcalcilication of the axial skeleton may be observed as we trace 
icnbcn of the order from the I'alicoroic upwards, the oldest 
hot having the notoohord differentiated into distinct vertebral 
iof which wc lind ihu liisl complete types in the Iawct l.ia.'i^ic 
buuLT. Here, however, the centra merety assume the form 
pble ooMSi, only very slijjht traces of the pcriphcra] calcifica- 
Iroesury to form the biconcave centra of the later types being 
I In the Lower Kinicndgian lithognipluc &tone of the Con- 
[eentra of the complete tuUrospomiylit type are fin>t met with. 
) same authority mentions that the Palxozic types arc, as a 
(faaiacterised by the great de%'elopmccit of the cxosltelcton, 
Ui a few forms bkc PUuraianlhui appear to have had naked 
\ These early dermal structures are also noticeable for their 
lie sculpture, this being equally developed on the dermal 
K-^lbagreen), and on the fin and cephalic spines. Smooth 
P^ippear to be very rare in the PaI;eoi:oic, and it is not till 
{bh the Upper Trias and Lias that spines completely covered 
{Booth (pnoine are met vrith. 

Begard to the teeth, Mr Woodward observes that "pointed 
■kd obtuse teeth occur among the earliest Elasmobranchs ; 
I fonDcr as well as the btter arc firmly anicuUtcd together, 
fast always ha%-c formed part of a dentition in which several 
Were functional. Though the teeth of Cladodus and Difi/o- 
)huratamthus\ are as sharply pointed as those of most recent 
^ the piercing crown is placed upon a broad HoriAonially- 

Ebose, permitting of a ronsiderable amount of interlock- 
cn one tooth and another — an arrcingenient most nearly 
in the 8U^^•ivi^g CHtamydoseleukt. It !!> evicltvit, indeed, 

I ihc modern types of dentition, in which not more than one 
RACries of teeth are siinulianeuusly functional, are highly specf- 

ioru of this primitive arrangement ; and the change 
[from the deepening and bteral compression of the root of 
ih, rendering its base of support less fixed, an<l often not 
ing its coming into use until after attaining the summit or 

II the outer Mde of the jaw-curUlage. 

pth r^ard to the dispasilion of the iccth in the mouth as a 
Itfae modem Rays — most Stylliiia and Ch/amyJose/ack^ — may 
ked upon as retaining the most primitive arrangement. In 
fedaceous Sharks there has been a tendency towards the rela- 
nlargemcnt of the prehensile teeth upon the symphysis ; while 


in the Cestraciont Sharks the symphysial teeth have become 
though prehensile, and the lateral teeth well adapted for trituiatioi»— 
The former arrangement is particularly characteristic of moden^v 
times ; the latter, it is interesting to note, attained its maximum o^~ 
specialisation so long ago as the Carboniferous period. In mu^ 
early Carboniferous genera the series of lateral crushing-teeth began 
in part to fuse into continuous plates {Pleuropiax) ; two of tboe 
plates often amalgamated {Padlodus) ; and in the most special- 
ised of these Cochliodonts {e.g., Deltoptychius), all traces of tbe 
boundaries of the original components of the dental plates becune 

Suborder i. Ichthyotomi. — This name was proposed by Pro- 
fessor Cope for a group of primitive Elasmobranchs, ranging frran 
the Devonian to the Permian, but perhaps also surviving to die 
Trias, and showing the following characteristic features. The endo- 
skeleton has granular calcifications extending equally throughout 
the cartilage ; the notochord in most, or all, cases is not constricted 
to form distinct vertebras ; and the calcification of its sheath in the 
precaudal region does not extend beyond that very incomplete stage 
to which the term rkachitomous has been applied — the explanatiffl) 
of which is given below under the head of the Labyrinthodont Am- 
phibians. The neural and hsemal spines of the vertebrae are long 
and slender, and no intercalary cartilages are developed. Finally, 
the pectoral fins have a long segmented axis of the archipter^^ial 
t)-pe (fig. 850). 

It may be mentioned here that Dr Koken is indisposed to admit the 
right of the Ichthyotomi to form a group of equal rank, with that un- 
bracing all other Elasmobranchs ; since he regards the primitive fca- 
lures exhibited by the vertebral column, and the nature of the caudal fin, 
as only one degree removed from those found in certain SelachiL This 
writer, indeed, regards the Pleuracanthida and Cladodontida as so 
closely allied to the Notidattida and Cestraetontida {Hybodontida) that 
he would class the whole of these families in a single group, for which 
he proposes the name Proselaehii, and in which the Cockiiadtmtida 
should perhaps also be included. So far as regards the slight import- 
ance from a class ificatory point of view of the imperfect calcification of 
the vertebral column, Dr Koken's views are in harmony with those 
adopted below in the classification of the Labyrinthodont Amphibia. 

Family pLEURACANXHiOiB. — In this family the body is slender 
and somewhat depressed ; the mouth differs from that of all the 
Selachii in being terminal ; while the caudal fin is diphycercal. 
There is a long and low continuous dorsal fin ; while the pectoral 
lin has a biserial arrangement of rays somewhat after the fashion of 
Ceratodus. The type genus Pkuracanthus has received an almost 
bewildering number of names, of which it will suffice -to mention 
Diplodus, Ortkacanihus, Xenacanthus, and Didymodus ; some of 



: -'■«, B tieir ttrtninations indicate, having been applied to ^ncs, 
nuj olfKn to Weth. 

Scceal dt»ovenc3 hare enxbtcd us to .illain to a nearly complete 
koowtedjc of the anatomY a( thi» icnuirlublc ^'cnus, and a restoration by 
M. Brm^nian of one of the spctica w ihown in the iicfomjianying wood- 
I at. Tkt ikxa was quite naked ; lite body elongate, and the snout 
lobiBK; Tbc tccih oavc a thick and dcprc&scd root, with a crovt-n 
Iwncd hy two URec|u<il comers diverging like a V, with a small denticle 
aifacbaicDf [be two, and not unfrcqiicntly a minute flaiicncdmnmniilla 
tfmtamiy. In the male the pelvic fins carry a robust "cliispcr." At the 
npflfllKbcad tbeiewait a Utkc Ijarbed spine (fig. Sjz, i). with a doutiic 
n««f)ernuioa5,aad,«cconlini;to ibc restoration (fih-. Sjo), aupportini; a 


t^t^—Ittauntioa al ih« iltlMen «f Pfnr^aiMiu C^tJiyi: ham ib« CkrtaaitMOW 

tf Pr»a«e. wtluocJ. <At«r Prvogiikji. ] 

Uc fin. In the ik\M, according to I> Knben, there was i tlistmct 
dtbular, but the pnUtouien-gmd \nt had a direct connection 
iriw poctorbital pmcesfc of ine cranium, as in Notidatius. Di Kokcn 
! iHjffady modily the structure of the pecioml (in from that given in 
^.<5a Tbedoml fin is of great length, extending backwards iis far as 
IM uphyeercal caudal, fiom which it is scpantcd l)y a deep milth. 
Anaraio]; to M. Brongnian's restoration the anal lin was double, 
and iis two divitions had a strwiure curiously like thai of limha ; 
Dr Koken eonxiders, howc^'er, that rhe restoration is incorreei in this 
puocnlAT. Specimens of the figured species attain a length of mor« Ihan 

In ttne this genus extettds from the Carbon iferous to the I^jwer 
Pereiian; while in space its range embraces botK Europe and 
North AmericsL Chondrtmhtip, from the Ijjwci Carlnoniferoiis of 
DunlneMilure, which is provisionally lefcrrcd to the same family, has 
no ccpbalic spine. Deiadied teeth from the Kcupcr of Sonicisel, 
described under the name of Dipledus, apparently indicnte the RUr- 
of a foTTO ajlied to Plturacantkus in the Triassic period. 
F\uii.v Ci-vDOi>ONTiDJ». — ^Thc second family of this suborder is 
■ impetfectl)^ known to admit of definition ; but it appears that in 
: type genus the pectoral fin had only one series of rays, and was 
imermediaie between that of Pleuracanthus and the fins of 
; ScbchJL The type genus Cladodm had a broad and depressed 


head, with the teeth arranged in numerous rows. The cnw^f 
of these teeth present some resemblance to those of the Seladi*^*' 
Hybodus (fig. 865) ; consisting of one large cone, flanked on tlAi^^ 
side by one or more smaller cones, of which the outennoit il^ 
generally the largest This genus is exclusively Carboniferous, nf 
occurs both in Europe and North America ; a large number flT 
species being known. Dicentrodus, of the Scottish Carbonificni^ 
is distinguished by the teeth having only a single lateral cone 
developed on one side. Phtxbodus, of the Devonian of Iowa, B 
an allied genus, with the lateral cones of the teeth at least as bqe 
as the middle cone ; while in Lambdodus, of the North Amoioi 
Palaeozoic, these lateral cones are totally wanting. In Dknttohi, 
of the Carboniferous of both Europe and North America, we hne 
a modification of the Cladodont tooth, in which the central cone b * 
compressed, with the cutting-edges serrated ; while the lateral cooei 
may either be two in number, or absent. Finally, the Noift 
American PalEeozoic genus Hypocladodus differs from the preceding 
by the absence of serrations on the edges of the central cone ; fl* 
lateral cones being invariably absent. 

Suborder 2. Selachii. — The existing Sharks, Rays, and tbot 
allies are characterised by the endoskeleton being, as a genenl 
rule, only superficially calcified ; while, except in some of the eaiiid 
forms, the notochord is constricted at the centre of each vert^Ht 
The neural and hsemal arches of the vertebrae are short and stout ; 
and intercalary cartilages are nearly always developed. The pectonl 
fin (fig. 846) has not a segmented axis, being of the ichthyopterygial 
type ; and the axis of the pelvic fin of the male is produced into the 
so-called "clasper," which is connected with reproduction. Some 
of the other features of this suborder have been already mentioned 
at the commencement of this chapter. The Selachians may be 
divided into two sections, according to the structure of the vertebral 
centra. In the one the anal fin disappears, and there is a tendency 
to the depression of the body and the enlargement of the pectoral 
fins, but there is no diminution in the size of the spiracle, and the 
vertebne, when fully developed, are of the type known as Uetospon- 
dylic. In the other section the anal fin persists ; the shape of the 
body is always rounded — not even excessively flattened in the 
Sciiiiida; the spiracle tends to abort, and may be almost or totally 
absent in the most specialised forms ; while the fully developed 
vertebrae are of the type known as asterospondylie. The type of 
vertebral centra known as (ydospondytic occurs in the immature 
condition of both sections. 

Section a. Tectospondvli. — In this section the vertebral centra, 
when fully calcified, have the concentric laminae predominating over 
those that radiate from the centre. The anal fin is invariably want- 



>n shows itseir in the depression of the body 
f tbe pectoral fins — the spiracles being always 
I This Ecction includes the Spiny L)og-li»he», Saw-Rsheit, 
iji, uid Rnys. 

tv £i>ixAcio.«. — In this family, which includes the exist- 
ij Dc|g-fishea (Ag. 849), we have generalined forms, with the 
« oc less rounded, and but slightly depressed. The leeih 
ted ; the pectoral 6ns are devoid of .1 notch ai their root, 

not ejtpanded anteriorly; while the gjlls are small nnd 
ind the spiracles large. One fossil species referred to the 
Meditenanean genus C^n/riiia has been recorded from the 

of Italy ; but this determination is not absolutely certain. 
u (6g. 849X of whieh two species are found at the prL-scnt 
Ktrmpenkte seas of both hemispheres, occurs in the Chalk 
ebonon, and also in ihe Miocene of Wurtenit>erg. Another 
n from the Lebanon has been referred to tbe existing genua 
hwj, but it may pethaps belong to AeanfAiiis. The exist- 
IS Spinas hii& been recorded from the Italian Pliocene. 
■mnus we come to .mother exisling genus, differing from all 
it precede by the absence of fin-spines. It occurs fossil in 
lene of Italy, which has also yielded remains referred to the 

Y pETAtxiDO!fT!n*_ — The Pclalodonts form a family ex- 
Carboniferous, presenting the following characters. The 

( somcwhai depressed, while the pectoral fins were large, 

luced forwards in the direction of the head after the manner 

Rayi. The leetli (figs. 

>) fanned a close pave- 

the mouth, and are com- 

(rom before backwards, 
crown raorc or less bent 

fe, and cither blunt and 

ir with a rutung<dgc, the 

ng often large. In the 

^afossa {Ciimaxodus or 

u), vfaich is common to 

ni>d North America, the 

t- 851) are so thictcned 

tct^ that the complete 

rms an almost entirety [ritiirniing Burtice. These teeth are 

in three chief rows, as in the figure, which gradually 

in size anteriorly, and are flanked by one or more smaller 

less thickened teeth. The body is covered with fine sha- 

Thc Nonh Ameri«m fnsodus has the margin of the crowns 

ttbdcft into two 01 three jwints ; while in PetaJorhywHus, 

yit- tsi ,—Potlttiur Mp4CI oltome rf the 
ttx Kngliih Cai' 

(cniraJ Wiih of Thihum limtf^/rrmh; (rain 



which is found both in Europe and North America, the teeth 3f^^ 
the same general type, but have more compressed crowns, withf 
undivided roals. Again, in fetaicdns, which has the sanw distribuD 
AS the prcccdint;, the teeth (fig. 852, 4) arc much elongated 
verscly, and compressed from before backwards — the crovm b«n 
petal-shaped, with a smooth or slightly crcnulated cuuing^ge. V)ff 
nature of the artangement of the emire scries is, however, unknomL 
The teeth of Oettspty<hiHi (fig. 851, 6) arc distinguished by the 
coarser denticulation of their tmting-cdgc — those of OtUofrisitia 

Tif. Is«— Si-iiin And mill of CtrtAnifcruiu fc-lwHnolifsncJiei. i, Nuehul ■pi"* *• Pitm'»rtt- 
iMmiirrUumiu; i, Flniiiine or6>rsirii>fit«); ). Ilex uf JjMraici tmtkiul 4, Tomh of/VMAifia 
««Miiuru. itva item lh< Ude ; j, Di>. of PuaimidM, mob fran aluH ; 6, Do. oT Cttm/ftf 
[Mm. 1-j ore rediKstl. 

differing from the latter by (he absence of etumel^blds at the tnse 
of the crown, and also by ihe nature of the root I( is probable 
that both these genera occur in North .America as well as in Europe. 
In Polyrhizcdus {Datiyiodits), of both Europe and North America, 
the teeth are extremely stout, with low crowns, usually having a 
sharp cuLting-cdj;c devoid of crcnulations, and a large root diridcd 
into a number of rootlets. Other allied genera from Europe are 
known 05 Gh$sodus and Maohpfiodm. 

Family rRisTODONTto.«. — This family is only known by the 
genus Pristodui, of the European Carboniferous, in which the crown 
of each tooth is thin, plate-like, and symmetrical, with hollows corre- 
sponding to elevations in the opposing tooth of the opposite jaw. It 
has hccn .■suggested that cich jaw carried only a single tooth. 

Kamilv Squatikid*. — With this family we come to a group rep- 
resented at tlic present day by the Monklish, of whicli the skele- 




if \he fiKtoiai fin is shown in fig. 846 (p. 930). The body is 

Ktd, mih the mouth pL-iced anti-riorly ; while iJif iwcloiral 

M scdt pnxluccd aatcriorly, although unconnected with 

Bad. Tbe teeth are conical 

Vattd ; and the dorsal fins are 

rqiine;aod pbcedon tbe tail 

lly koovn genus is Sfttatina 

\ which ranges from the 

1^ Upwards to the present 

hfaough lotne of the fossil 

hjre been described under 

umes, siKh as TAaumas. 

lUy pttscrvcd skeletons occur 

Lower ICimendgian litho- 

bmestones of Bavaria, one 
I a shown in the accompany- 
idcnu Other species hive 
lined from the Chalk of the 
\ and of England, and also 
c Mioctne and Pliocene of 
tinent ; while detached teeth 
id in tbe English Gault, the 

Clay, and the Red Crag. 
uv PttisntML — In the true 
ts the bod)- is scarcely de- 
tbc pectoral fin& ate not 
[panded, and the gill-slits are 
nferiorly. The most charac- 
feature 1$, however, the so- 
'saw," which is a long and 
cified prolongation of the 
iTiDca wnn a sCTies 01 large ofi-,„„'„,_^..,„,^. f,„m ih= Hiimnij. 

teeth on either border. Re- «'*" "' b«™u, i«uii»;rj. nmimi >«*. 

, , . , n. Muidibia: 0, Pcclntal ijinll* ; t, Pvc- 

H the single existing genus lomlUn:.^, I>e1vi>,;<, PdvUfin. 

KCttr in the Middle Eocene 

m and Braclclesham ; in the Middle and Lower Eocene and 
C of the Continent ; and the Eocene and Upper Cretaceous 
h America. PropriHh, from the Kocene of Kgypt, is said 
by the absence of calcificacioiv in that part of the "saw" 
tupporU the teeth ; Am^ypn'ilis^ of the same bcJs, has 
and broader teeth than Pritth : while Sc/ervrAymrAus, which 
long to the Pristiopharida, differs in the structure of the 
and the small site of the rostral teeth. The latter family, 
tas bteral gill slits, may be represented in the Miocene of 
by a species of tlie existing Prishcphorut. 



ULT RHn(OBATiD.f: — Wiih this ramily we come to Fishes lo 
Ac gtnml term Raj"« may be applied; all of them are 
lerised !>>■ their extremely ilcpressed bodies and ihc great de- 
lent of the pectoral firvs, so ihat the body \>iopvT wiih its fins 
ed ihc "disk." The teeth always form a. Virtl of pavement 
i^' SS?)- In the prvsenl family the tail is long and stout, 
D wrll-dcvdoped dorsal fin& ; while there is also a caudal fin, 
br^iudinal fold on either side. The disk in not excessively 
led; the rayed portion of ihc pectoral fins noL extending to 
BL The ij-pe-genua /ihiiitfbatis {SpatAodafis) is represenied 
pnsent day by about a do£en specict, whreh inhabit the 
seas, and attain huge dimensions. The snout is produced 
pog rustrum, wbkh is conncctoJ with the pectoial fin by a 
Bums expansion (fig. 854). The teeth arc obtuse ; and the 
Ens arc devoid of spines. In time this genus ranges from 
peridgian to the present day ; species being recorded from 
pgiaphic limestones of Bavaria (Ag. S54) and France ; from 
rtlamtian of France ; the Chalk of the Ixlianon and Italy ; 
Idle Eocene of Monte Bolca ; the Miocene of U'urteml>erg ; 
rr localities. The Austral* 
fMfirhtma, which differs in 
tcture of the nasal valves, 
10 occur in the Middle 
<of Italy. Finally, the gen- 
vmnodatii ant! Anerodrrmus^ 
Wcly from the Kimendgian 
ice and Bavaria, appear to 
I the present with the foi- 

|LY Raiip^ — In the true 
>T Skates (fig. 85s) the 
i broad and rhomboidal, 
■ally has dennal rugosities ; 
be rayed ponions of the 
I &RS extend to the com- 
tly short »iout. The tail 
ibortcr than in the RAino- 
, with a longitudinal fold ; 
» development of the median 
ihjcct to considerable varia- 

Thc type genus Xaia (Ae/iaa^/is) has ihc tail veri,- distinct 
Je body (fig. 855); the pectoral fins not reaching to the 
ity of the srtoot ; two dorsals, and either a rudimental or no 
fin. The dentition (figx. 856, 857) often varies gre-aily in the 
some Of all of the teeth in the male being sharp, while 

7« iV 




fig. Ssi,— Til* Hmiiiii Skate iJlaia maf 
('w4hi)i nor ^ «i-rlfa luiunl Usv^ Rvcvntn 
IfllUii. (AJiti GmK.) 



all those of the female are obtuse. Some of these Skates me 
upwards of seven feet across the disk. In a fossil state this 
occurs in the Chalk of the Lebanon, and the Upper Eocene 
Hampshire; and also in the Suffolk Crags and Italian Flio. 
where it is represented by the living R. clavaia (figs. 856, 8SfJ^ 
The extinct Dynatobatis, from the Tertiary of South America, is dil^ 
tinguished by the enormously expanded bases and the small : 

Fig. 856.— Froal vi«»af lhejawsorihcnulcThorab>ck Skue <AaM cmbwXbX lUdaoi 

of the dermal tubercles with which the body is studded. AcomSi*- 
baiis, from the Middle Miocene of France and Wiirtemberg, has t>B 
dermal tubercles, with small bases, which often fuse together into 

t'ig. 857. — Front view of the jawi ol the female Tbombadt Skaie (A'o" ctmalm^. Radnotd 

groups ; while Onco&atis of the Pliocene of Idaho, which has these 
tubercles of a pentagonal form, may be identical with the type 

Family Torpedinid*. — The Electric Rays have the disk broad 
and smooth ; the rayed portion of the pectoral fins not continued 
beyond the base of the snout, and the median fins well devel(q)ed. 
The peculiar electric organ is placed between the head and the 
pectoral fins.' Extinct species of the existing genus Torpedo occur 
in the Middle Eocene of Monte Bolca, near Verona. 

Family Psammodontid^. — This extinct family is known only by 
portions of the dentition, so that its definition is at present impos- 
iiible. It appears, however, from the parallelism of the mandibular 
rami that the body must have been depressed like that of the Rays. 
The teeth (fig. 852, 5) are of a flattened quadrangular form, with 
the root nearly as large as the crown, and were arranged in one or 
more longitudinal rows, which were arched antero-posteriorly with 



of the lectli of adjacent rows. The genus Co^fdus 
inancs] teeth, with the jKutero-bteral angle-!! of chc rout, 
Mimttako of the crown, (icoduoed tNtclcvrards ; the tcvth 
■oowot in Troni, with the aiiteriur irdTgrn usually eiiher 
croonvex. The crown surface when unwom U tu^o^c, aiid 
bj- a more oi less iratisverse line, which sometimes jierniits 
Ipoftions to be separated. This genus occurs in the Car- 
ta (if Europe. In the typical Psammodus ihe [eeth (fig. 
ire quadiangular, generally more or less oijlong, but occa- 
leariy square, with ihc root much thicker than the crown. 
eb it is cisity separate ; the surface of the crown b«int; 
marked by tnuisvene wrinkles. It is probable that ihc 
e arranj^ in the jaw-s in four ion^tudinal rows. Psam- 
represented liy a considerable number of spedcs from the 
Irous of both Europe and North Americx Lastly, Ardus- 
Va the CarlxmiferouK of Indiana, 15 an allied fnrm, with 
I pawment-Iike teeth arranged in several rows, of which 
I ntrbces arc somewhat excavated lo fit tbc curvature of 

r MvLioBArii>,E- — Willi the Eagic-rays wc come to an exist- 
r well represented in a fo^il state as far donii a-s the I^wer 
vcitcbne frotn Crctaccoux auid Jurassic beds having been 
*red to thi& family. The di»k is very large, owing 10 the 
dopmcnt of the pectoral fuis, which stop short at the •i\^<ii 
sad, but reappear at the extiemity of 
t in the form of a small single or 
tpholic (in. The tail is extremely 
u>d KKmbles a whip-lash ; and the 
, wbcQ present, forms a complete 
t The tNTW genus Myliobatis has 
I liCc from the di»k, and a single 
fin. The lecth are L-u^e, flat, and 
I, and arc arranged in seven loiigiiu- 
^; tbc middle row in the adult being 
) broad, while the lateral tows are 
Hpcd (fig. 85S). In the young the 
DW of teeth i^ not larger than the 
es, and there is a gradual increase in 
re breadth of this row as the lish in- 
) age. The upper dental plate is cx- 
onTcx from before backwards, but the 
t is quite flat In addition to the 
wrrtebrx mentioned atK^e, thi.s genus is known continu- 
D the Lower Eocene upwards, and has a wide distribution in 
in the Eocene, it is recorded from Europe, India, and 

Li>w<f <J<'jii*I jiU*' '">i -Wr- 

MiiMk X'-'-.^vt of T^racl(I#- 
OujiL. >u>MCii, Kii!utcr!t' 



North America ; and it has also been dcscrihcH from ihc Teni. 
lew Zeaknd A very largu numbvr of specific names have 
ilied to [he fos»i) forms, but Mr S. Woodwatd has shown that 
number of valid ijiecies may he greatly reduced, since many of 
characters on which tlioy w«te founded are solely due to difl'crenoe* i(^ 
the ago of thr spt^cinien*. In RhittupttTH {Xy^^hatrs), while the \ja& 
is still free, there arc two cephalic fins, ai]d the teeth arc arranged ffl 
fiv« or more rows, of which the middle series and the adjacent j/k 
arc broad, while the one or two pairs on the borders fonn regilH 
hexagons. This genus is know-n from tlif London Claj-, the Eoccm 
of South Carolina, the Swiss Miocene, and the English Crag, audit 
now rvprescnied by seven species from tropical and suh-iropical «u. 
The existing genus Aitobalis \a distinguiirhed from the preceding 
by the teeth being arranged in only a single longitudinal row, whidi 
is often bent, and corresponds to the median row of MyJiobaHi. 
One species is found from the London Clay to the higher Eocene 
of Barton, and the genu* is also represented in the Swiss Ktiocene. 
The existing genus Ctraloptcra has no upper teeth. 

In this family may In- provisionally included the Cretaceous genu 
PtyiAodui, which was long considered to he a Cestraciont Shirt, 
but lias been shown by Mr Smith-Wood- 
ward to be a Ray allied to Afyiiohatit. 
']"he connection between Piychodus and 
the existing A/yh'oim/idic is shown by the 
geniis Apocopedim, from the Upper Cre- 
taceous of Braxi^ which has teeth of an 
intermediate type ; whili.' some of die 
Eocene species of Myliobatu have teeth 
of nearly the same form as those of 
Ptyihodus. The teeth of this genua (%. 
859) have quadrangular crowns, with the 
enamel of the central region tfannm 
into a number of transverse folds, while 
the root is smaUer and lower than the 
crown. The two sides of each jaw are 
parallel to one another, and the Icclh *« 
arranged in several parallel rows running 
from hack to front, as is shown in the 
accom|Mnying diagram (fig. 8(io). It will be 
lieen, moreover, that each jaw has a sirtgle 
median aeriea, composed of very small teeth in the upper, and 
of very large ones in the lower jaw ; while on either side of this 
median row there is a series of iccth somewhat )e» large than 
the median row of the lower jaw. The five external rows gradually 
decrease in size towards the outer !»ide of tlie jaws. A com 


ntwi of in* cra*n ot a tooih of 

Upper lincnMUiil of Kevcin- 
bvJv- (.Uler ZtlKl.) 



4^ ihf< (wvh in the u|njer (4} mi J ivv«r (11) 
am at fIjtJuiliu ittMrinu. (Aflcr S. 

rob % Ij* will show that the general plan of this arrangement 
rflierctth is essenlialljr the same a& in Atyiiob^is. This genu* 
All J wMle dKtribuiion, being 
kmA \a the Upper Cretaceous 
(ri" Enropcv India, and North 
Famiit Tkygosid^. — In this 
jFv t)i« pectoral Ans continue 
I the «trrmitjr of the mtuzie, with 
rlich ihejr ate confluent. 'J'hc 
i> slender, and sharply di«tn> 
from the disk ; while the 
cicil lint are imperfeci or at>- 
~MR, and may be replaced by 
Ktr ated sptncs. Tliere U some 
noemmtf j>s to the occunence of 

Tfygam in a fossil suttc, l>ut it is probable that either this genus or 
tfce iIU«d Tamura date frum the Middle Eocene of Monte Boica, 
•fld tniuint rcrcrreil to the fornKt have been dcacribLt] from the 
L'pper Cretaceous of Newr Zealand. The extinct Xtphatrygmt, from 
the Eocene of Wyoming, is distinguished by iu cuspidate teeth. 
Renuins referable to the existing tropical genus Urolopkus occur in 
the Uiiidle Eocene of luly, and perhaps in that of Helgiuni ; while 
CydcSatis is an extinct genus from the Cretaceous of the Lebanon. 

Section B. Aaterospondvu. — This section is di-tlinguished from 
the but bf having the ndiaiir>g lamins predominating over ihe con> 
centric ones in the fully calcified vertebral centra, io that a seaion 
Aaw% a star-like arrangement. The anal fin is always present : 
■bile specialisanon docs not tend to a flattening of the body, or to 
El eipansioa and forward growth of the pectoral fins, and the spir- 
adca are small, and may disappear. This section comprises the 
XxvK Sturks arkd the Dog-fishes. 

An these (uhes have clon^lcd and subcylindrical bodies, and a 
Hraoe tail, well adapted f»r sirimminx- The untcriur, and vco' fre- 
qaenny all the lecth are formed oit die type of a laterally comprc^&cd 
coae with trenchant cdccs, at the base of which iwo or more minor 
cooea nuy be dcvclopca. In many cases, however, ihc hinder (ccth 
have obtuse cronns adapted for crushing. The two nimi itf the jiiwn 
arc never pamtlcl to one antrther, in ctmsequence of which the iccih 
e a]va>^ set in nbliuue ri>WA (fiK- 864^, and never fonn the straight 
lCTO-p04teriof rows. like ihoM occurrini; m the Kftys(%, 860V iiharlis 
)i«nroD»,and of active pelagic habits ; and arc mnst niimcn)us 
" *ea», alihoujjli ihcy ascend tidal rivers, and arc even found in 
id lake in the Fiji Islands. 



there is but one dorsal fin, and the number of the gill-ctefts eiceedi 

Family Notidanid^ — The single family of this series is repn- 
sented by the genera Notidanus and C/Uamydoseiache, both of whidi 
inhabit the wanner seas. The dorsal fin has no spine, and the VexA, 
of which several series are in use at the same time, have shaipif 
pointed cusps. Notidanus {Hepianchus, Hexanckus) is readi^ 

^f Orb 


Fig. S6i. — Left laural view of the ikull at Ntiidama. Recent. Reduced. R, RoMiaa: 
AF, PF, Pre- and poiiarbiial proceuei ; Orb, Oibii; NK, Nunl Canute; +. AnioUatiM d 
palatopterygoid i.Pg\\ t, AnicuUlioo of Meckel's c»itil^ {Md)\ Z, Teeth; If' .T, Vcnebnl 
column. (After Wiedersheim.) 

characterised by the inferior position of the mouth and the pecuKar 
form of the lateral teeth. These lateral teeth (figs. 86i, 863) ue 
comb-like, consisting of a series of compressed cones, inclined in 
one direction, and fixed upon an elongated base ; the anterior cone 
l>eing the largest, and frequently having cusps at the base of its 

anterior border. The lower teeth 
are more complex than the upper, 
and the anterior teeth in both ja«s 
are simple and awl -shaped. The 
primitive position of this genus is 
shown by the persistent notochord 
and the characters of the skull 
(fig. 861). The latter approaches 
the so-called amphistylic type, differ- 
ing from that of all other existing 
Selachians by the slendemess of the 
upper part of the hyoid arch, which 
does not support the palatopterygoid 
bar and mandible. The palatoptery- 
goid is connected, however, with the cranium by a distinct fircet 
articulating with the postorbital process. This structure is re- 


V\%. S63, —Lower leeih of NetidoMut 
gigai ; rrom Ihe Red Crag. The an- 
rerior cu«ps are worn away in the lower 
figuR. (After Smith- Wood ward.) 



by Mr SmUh-Woodward as but veiy slightly iemo\-«! 
tbe otiginal primiiivc condition ; Catradon, in which the 
biilar becomes tUstinctly differentiated, lx:iiig a stcj) in 
iTaim. In time this gcnu» is definitely known to range 
the Middle Jura.s&ic of che Oxford Clay to the present 
tl hu indeed been recorded from the Lias, but the dctcr- 
is more than doubtful. Nearly all the described species 
Eeropean, but one has been recorded from the Teftiary of 
Zcabnd. Some of tbe fosNil torih tihow sii^ns of tvc^r at 
smnitB, And it thus seem« they must Ktve been firmly 
loicd in tbe jam h'Vc those of the Hybodunt:^; specialisation 
Iniog apparently tended to produce a loose dental articul-ition 
riBDqgfaoet the section. The genus CMamydoselaefu has a terminal 
nmdi, with Utera) teeth similar in butli jaws, and consisting of three 
ibdtT rottcs separated by smaller cusps ; the notochord beint; partly 
This genus is now known by a single living species from 
aea3, but lecth from the Pliocene of Tuscany have 
referred to it It will l>e ohsened from the figure that the 
feundibular articulation of Notidanus ia placed for hchind the 
proper, and it is twtewonhy that a similar condition ob- 
bus io fimratanihvs unong the Ichthyotomt. 

FxiuLV CocHUOimNTiDit — With this family we enter the second 
Kries of the section, in which there arc two dorsal fins and five gill- 
defts. The present extinct family is an ill-dcfincd one, apparently 
lOied to the Ctsfmei&ntida, but with a more specialised dentition, 
Tbe dentition is formed from that of the CtUradovtida {infra) by 
Ihc welding of at least ot>e of the oblique trann'erse rows of teeth 
encircling each ramus of the jaws into a continuous airvcd plate 
(fi^ 863), which may have either a smooth crown-surface, or may 
be marked, as in ttie figured example, by grooves and ridges, indi- 
catng its compound origin. These dental plates grow hy additions 
to their inrjcr borders, while the outer borders are usually involuted. 
Tbe dorsal fins were provided with spines. This family eoniprifies 
a number of genera from the Carboniferous, only sonic of which can 
be Twy briefly noticed in this work. The one most imperfectly 
ktuiwn is fleioduSi fmm the Hnglisb C3rlK)nift'rous, which appears 
allied 10 tbe ttext, but does not seem to have tiad the tt-eth welded 
pbtca, at»d therefore differs from the accepted definition of the 
ily. Pkarofiax {Pteurodus) and Fsephodus are more tyi>ical 
the latter occurring bolh in Euroiw and North America. 
dental plates of Fsefhodus form at least one scries of smooth 
cnncd teeth, without coronal ridges or involution of the outer 
bonier ; and there were also smaller rows of lateral teeth, as well as 
:ie prelteiistle teeth at tlic cxltcmities of the jaws. Sanda/^dur is 
by the huge dental plates, which arc of an elongated Irian- 


gular shape, with the outer border sHghtly involuted, and an uods- : 
lating coronal contour in the upper jaw ; the genus occurring n . 
Europe and the United States. Other genera which can be modf ■ 
mentioned are Tomodus, Xystrodus, Deltodus, and Ptedlodms ; the 
three last being common to Europe and North America. Teeth 
referred to Pmdlodus, and to the above-mentioned genus Psephsitt, 
have also been described from the Carboniferous of Northern Indik; 
and the teeth from the same beds described under the name of 

Helodepsis are likewise referable to 
the present family. In the tn" 
genus CochHoduSf of Europe and 
the United States, there are tw 
pairs of dental plates in the nun- 
dible (fig. 863), in which the ouia 
border is much involuted. The 
posterior plate is elongated antero- 
posteriorly, and has its anteiiot 
Fig. 863.— u««r dentoipuittof Cm:*/!- and posterior borders converging 

ffrfm ceMlerlui: from ihi Carbonifcroui of „„t„„,ji„ ,„j , „,„„;„„_, „l,i;„„. 

Armagh. One-taif i»tu™i .!«. Tht iped- outwardly, and a prominent oDliqoe 

men ii viewed frODi ihc anlerior upcct, uid ri,^„fi lilrpuriu' narrrvm'xrto in thr 
Ihe whire line or ihe left lideihowiThedivi. "°g^> '"'eWlSe nanOWmg m UW 

uon of ihe iwo plates. Same direction, in the middle of 

the crown. The anterior plate is 
narrow from before backwards, with a ridge on the hinder border of 
its crown resembling the median ridge of the larger plate. The 
remaining genera of this family are Streblodus^ DeltoptyJtius, Diplac- 
odus, and Cyrtonodus, of which the two last are doubtfully entitled 
to distinction ; while other names have been applied to detached 
anterior teeth. 

Family CESTRACioNxiDiE. — This family, which may be taken to 
include the Orodontidtx and Hybodontida of many writers, is repre- 
sented by a considerable number of genera ranging from the Car- 
boniferous to the present day; all the existing species being included 
in the tjqie genus Cestradon, of which the upper dentition is shown 
in fig. 864, The family is characterised by the presence of a spine 
to each dorsal fin, of which the first is placed immediately above 
the interval between the pectoral and pelvic fins. The teeth are 
generally more or less obtuse, with several series in use at the same 
time, and those of each oblique series never fused into continuous 
plates. One of the oldest genera is Orodus (more correctly Oreodus), 
of the Carboniferous of Europe and the United States, in which the 
teeth are of the general type of those oiHybodus^ and are only r^arded 
by Mr S, Woodward as entitled to separation on account of the 
absence of other associated remains which are characteristic of the 
Mesozoic genus. It is probable that some of the dorsal fin-spines 
originally described under the name of Cltnacanthus belong to 



Mom of these spjnes (fig. 852, a) are, however, rcfcrahle 
dte allied CuiJooifcrous genus Sfi^naantt/tas ; and ihcy arc 
~: by their ortumeiiution of robust longiludina] ridges, 


dts^ (AIVaO«n.> 

fcji-h are partly nodose. Ot^cr Carboniferous getxra are Campodus, 
xUst^us, and Tntfy<Aiiis, some of wliicli liavc several synonyms. 
In the Thunngian I'crmiar this family is rtprtacnled by the genus 
IWkbfai, in wbJch the teeth have large and smooch eionns vrcll 

iht LoKcr Cnunniu. Rnluctd. 

idapted for crushing; while in the Muschclkalk, or Middle Trins, 
have the imperfectly known PalaoAatis, wiih iccih wry like tliuse 
tsienuaniAtti. With the genus Jfyhodus, ranging from the Mus- 
elkalk to the Lower Grccnsand of Europe, we come lo a type 
U now almost as well known to us as existing Sharks, owing 



to the bttautiful preservAtion of uuiiy of the specimens From I 
l,ias in which deposits these fishes were nbundant. Many ofl 
numerous species Attained very large dimCMixiions ; and the 
may be dcdned by the following chaniciers. The teeth (fig. 
arc conical or cuspidate, the crown being moK or less striated, 

one main cone, flanked by one I 
more Literal cones, and the Wtj 
hL-ing more or less dciiresscd. ■ 
'I'lic tet'th at the sj-mphysis an 
Iar}cc and few in number ; while 
the jinspines {fig. S66) are lon- 
giiudinally grooved and ndgedi 
with iwo rows of denticles placed 
near the posterior borders. The most leitiaikablc feature is, how- 
ever, the presence of iwn large hook-like spines immediately behind 
each orbit, which hav*: been described under the name of Sfiitn- 
onckus. The notochord is persistent. 

Fin B*; — Toolh <i( jlcmtui natiil/i. from 
the Lavcr LJAitdf L^iiir-Rr|r^h 

SjMcimens of the I-nwcr Liassic fi. IMafift/i^i show the whole of tbc 
dentition in situ ; and it apiiKio. from tiirsc that ihere ivas no medtao 
8ymph>-sial row of nwndibdar teeih, but thai there were ten iransverW 
rows of lower teeth, with five teeth in each n»iv, while the number of rowt 
in the upper jaw was cither nine or ten. In ihe later h>m\s, such a* H. 

finsfiKus of the Wealden, ihe teeth 
differ from those of the typical 
I.iassic forms by the taller, com- 
preM(nl,and ncarlysmooih crowos, 
and Mr WniidviTird sujrgcsts thai 
it may evemiially he a(i^■I»lUe ta 
refer these tyqtes to :i dittinci 
pemis. The orhita! ho»k« found 
tn the Oxford and Kimeridge day 
indicate lishes of very lar^ di- 


The genus Atrodtis is closely 
allied lo the preceding ; but 
the teeth (fig. 867) are non- 
cuspidate and more rounded 
Tliis genus i» abundant in the 
•u. Lias, where it is represented by 
""■ the lai^c *-(. Hvf^Iis, which is the 
type ; but it also extends down- 
wards to the Mu»chelkalk, and ranges upwards as high as the Clialk. 
The majority of the speries are European, but the genus has also 
been recorded from the Cretaceous of North America, as well as 
froin certain beds in that coimtry which have been stated to be 

-Pnn ffthr |-.i 

J, Irom the III 

Hultieol. (A(WrO««n.) 



mthia {Str^kodut) agrees with Hyfxidus in the persistem 
prd and the presence of orbital spines, but differs in the 
lEt3 of the liii-fipines and (ccth (ligs. 868, 869). 'Itic prin- 

eeth fonn irregular rhomboids, with shghlly arched and 
)d crcMms, marked by a reticulate omainentation — the sym- 
tectb being large, few, and simple. 'Jhc fin-s[iincs an: 
I by uellsue tubercles, which are somelimes fused, and 
ini poKeriar nxstal series of dt.-iiticlcN 'ITie lyjic species 
68) b of great sixc, and occurs typically in the Orcat 

of the Continent and England, but ranges upwards 10 
mwfidge Cby, and thus presents a remarkable instance 
Intcncc. Thb genus is also represented in the Purbeck, 
extind genera are Palaespinax, from the Lias, and Syntfh' 
Om the English Chalk, both of which have teeth closely 
\m% those of Hyhodus ; 
I firt-fpines, at least in 
XDCT, were smooth, like 
If Ctstroiiori. The Cre- 
i genus is tbe more 
Iscd, and approximates 
nial structure to AW/- 
; tb« palatoptexygoid ar- 
ng directly with the 
Finally, the esisting 
iffti {\n which may 
forms described 
names of Gyropieu- 
DreffanefiMonu) oc- 

Europe from tbe Kim- 
of Bavaria to the 

Clay, and is now represented by four species in Uic 
Bin and Japanese seas. 'ITiis is a specialised genus difTering 
U/erxuanthus by the absence o( orbital spines, the numerous 
hU lymphysial teeth (fig. 1164), ihc smooth (in-spincs, and the 
ctftcatton of the vertebral column. 

ILV ScTLUiD*. — This i» a family of sniall extent, and most of 
embers of which are of relatively small dimensions. The 
fins have no »pir>es, and the first is placed above or behind 
hric (in ; while the teeth arc small and rii«pid.ite, generally 

series being simultaneously in use. The living furms arc 
jnlj' knowTi as iJog-tishes. Paia&seyltium, of the Lower Kim- 
Bi of Bavaria, seems to he allied to the existing GiHxlynrosloma 
y/iittm. &jtIiodus, of the KnglUh Chalk, is an imperfectly 

form with teeth like ScyUi"'", but with vertebra: approximal- 
^Xamna- SeylUum itself is represented in the Cretaceous of 

I'ic t^' — LaKnl and cial iuif«-r> of ft loath 
tit A ilfr,Kanritt4 dFa.«/i'Mt<-iu. fronib* CofSlliui 
o( llie Cuniinciit- (Alicr Ziiwl.] 



the Lebanon and in the Continental Miocene ; while allied es 
genera are FrisHurus, from the Lower Kimehdgian of Bavaria, 
Mesitia, from the Lebanon beds. Finally, the existing Chik 
Hum occurs in the Miocene of Wiirtembei^ ; while Gin^ymosi 
of which some of the existing species attain a length of 1 3 fe 
represented in the Eocene of Alabama. 

Family LAMNiDiC — This family comprises the largest ol 
Sharks, and is characterised by having the first dorsal fin ^ 
above the interval between the pectoral and pelvic fins, 

without a spine. The teeth are 
minate, and when fully adult 
solid throughout The earliest 
which has been referred to this fi 
is Carckaropsis, known by deta 
teeth from the Carboniferous of Ei 
and North America. The type { 
Zamna, in which Otodus may b 
eluded, comprises the existing 
beagles, and has large lanceolate 
(fig. 870) with basal cusps, but wii 
marginal serrations. Teeth agr 
with those of existing forms in 
general contour are found in Ei 
from the Lower Miocene (fig. 
upwards, the so-called L. acumi 
from the Chalk, belonging, how 
to the next genus. In another group of this genus, formerly ki 
as Otodus, the teeth (fig. 871) are distinguished by the great 

Fig. 870.— Toolh of Lmmita aa- 
fidiUa, (tarn Ihe Lower Miocene 
(Oligoccne) of Gcnnany. (Af\er 

Fig. S71. — ToDih of Lamna a^ftndienlttttt, 
fiom the EngUih Chalk- 

Fig. 873.— Toolh o( Ojtyriima ^ia 
fiom ihe Hungariui MioCBOC 

pression of the crown, the large basal cusps, and the shortne 
the root. Teeth of this type occur in Europe from the Ga 
the London Clay, and are also represented in the Upper Cretai 
of Southern India and New Zealand. The nearly allied but e: 
genus Oxyrhina is characterised by the still greater compressi 


ODras of the teeth (fig. 872), which are generally devoid of 
ciBpi. Il has been recorded from ihe Jurassic, and occurs 
hfilJj in the Challi of Europe, Indiii. and New Zealand, and 
» rrpnsented in ihe European Miocene. Teeth from ihe 
' Octtceous or Europe and India, differing from ihasc of 
by (he tiearlj: circular section of their crowns, have been 
to tbe existing genus Odem/aspts ; but Mr S. Woodw;trd 
that tbcy belong to a genus from (he Crctaccouit of the 
originiilly described under the preoccupied name of Hhino- 
I, but nov known as Sea^ntfrhymhus. Odontaj/di iti;e1f 
n the Eocene. Teeth of a long and slender type, from ihc 
urassic arvd l^wer Cretaceous, have been described under 

y~-T*9t^^ C^nharrim mttaM-t, bom ih* M iocm* gT M ill*. (\br Ziiigl.] 

DC of Sfhtnodus, now changed w OrlAawdus. Other teeth, 
le Comtinenial Miocene, have been referred to the existing 
Aiofetms or "Threshers." With Carehawothn wc come to 
of ertotmous size, characterised by their large, ]1al, and rcg- 
lhan(;ular teeth (fig. 873), in which the edges are scrraicd. 
Eie are no basal cusps. The one existing Mijecies attains a 
of 40 feet, and has teeth measuring a little over 3 inches 
margins, with a basal width of 1.8 inches. It occurs fossil 
liocetie of Europe. In tbe Red Crag, and also at the lx)ttoni 
teetb arc, however, found in which the corTe*pon<ling 
are 5 and 4 inches, and thus indicate enormous indi- 


viduals. Smaller teeth (fig. 873) also occur commonly in J^2 
Miocene of Malta, the Tertiary of New Zealand, and still ai a*** ^ 
forms (fig. 874) in the Lower Miocene and Eocene of the OC*^ 
tinent ; and the genus is also represented in tf** 
Pliocene of Burma. Its earliest representatif^ 
is, however, C. longidens, from beds which appetf' 
to be of the age of the Maastricht or topmo* 
Cretaceous. Small compressed and triaiignhi 
teeth, usually with serrated edges, from the Ox^ 
of Europe and India, to which the name Gmu 
CaXl^'^'^Zi has been applied, probably indicate Sharks allitd 
tut, from ihe Lower to CarcharodoH, atthougti in external contov 

Miocene of Ihe Con- , . ", i ,. . 

linenL. Reduced. they approximate to the teeth of the not 

family ; they have, however, solid crowns. The 
genus Cetorhinus {Se/acke), now represented by the huge BaskiDg 
Shark, dates from the Pliocene. 

Family Carcharhd^, — The last family we have to menticoiii 
distinguished from the preceding by the presence of a nictatiiig 
membrane to the eye, and also by the hollow crowns c^ the teedi 
It is unknown before the Upper Cretaceous, and is dominant at the 
present day. The teeth have triangular and compressed crowns, 
usually with more or less distinctly serrated edges. The genus 
Hemipristis, of the Upper Chalk and Lower Tertiary, is chaiac- 
terised by its tall lanceolate teeth, the crowns of which have botfi 
edges coarsely serrated, except at their summits. The existing 
genus Gakocerdo is first recorded from the topmost Cretaceous of 
Holland, and occurs throughout the European and American Tct- 
tiaries from the Eocene upwards ; the existing forms being known as 
" Topes." Carcharias, including the well-known Blue Shark, has 
small and generally triangular teeth, those of the upper being very 
different from those of the lower jaw. It may be divided into 
several groups from the structure of the teeth, which in some forms 
have smooth edges. It first occurs in the London Clay, and is 
thence found throughout the European Tertiary series ; it has also 
been recorded from the Egyptian Eocene, and is found in the fiesh- 
water Pliocene Siwaliks of north-western India. The strange Ham- 
mer-headed Shark, the only representative of the genus ^hyrma 
{Zygana), has teeth so closely resembling those of Carcharias that 
it is almost impossible to distinguish detached specimens. It ap- 
pears, however, that there is sufficient evidence to prove the exist- 
ence of a species of the former genus in the Miocene of Europe and 
of the United States. 

IcHTHvoDORULiTES. — In conclusion, a brief notice may be given 
of a few of the numerous genera founded upon the so-called " ich- 
thyodorulites " or spines, of which the serial position cannot at 



tbc detcnniaed, and several of which arc probably referable 
>idci. Tbc earliest of these is Onehus (fig. 875, \), 
(&e Silurian bonc-bcd of I .u<lIow, to which genus may also 
[thedemul denticalex (i^iJ., B) described under the name of 

ni Hv ' ggtit («1 0* OiU*mw liwtbtrianu, an J [ »} Utt mal p^aU* o( Tluiftm ! 
bum ibi Sllniin of Ludlu". 

ThtUgt. In the Devonian of North America, and aUo in the 
ITppa Silurian or Devonian of Hohcnna, large spines have been 
abed under the name of Miukaracanthus (fig. $76), which 
dy belonged 10 the dorsal fms, Gyratanthus is ba-^ed on 

OwkaU HtiukI liM. (Aflct Nealicrry.) 

ical Spines (fig. 853, 3), which, it has been suggested, 

_ Jong to the pectoral fins. Other Carboniferous specimens 
nt been named Lophaeanthus and Ofafanihus (England), Eu- 
tam/Jius (Russia), Xystramnthui (Morth America and India), and 
%tMma{aiiHitftus (India), 
be spines described as 
racamtkui amrtoK Icnown 

belong to the bead of 
Selachian, and not, as 

first kupfxncd, 10 the 
il of a PLicodermtc (la- 
id. It will be un- 
Ksssrjr to mention a 
mber trf other t>'pcs 
Kii varioOE deposits ; but _ 
■ must not conclude ri<. i;?.- P«n ■'*Caii<i=.ii tpin* -rf Atrf«i-»»»r: 

. , . ■ txrni 111* C»rtuQHct"ii« "f NorUi ttmava,. Onc-halr 

tfafHIt referring to the ntnud >«■ (A/i*r NewUcry.J 

oarkahic specimens (fig. 

f7) from the CarlKjniferous of North America and Australia, to 
uch the name E-ieiius has been given, These have a highly 
1 axti| bearing compressed lancet-like leeth, with serrated edges 


either on the convex or on both borders. It is suggested that tiny 
may have been carried on prominences in the caudal region. Aid 
after describing the best preserved remains of this genus, Dr New- 
berry concludes as follows : " Hence, until further li^t shall be 
thrown upon the interesting question of the homologies and functioni 
of [the remains of] Edestus, we may regard them as the post-donl 
spines of large cartilaginous fishes, of which the other puts are yet 
unknown, and may suppose that they were used for attack and it 
fence, like the spines of Trygon or Acanthurus." 

Spines of similar type from the Carboniferous of Russia, ori^ 
ally described as Edestus prot{^irata, have been subsequently nude 
the type of the genus Protopirata, although it is doubtful irtietiiff 
this term, having been first made a specific one, is admissible as x 
general one. 



CZASS PISCES—conlinued. 


Ill Crimeroioki. — ^The ChJtnxroids are marine fishet, 

by some wriiers as a sulx>rder of Elasmobmnchci ; but 

they resemble Sharks in cxtemni cuntour, in tlic presence 

"daspers" on the pelvic fina of the m.ile, nnd in the structure 

I CB-capsules, yet ihcy present such important differences as 

' the propriety of referring them to a distinct order. 'I'he 

b entirely canib^inous, And the vertebral column only 

iB|iafeiJl]]r segmented; the notochord lieing surrounded by n series 

«f GBtilagtnous rings, which may be partly calcified. The skin of 

ibe lyptol forms i% ui^ually quite naked in the adult, but in the 

yOKig there is a row of sniall dermal ossiFicittuns on the tiack. 

Tht skull is movably articulated to the vertebral eolinnn, and has 

Ac hyomaodibuUr fused with the palalopter>-goid bar, and the latter 

inly united to the cranium, with which the mandible conse(]ucntly 

MicaUtcs without the intcn-cntion of a separate suspcniionum — 

iIm xrrangement being tenned autostylic. The gill-clefts are four in 

DBuber, and protected by a fold of ikin containing a cariilaginouh 

^l-cover ; their communication with the exterior being effected by 

> angle aperture. The mouth is always terminal; and in the recent 

fiwiDS each jaw carries one pair of molariform teeth, roipectively 

uuched to the palotopterygoid and Meckel's cartilage (mandible), 

with the addition of a smaller anterior pair of vonterinc rutiing-icelb 

in the tipper jaw — all these teeth i>ersi.«ting throughout life. The 

bu arc similar in structure and position to those of the Sharks -. the 

fint dorsal always carrying a strong spine, which articulates with 

the neural spines of the vertebrae, and is thus susceptible of niolioti. 

In the absence of a swim-bladder the Chimeroids, again, agree with 

ifae Ebsmohranchx. There is a laicral line stren^thetied by carti- 

lings. From the absence of any membrane bones, the 


massive teeth, which are strictly comparable to those of the 
constitute the whole of the solid part of the jaws. 

This order may be regarded as in some respects connectittgl 
Elasmobranchs with the Dipnoi — the autostylic cranium and 
dentition being essentially Dipnoid. ChimEeroids have existed I 
the Lias upwards, and not improbably date from the Devoma| 
while, as is usually the case, some of the extinct genera show : 
more generalised affinities than their existing representatives. 

Family Squaloraiida. — The extinct genus Squahraia waskii|j 
regarded as an Elasmobranch of somewhat uncertain affinities; Mj 
according to the observations of Dr Traquair, it should find a pbee : 
in the present order. In this genus, which is confined to the IJi% 
there is an elongated body ; while the skull is produced into a long 
flat rostrum, and carries a basal pair of teeth separated at the sym- 
physis, in advance of which are two small vomerine teeth of the 
normal Chimxroid type. Further, the skull of the male has a pre- 
hensile spine on the upper part of the snout, resembling in structure 
that found in Ischyodus. The " lateral line " agrees with that of 
other Chimseroids in being open, and protected by cartilaginous 
rings ; while the skin appears to have been entirely naked. The 
vertebrae are of the Tectospondylic type of those of the Rays. The 
skull has been described as hyostylic, but Dr Traquair considen 
that this is due to crushing, and that it is really of the autostylic 

Family ChiM/Erid£. — Nearly all the remaining forms, from the 
Lias upwards, may apparently be included in this family, which ii 
now represented by Chimara and Callorkynchus. The teeth aze of 
enormous size, those of opposite sides meeting in a median sym- 
physis ; and each tooth has one or more triturating ridges, w pro- 
minences, difTering in appearance from the rest of the tooth, whidi 
may be conveniently termed tritors. The type genus Chimara has 
the teeth adapted for cutting ; those of the mandible being thin and 
plate-like, with one large median tritor, and two tritors near the an- 
terior extremity, and an outer series in the form of dots ; while the 
palatal tooth varies considerably in shape. This genus is repre- 
sented at the present day by three species, and has also been re- 
corded from the Pliocene of Italy and the Miocene of Bavaria. In 
Elasmodectes [Elasmognathus) of the English Chalk, the mandibular 
teeth are likewise of a cutting type, but without the median tritor. 
The extinct Ischyodus, which in England ranges from the Upper 
Jurassic to the Chalk, but has also been recorded from the Eocene 
of North America and the Cretaceous of New Zealand, appears to 
connect Chimara with the next genus ; the teeth being more adapted 
for crushing. The mandibular teeth are, indeed, more massive, 
and generally have two well-marked tritors externally to the large 



95 » 

one; while Uierc is a spine on ihe rosinim. Edaphodm, 
I laqgcs from the Lower Crcensand to the Middle Eocene of 
bam, like ibc last genus, attains gi^cantic dimensions, but 
I teetb adapted tintirely for crushing, 'J'hc mandibular tooth 
:8;8)isTery massive, and has its sj-mphysial surface (which in 
' pnceding genera is narrow and grooved) very wide and quite 
irhile there are two outer artd one median triiors, ■.%% well .is a 
ntnal tritor which is not shown in the figure. Each pahial looih 
1 Anwhcd with three triiori. TCTrth of this genus are common in 
Oetnjceous and Tcniary deposits. Elastnodus^ which is 
in ibc l.owcr and Middle Eocene of England, appcan to he 

nt- tii.— lNBBiaiC>asri)>cn[ti>nBti<iibiitkrtwikor£^/tm(mjr^Wf,- TriMD lb* Englith 
OaIL. R*iIuc*iI. (Aftn Ecmon.) 

allied to lh« preceding genus ; while in the existing Cnllorhynehus 
we apparently have the most specialised representative of the group 
with entsbitig teeth. The latter genus is represented by an existing 
sfiectes in the Southern sea^ and by a fossil one from the Ixiwer 
Crecacecus of New Zealand. The mandibular tooth is inaN:>ive, 
with a narrow symphysis, and only a single tritor, representing the 
large median one of Edaphothn. Other European forms are Can- 
uims^ frDtn the Lower Jurassic of Stoncitfield, in ultich the tritors of 
the mandibular tooth are condueni ; and Myrttuanthus i,Pri'^aSk- 
tdu « Mttp/acanikut), of the l.ow« Lias of Dorsetshire, which is 
known both by the spines and the teeth. Its piemaxill^iry tceili ;ite 
chisel-like in shape. Le^atanlhm of the Lias, and Difristit of tlie 
Middle ^[iocene, are imperfectly known forms probaljly referable to 
tki» fiunily. In North .\mcrica the names Evuiyiodus, Lepti^myius, 
Byaaitais, Dtphrissa, luttitnitt, and ^^kagtopxa have been applied 
to CrOceous forma, while a Miocene typt- has been called Myh- 
gmmUtMS : but there is considerable doubl whether all these forms arc 
rtaOy <li»Tinct from European gervera. 

As genera of which the family position is uncertain may be mcn- 
Ckimtarfipsis from the Lower Kinieridgian of Bavaria, whieh 
iroin existing forms by its shagreen skin, and apparently also 

VOU It. £ 


by the presence of a remarkable spine-like tooth placed in front ^i 
the normal tooth of the mandible. The fin-spines of this gO- . 
approximate to those of the Elasmobranchian genus Asteraau^i^, 
Rhynchodes from the Devonian of Ohio, and Ptyciodus from thtt^ 
both Russia and Illinois, are genera founded upon teeth which 
describers refer to this order. Some of the genera founded 
the evidence of detached fin-spines, a few of which are noticed it 
the preceding chapter, should perhaps also find a place among the 

Finally, the genus Calorhynchus may be mentioned in this as- 
nection. It was founded upon spines originally r^arded as ihft 
rostra of Sword-fishes, but which are evidently of dermal origin, and 
are considered by Mr S. Woodward as being probably fin-spins rf 
Chimseroids. These spines occur in the Chalk of England tad 
Maastricht, in the Lower Eocene of England, Egypt, and India, and 
also in the Middle Eocene of Bracklesham in Sussex. 

Order IV. Dipnoi. — The Dipnoi, which Dr Giinther r^aidi 
OS a subdivision of the Ganoidei, are typically freshwater fidu^ 
usually presenting the following characters : The body is covered 
with imbricating cycloidal scales, while the vertebral column is car- 
tilaginous, and there are both anterior and posterior nostrils placed 
more or less within the mouth (fig. S8i). The primitive cartila- 
ginous cranium persists more or less completely, and, like that 
of the Chimseroids, is autostylic ; cranial membrane bones are, 
however, always developed to a certain extent, and there are also 
splenial and articular bones in the mandible, while the cranium ii 
immovably connected with the vertebral column. The palatopteiyg- 
oid bar persists as the functional upper jaw, and, as in the living 
Chimseroids, carries a single pair of molariform teeth, while a cor- 
responding pair of teeth are placed on the splenial and articulai 
bones of the mandible. There is also a smaller pair of vomerine 
teeth ; in advance of which there may be other minute teeth. The 
paired fins have a long, cartilaginous, jointed, median axis (fig. 845), 
and the tail may be either diphycercal or heterocercaL There are 
no functional hranchiostegal rays ; and the five or six cartilaginous 
branchial arches are more or less rudimentary, and their single apo"- 
ture is closed by a gill-cover. The teeth agree with those dl the 
Chimseroids in having no successors ; but, from the presence of 
membrane bones, do not constitute the whole of the jaw. 

In the structure of their skull the Dipnoi show affinities to Chi- 
mferoids. Ganoids, Teleosteans, and Amphibians; the autostylic 
feature connecting them with the former, and the double nares 
with the latter, in which the skull is also autostylic The lungs are 
formed by the connection of the swim-bladder with the gullet by 
means of a duct, and these fishes can thus either breathe by means 



fiOi in water, or by their lungs on land. In Ihe existing 
\hva^Urmi iher€ arc cxicmal luanchial tufts, like those of 
of the Amphibia. The stnicture of the peJvic girdle and 
ihubecn already noticed in ilie iniixxiuctory chapter on the 
i; hut it may be observed ihal allhouyh the skdcton is cssen- 
Dt ctnibginoas, yet imperfect ossifications may occur in the 
bal jpioes of the vertchm:. as well as in the ribs and fin-rays. 
he scales may he cither ganoid or cycloid. 

71ut thb order is essentially nn old one, may be itiferred not 
djr bom the widely scattered dtstiibution of the three exiting 
QCfai, and their paucily in Kjtecies, and KDiiietime.s in individuals; 
It also From the geneiali.sed structure of its members, and the 
e on enee of one of the exi&ling genera in the I'lias of Europe, and 
; nrfiuttd Pcmiian of North America. 

KvMiLY Lepim>mrenid.«. — This family is only known by ivro 
nting gciMni, and is characterised by the perbi^^tcnt chondro- 
. carrying a few large metnlwanc bones, hy the cyct<iid scales, 

ri^ \ f ^ 'Lif i icli nm PMrad***, fram Sa1■I^ Amctjciu Reduced. 
/, Fectonl ; i>, Pilvk An- 

e absence of ji^lar jilates, the continuous vertical fin, and the 
m»r central axis of the paired fit», which are reduced to filaments, 
le two genera are U^donrtn (fig. 879) of ihe .Amazons, in which 
f paired (ins are not frinKcd j and ProtffpUrus of the rivcrK of 
~^ ' Africa, where those fins are furnished with fringes. There 

Ib'-'I'W BfCTfliiiaAi tConarfw ftrtUri), hdn (Juntutand. Xtduced. 

small conic&l vomerine teeth, and larger cuspidate teeth 
I the palate, while the body \» eel-like. 

Family Ceratodostid*. — The genus Ctratodus (fig. 880) has 
eo generally placed in the san>c family a.^ the preceding forms, with 
jich it agrees in the coniinuous vertical fin, the cycloid scales, the 

of jugular plates, and the few cranial bones. In many 



respect!), and more especially as regards the dentition, CWnto^^ ^ 
pointed out by l>r Kritsch, is, however, much more nearly allied 
under- mentioned family Diflerida, and il seemK ndns:ihle to 
it as rciiresenling, at least provisionall)', the t>'pc of a di&tinct S^' 

Ceratf'iius is one of the very few insUnces where a genus fo" 
upon the evidence «f fosiil ipecimeiis, has ;(ubaequcntly been d^*-* 
ercd in a livinjj condition. Fossil teeth of this genus were long kra 
but it was not until the year 1870 chat the existence of a living 
resentativc wis brought lo the notice of science. The body oa 
Barraniundas, as thcac fishes arc termed, is lateraUy com; 

with one continuous vertical fin ; while- 
paired fins arc paddle-shaped, with a 
fringe. The vomerine icclh (fig. S81, 
are shaped like the incisors of many moiW 
mals, while those on the palnle {iiiJ., at 
and mandible have an inner smooth con* 
vex border, and externally bear a nta^ 
ber of strongly-nurked ridges or bomL, 
In the e\isting species the teeth of o 
site sides are separated by a distinct 
tervol (f[f,. SS 1 ), but in some fossil fo 
they were in contact. Again, in the living 
species, the palatal teeth bear six distinct 
horns, while the b-ickward production of 
the inner margin fonn^ an incomplete 
seventh horn (fig. 881): but in the man- 
dibular teeth there arc not more than *c 
typical six horns. In fossil forms, accord- 
fng to Professor Miall, the mandibular teeth are slightly smaller and 
narrower than the palatal ones, and have not more than four horns; 
vrfiile ihe palatal teeth have either fi%-e horns, or four and a rudi- 
ment of a fifth (fig. 88j); 
this simpler structure of 
the tcelh in tbe eailiei 
forms being analogous to 
that which we have already 
memioncd as obtaining in 
the Selachian JVoHJamms. 
Some of the fossil teeth 
indicate individuals of two 
or three times the sice of 
the Rarramunda, of which 
the largest specin\ens at- 
The position of these upper teeth 
on the pa]atopter)-goid bar is wcU shown in fig. 881, which also ei- 

f*t>eri. Kcdneed. ■, Vomt- 
tini; 11. I'sUthI: asi, Mandi- 

bul^r tmh ; ■, oDiarinr ; u', (■»• 

Pl|. tS>.— Thi rich) palslil lootli <A Crrattdmi llif 
l»/(tmmi, fcoin ihc Ixiwcr Condim 
Tm kR *ida it ih* anivria htknlu . 

fcoin ihc IxiwcT CondimuA oT ai tleri, IncIU. 

tain a length of nearly six feet. 




ilhtlBse puasphenoid, and the anterior and posterior ante*. 
inpiinis distribation in past times, nc meet wicli remains of 
(ptnailie Jurassic of Colorado and Montana in the United 
lUdibo in tlie Lower Jurassic of Stoncsliclcl near Oxfuid ; 
lOMBofFT, abundant in ihe Upper (Keuper) and Middle (Mus- 
^}Tnu of Europe, and has been recorded from the Lower 
iBmlcr) of that formation ; it also occurs in Illinois in 
,«kithaK thought to be probably Triasstc; while in India 
kvrmycoaiirKin in the Maleri group oftheGondwana syiiicm 
[ivCoitnt Provinces, which is not improbably also of Triassic 
I'wilwT. Professor CoiK' lias recorded the geiius from strata 
laid Ancttca identified by him with the Permian ; hut the 
ifettD the European Permian originally referral to this genus 
1 to Ctenedus, 
funvi pRANEROFi.cvRiD£. — In this family, which has been 
[fiRcl bjtmat in the Crossopicrj-gian Canoids, w-v Ktill have the 

■Miuial litt. A. 'iaXt nusnilisil. 

conttnuoos vertical lin, the diphycercal caudal lin, and the narrow 
am of the paired fins characteristic of the preceding families ; Imt 
the scales arc said to liave a j^anoiJiil structure, jugular plates ate 
present, and there is a scries of minute teeth in the margins of the 
j»w». This family is tyincatly represented hy Phantroficuroit (fig. 883), 
of the IVvonian and Carlwnifcrous of Scotland and the Devonian 
of Canada, and the allied Urtmtmus of Ihe Scoliish Carboniferous. 
W'e (nay, however, here menlion the genera Mtgapteumn * and 
C»meiufioma from the Permian uf the Continent, which should prob- 
ably find • place in this or an allied family. The former genus has 

■ Tbc (taaiBlicndal icale* which have lieen dtKritwl it )K!:loTi(!iii|; 10 this gcniu 
UK iboM of ■ Ganoid. The nnnte ttiould ptopcrly be At<s^l«plmriitit. 



OSBiiied n'bs, large operculars, and Dipnoid teeth, but th« foniii 
the fins and tail is unlinau-n ; vrhJIc the latter agrees with 
pfeuron in the structure of the teeth and tail. 

Fa-MILV DirrERiDjT.. — This family is characieriicd by the 
development of the cranial bones, the more or less ganoidfll 
ture of the scales, the presence of jugular plates, the heter 
tail, the two distinct dorsal fins, and the greater sim and breadthl 
the scaled portion of the paired (ins. Its range extends from 
Uevonian to the Ferminn, and it shows signs of connecting 
preceding family with the CrossoptcrygJan Ganoids. The teA^ 
(fig. 88 5) are of thv same general type as those of CcraUtla, 
but may carry a larger number of smaller ridges, which in toot 
insitances (lig. 885, t, s) are ornamented with a number of cu^ or 
dcnticulcs. The lyijictl genus Diplcrus (fig. 884) comprises fiafao 
of small or medium sixe, with circular scales, and both the dotal 
(ins placed in the hinder Uiitd of the body, the Jirst being mud) 
smaller than the second. 1'he |>ectoral fins are long and paddlB- 

Pig. O^^DifHm yaJnrlKHuii : tmrn llie Dtmnion oT Runig, lwo-4biid( 
nalual liic. (Ah«r PBiider-) 

shaped ; while the pcK-ic pair, and the anal, are respecli\-cly placed 
beneath the first and second dorsals. The quadrate is ossified; 
and there are also o^sitications in the fin-rays and ribs. The leeth 
(fig. 885, 1) carry numcrou.'i dcnticulcs on their ridges. This geiUB 
is characteristic of the IJei'onian (Old Kcd Sandstone) of Europe. 
Ctettoiitts attains considerably larger dimensions than the preccdtnf 
genera ; some of the s]tectes reaching a length of nearly five feM. 
The leeth are characierised by their ridges carrying many cusps; 
and the scales are targe and thin, with a rhomboidai contour, and 
bearing traces of rows of dcnticulcs, with vascular grooves on tiw 
inner side. The skeleton closely resemble* thai of Ctratodus, but 
is more fully ossified ; and l>olh this feature and the more numeroiU 
cranial bones are regarded by iJr Fritsch as characters of greater 
specialiwtion. The teeth (fig. 8^5, a, j) arc frequently simpkr 
Ihan those of Dipltrus, and thus approximate to those of Certttodms; 
while the form of the parasphenoid and palatoptcrygoid differs con- 
siderably from that in the former. This genus ranges in Europe 
from the Carboniferous to the remiian, being very abundant in the 


ox) h b also recorded from strata in Illinois and Teras 
ire correlated wjih the European Ptrraian. 
}A3immhtn of thU order, but perhaps indicating one or more di»- 
I bmilei, we may class the following genera : Pala^aphns, from 
[Zteruaian of Europe and Ohio (ihc apecies from the latter area 
been sepanled as ffeJioJus), is a moderately brgc form in 
ihc mandibular teeili (fig. 883, 4) are very broad, and carrj* 




\oma jam nf t>ifum Vaittttbtmeii : tram ibe ttrvoaUiL «K, 
:a>ih: a. ■. N>n>J jpnm-o: f, faUloMirKoid. *, I'aUlc of 
/»■'■ 4iboniftm». 1. ftUiadiLk nl Clnidmt imititaiat; CkrtioairaiMU. 

at rti^'t-'f^wi imtifii: Dtvonlui. All ndveod. 

ffoor low ridges. Hohdui, from ihe Devonian of Russia, is a 
fonii very imperfectly known, but apparently allied to the 
while Cotuhodus u known by teeth from the &ame de- 
sits in both Scotland and Kussia. Gunarhynckut. of which the 
rizon is unknown, may also be provisionally placed hcr«. From 
orth America we have Ptyemodtn and Gnalfwrhisa from the Per- 
iaa, and Mytoshma fiotn the Dc^'onian, of which Ihe full affinities 
^uire fitrtbcr elucidation. 


CLASS PISCES— continued. 

Order Ganoidel 

Order V. Ganoidei. — The Ganoids form an order exceedio^ 
difficult of definition, owing to their close connection on the ooe 
hand with the Dipnoi and (through the Acanthodea) the EIudxk 
branchei, and on the other with the Teleostei, and it is not impnib- 
able that it may be eventually necessary to divide them into at tent 
two orders. As mentioned above, some writers group them in a 
subclass with the Teleostei under the name of TeleotomL The 
body may be either naked or covered with shagreen skin, or with 
large detached bony scales, or completely covered either with trot 
ganoid scales, or with cycloidal scales of a ganoid structure. The 
vertebral column, again, may be either cartilaginous or fully ossified; 
and its termination in the tail may be either diphycercal or hetero- 
cereal. Paired and median fins are generally present, and the pelvic 
pair (with perhaps one exception) is abdominal in position. The 
skull may either be covered merely by cranial membrane bones^ or 
may be completely ossified. It is hyostylic — 1.«., there is a hyoman- 
dibular suspensorium — and the palatopterygoid is distinct from the 
cranium ; and, as a general rule, even in the cartilaginous fonns 
maxillae and dentary bones, which carry the teeth, are developed on 
the palatopterygoid and Meckel's cartilage respectively. The gills 
are usually free, and their single aperture is covered by an oper- 
culum ; while branchiostegal rays are very generally present In 
most cases a secondary pectoral girdle of dermal bones (clavicular, 
supraclavicular, &c.) is developed externally to the cartilaginous 
scapulo-coracoidal girdle, which alone exists in the preceding orders. 
Finally, there is a swim-bladder, with a duct into the pharynx ; while 
there are some other characters of the soft parts into the considera- 
tion of which it will not be necessary to enter in this work. 

Some very curious features occur in the ossification of the rer- 


oatumn of ccruin members of this order which call for 
notice Thus in £uryft>rmi4i the ossifications in the dorsal 
nraut of an upper and a lower hollow. i*-ctlgtvshap«I, semi- 
,wiih their pointed extiemtties Jnicrlocking, and ihc former 
the neural arches and the latter the ribs; while in the tail 
two centra to each neural arch. i'>om this it would appear 
piecci bearing the rilv> in the dorsal rrgion, and the caudal 
hich have no arches, correspond to the intercentro of the 
la, which arc rwticcd bclovr. The teeth of t'ranoids vary 
1 structure, and may be either conical and home on the 
of the ^ws, when they are continuously rcpliiccd, or flat- 
Eks aitarhed to the vomer, which have no successors. The 
of the 6ns are frequently fumii^ed with Uie modified scales 
& fulcra. 

[krablc diversity of view has prevailed as to the clasMfiratiun 

ds ; but the system of Dr rraquair. who lias paid especial 

to the structure of the order, a adopted in this work. 

[an)s their distribution in time, Ctanoids first apfiear in the 

nearly at the same time as the Hlaainobranchs ; and from 

Dtiian to the close of the Mesozoic they form a verj' large 

Da of the Fiih-life of those periods Their wane, however, 

to have set in during the Upper Creiaccou->, when ihc 

li bc^tn to be nuntcrically strong ; and from that date there 

n a rapid decrease to the present day, when we find only 

the suborders (Amioidca) rcprescnlcd by sevcml genera j 

the other three surviving sutiordcrs one is represented by 

Bra, and each of the other two by a single genus — these four 

Bch having a solitary »pccics. 

ROCANOID Srries. — The first three suborders may be con- 
|r grouped in a sJr^Ie series, and are mainly characterised by 
Secdin^y low development, their afliniiies being still doubt- 
rofessor Cope is indeed disposed to regard one of these 
n (Placodemaia) as more nearly allied to the Tunicata 
other N'rttchrau ; but this view b scarcely likely to find 
ion with the majority of pala!untalogtsts. 
RDER 1. CEPHALiASPinEA. — The members of this cxtraor- 
poap have the bead and the anienor part of the body 
with a continuous shield, while the test of the body is 
nth stnall anguLir plates or scales. No traces uf an inner 
\f lower jaw, or teeth, liavc >■« been disco\crcd ; but at lea.<tt 
tcoftid family there was a strong pcciorat fin at the hinder 
gr of the dorol shield. The latter has been recently showt* 
nmisbed with a sj-stem of sensory canals. This (roup is 
I to the Silurian and Lto-onian ; and it is suggested by Pro- 
[uxley that it may be allied to the Acipenscroidca. 



Familv Pteraspidid*. — The entire body in this familj u 
known i but tbe dorsal shield (figii. SS6, 887) i.t usually com^ 
of screral pieces united together, and has its borders thickened . 

FiK St&~)intwrftcl dond 
kU of C/ai*>Hi ■ - ■ - 
troai the Silurlao 


uriao d Ijidlow. 
(AfW Murabuwu) 

PttratfU miml* ; rroai *• 
Low«r Ikevonian of K«refcic4. 
Rodiwtd. (ATiet Unlualcr.) 

bent, and in structure approaches the scaIcs of Teleostean Fuhes. 
It has a. median posterior spine, and sometimes small lateral cornui. 
There is atso a small shield on the ventral aspect of the lull, 
which, like the dorsal one, is marked hy fine striations. In tbe 
type genus Pkrasfiis, the dorsal shield {fig. 887) is shaped like 
an arrow-hcad, and composed nf smrn pieces ; while in Qm/iJ- 
aspis {fig. 886) it is oval, composed of only four cleracnu, and ha* 
the posterior itpiiie very short ScafhasfU has been founded on 

fig. an.— Reduced tvtart,v]aiai)C Fttrmoit' 

remains belonging to these two genera, which are found in both 
the Silurian and Devonian ; the simple shields described under this 
name having apparently been placed \-cnlrally beneath the cnoie 
complex ones, on the evidence of which the other genera were 



Helttspis, from the I-owet Devonian of North Wales, is 
[datmguubed by the doruJ shield, coiuJsting of only a single 
[cknenL Figore S8S ts an attempt 10 restore the form of the type 

Fxmir CtM*ALASPlDlD«. — Thf Ccphalaspidts are regarded Iiy 
WW •rtere a* belonging 10 the PttraspUiJa, while by Professor 
Zaid ibey are placed in a distinct suliordcr. The dorsal cephalic 
; ilatid (^ 889) consists of one or two pieces, and usually has more 
' Of k» developed lateral comun. with a regularly eurvcd and flat 
lonr border — the eyes being situated near the middle ; in structure 
it rexmbles true borte. The body i:t covered with bony plates, and 
the caudal (in is heterocercal. The tj-pical genus Ctphalaspis (figs. 

AboM aecililR] nuont lu*. <Afl<r LMibetUi.) 

1^9, 890) has (he cephalic shield single, and may be divided into 
ihr Enttpkatasfiitifu and tJemictphataspidim groups, according 10 
Ihe d^m of development of llic cornua. It occurs in the Si- 
bill) aod the Devonian of Briuin, and has also been obtained 

Txz.tvf-iimattvtntiCtplmtmfitLyHU. (AAcrfajtcl 

fifooi the Silurian of Bohemia and the Devonian of Canada. Xen- 
atfit, from ibe same English furmatiunN, in dLstiiigiii»hed by the 
presence oC one or more quadrangular dorsal plates behind the 
head ; vrhtlc: Anchemaspis, of the Kuroi>can Silurian, has ihc cephalic 
shield transversely divided. Didymaspis, from ihc same formation, 
agrees with the latter in its double cephalic shield, which is, how- 



ever, unprovided with comua ; while other genera are 7%yesles uA 
Tremataspis, from the Silurian of Russia. 

Suborder 2. Placx)dermata. — This suborder is mainly dot- 
acteristic of the Devonian, although represented also in the Siluriai, 
and lingering on till the Permian. It is characterised by the out 
laginous vertebral column, and the at- 
closure of the head and the anterior pot 
of the body in bony plates, which ait 
covered with a radiate or granular sculp- 
ture; the tail being eiUier naked or 
clothed with scales. There is a distinct 
mandible ; teeth are frequently present ; 
and there may be a jointed pectoral fin 
enclosed in a bony covering like that of 
the body, but pelvic fins are invariably 

Family Asterolepidid*. — In the 
typical family the head (figs. 891, S9*) 
is rounded anterioriy, and covered with a 
number of small thin plates ; while the 
body is sub<|uadrangular and invested 
with larger plates, some of which are 
median and others paired ; and the tail 
may either be covered with much smalla 
scales or naked. There is a well-devel- 
oped pectoral limb, which articulates with 
the anterior ventro-lateral plate ; although 
it was long thought to articulate in some 
forms to a separate thoracic plate. The type genus Astervl^, 
which occurs both in Russia and Scotland, is characterised, accord- 
ing to Dr Traquair, by the anterior median dorsal plate overlapping 
both the anterior and the posterior dorso-laterals, and by the some- 
what depressed body. In Pterichihys (figs. 892, 893), on the other 
hand, the median dorsal plate, while overlapping the anterior dor- 
so-laterals, is itself overlapped by the posterior dorso-laterals, and 
the body is much more elevated. This genus has been recorded 
from Scotland and the Eifel, and probably also occurs in Russia. 
Detached pectoral fins of these genera have been described 
under several names, and were at one time regarded as ichthyo- 

The genus BotkrioUpts, from the Devonian of Europe and 
Canada, is distinguished by the different contour of the cephalic 
plates, and of the grooves of the lateral line system by which they 
are marked, as well as by the shorter limbs. No traces of the scaly 
tail have been observed in any of the known specimens, although 

Fig, B91. — Reduced restoration 
of the domal aspixt of Atltreltpii 
entala ; from ihe Dsvooian of 
Ru«ia. llie tail ii mlor«d from 
PUriehthfi. (After Pander.) 



Dr Traquair considers that this appendage was probably present. 
Some of the species attained very large dimensions. 

Ft-S^.— Redocol rulontton of ihe dona! (i) miid<icnmt{a)ii3pecttof P/rrititiftciimiittit, 
nkiit icDipturv ocnilivd^ from the DcvoniAn of Scotland- The ihin block line* indicate the 
cEQOiikd ed^o of ibe overlapped plates, and the double dotted linc« the sroovev of the lateral 
WiyitoB. m.KC, Median occipilaJ plale; i.tcc, Laleial do. : af, Angulac plate; /r.M, Post- 
■odUa do, : p'9m, Prvmedian do. ; /, Lateral do. : /-/, Extra-lateral do. ; mn^ .Mci^tal du. ; sA, 
ScBilaur do. ; n-m^ii, Anlcrior median dorsal do. : fi.7H.Jy Posterior do. do, ; a.ti-i^ Anterior 
tEv^laicra] do, ; f-d i, Pfnieriordo- do, ; n.r',/. Anterior ventro-Urera! do. : fi.T.f, Posterior do. 
^: m.T. .Median ventral do- ; ar. Articular plate of limb ; a, AiKoncal of do. ; c. Central of do. ; 
■. yirginal of do. (After I'raquair.) 

SfUrohraehius, of the Scottish Devonian, is an alhed form distin- 
guished by the smaller size of the pectoral hmb ; in this form also 

Fij. Bgj. — Reduced left lateral aspect of /'/rriV*;*i'« eomulut; from ihc Devonian of 
Scotland. Lctien a> in the preceding figure, (After Traquait-) 

10 caudal scales have been observed. The length of the head and 
arapace in the one known species is one and a quarter inches. 



Family Coccosteid.e. — The genus Coeeosteta differs so maikedy 
from the members of the preceding family that Dr Traquair has iti|^ 

gested that it should form the Qrpe 
of a distinct suborder. The bead 
has distinct bones and [dates difia<- 
ing markedly in their 
from those of the preceding 
but their structure has otdjr 
cently been rightly explained (fi( 
894). In the restoration given 
by Agassiz (fig. 895), although the 
arrangement of the posterior plata 
is fairly correct, yet anteriorly the 
grooves of the lateral line woe 
mistaken for sutures; whUe the 
ethmoidal bones at the muzde 
and the orbital notches are omk- 
ted. An approximation to a cn- 
rect restoration of the lateral aspect 
is shown in fig. 896, but the mouth is made too long. The caia- 
pace, or body-shield, consists of a long shield-shaped middle doisal 
plate, flanked by lateral plates, and completed by a median ventnl 
plate. The posterior half of the body is totally unprotected, but 
tnterspinous bones in the vertebral column support an anal and > 


Coccotttut dtcipitntl from the Devonun of 
Scoiiand, at. Ethmoidal, with naret of 
either side; f.m.x, PremJuilU ; /.a, Preor- 
biial, emernal to which i> the orbii ; pl.o, 
PoctorlHtall f, Ccntnl, m, Murgitm], n, 
Midiile occipital ; i.h, Luna] do. (After 

Fig. 995.— Donal view of Cttceittnt dtcifitnt, u rotored by Aeuui. In the anuiw pan 
of the head the eihmoiil4 and orbiii have bevn omiticdT and the black iinci moatly indicaic ibc 
grooves oflhe lateral line tytlem, and not lulureh The ihield-ihaped plale inmedialcly bifaiiid 
the skull ii the middle dorul, in advance of which ii the middle occipit^ of the aknU. 

dorsal fin. There appears to have been no pectoral fin, although 
certain forms which have been generically or subgenerically separ- 
ated as Brackydirus have been represented with such an appendage. 
Both this and the next genus are characteristic of the Devonian, 
Coccosteus being common to Europe and Canada. The allied Homos- 
teui includes gigantic forms from Scotland and the Eifel, readily 
distinguished by the form of the cranial and body plates. Thus 
the middle occipital plate is longer and narrower, and the middle 
dorsal wider than long, and not pointed behind. Professor Huxley 



jcooparo the anoour of the Coccostcans to that of the Siluroid 
T* fcoit c« n *; while Ncwbcny and Pander compare the ventral 
with the [jlaslTon of the Chclania and l-ihyrintIio<Ioniia, with 




tnd mouch ue insda coo [one- 

vtiich. hoirever, they also compare the true pectoral girdle of the 
SMTOpter^'gia. The nature of the dentition is unknown. 

Faulv t>ixicirrHYtD«. — The genus DinicMthys, from the Dc- 
noiui (Horonian) of North America, may probably I»e regarded as 
Innning the type oT a family allied to the preceding. In this genus, 
fJ which the type species has an cslimBtcd length of from 15 to 18 
Im, the dentition (fig. 897) is remarkably like that of the Dipnoid 

f^fcf. — DUpBinna^anleriM wpKioTikK i«<n of Z>ffi*fl/4H llt-tfi; Irani lb* 
CWMiiBa tf Konk fltiln OneonlAh oatunJ iLk. (Afltr Newbury.) 

|BBB Prota^nii, from which Dr Tiaquair concludes that there 
m probably a close conrwction between the present group and the 
Kpooi. Other gigantic fcoms more and less closely allied to the 
tfpc genus are Titamtktkys, Liognafhus, and Diplopiathus, from the 
Oeronian of Ohio; while lypodtts, from the same formation in the 
EifiK, may perhaps belong to this ^mily. 

Ai Placodcrmata, of which the serial position is uncertain, may 
be mentioned Mcnasfris, from the Pemiian of (!t:nnany; Atanih- 
Mtfiis, Atantht>!epis, and Aspidkkthyi, from the Devonian of Ohio ; 
fiugea^mtAut and Lecracanthm, from the Carboniferous of Iowa ; 
SkAMMmihu and Pk&deraiantkus, from the corresponding fornia- 
tioa of IreUnd and Belgium ; and AtwrnatUhtkys, from the Devonian 
of Germany. 

SuBOROBlL 3. AcAKTHODEA. — Thc last gfoup of thc Proganoids 
ife abo PatKOioic, ar>d ranges from ihc Devonian to thc Permian, 
altboi^ it is not improbable some of the genera, founded upon 
rhich arc noticed under the ELisraobranchci, iliould be 



placed in it The Acanthodeans appear to be in some 
intermediate between the Ganoids and Elasmobranchs, and 
perhaps constitute an order by themselves. They have the 
which is more or less elongated and compressed, covered »ift1 
shagreen-like scales, and with the lateral line running between tvoj 
rows of such scales. The tail is heterocercal ; and the fins hnc [ 
strong spines, which, except in the pectorals, are merely insertei) 
between the muscles. There is considerable doubt as to the pi» ] 
ence of cranial bones or of a gill-cover ; but there is a ring of bono 
round the orbit The vertebral column is cartilaginous, and teedi 
are either wanting or are very minute and sharp. \ 

In their cartilaginous skeleton, the not improbable absence <rf lo 
operculum, the structure of the scales and position of the latoJ 
line, as well as in the spines of the median fins, the Acanthodo 
approach the Elasmobranchei ; but the articulation of the pectoid 
fin-spine to the pectoral girdle is a character of the Teleostean ^- 
lurida, while the orbital ring is a character of the higher Ganoids Kkc 
the falaoniscida. 

Family Acanthodid^e. — All the genera may be provisionallf 

Fig. I9B. — AcoHlheJtti from ihc Permian of Europe. (Afiei Kner and Rocnwr.) 

included in a single family, of which the type genus AattUkodes (fig. 
898), as now restricted, ranges from the Carboniferous to the Per- 
mian of Britain and the Continent. The head is very short and 
blunt, and there is but a single dorsal fin placed immediately above 
the anal, while it is thought that teeth were absent Mesacanthus, 
of the Scottish Devonian, includes small fishes distinguished from 
the last genus by the presence of an intermediate pair of small 
spines between the pectoral and pelvic fins; it is represented in 
the Devonian of Canada. Closely allied is AcantkodopsiSy from 
the Carboniferous of Northumberland, in which there were numer- 
ous minute teeth \ while Chiraeantkus (fig. 899, i), of the Scottish 
De\'0nian, is distinguished by the dorsal fin being placed in advance 
of the anal. In Dtplamnthus (fig. 900) there are two dorsal fins, 
of which the second is placed above the ana! ; each pectoral fin 
has two spines ; while there are minute spines between the pectoral 



pelvic fins, and ihc yx»i% are furnisSed mth small coiiica] 
This genu« is represeiiied by several species in the 
aiin of ScoOuhI, and not imprubably aLso occuis in thai 
^ Ca n ada. JUka^Haaini/itu, from [hi; Scottish Cirbonifciousi, 



k— I, CUraMMIiju jVKFT*iMHit: t.CUiHAHu* tntlgtr : 3, iKJaattiUkla ftttJli i 
from i^ EkevMilanoTScbdud. Redund. 

nro in pbce of Tour pectoral spines; while in /sfhnaMHfhus 
g. 899. 3). of the same depovtts, the small intermediate sjiinc^i 
K nnting, althovigh the>' arc introduced in the figure- Allied 
lo ihc bst are Eu/fuKaHlfsas and Parexks, from the Scottish De- 
nun, the fonner having the second dorsal in advance of the anal 


r^ «■» —IN*!aiMUkmi if<iu%i : hom ilw P^vonun of ScuHand /, ]'«CMrjil ; v. VtiiU^ ; 
«.Anl;L, CMiikli ^t,^rimHvdHO»lil<><ml<n>. KvdiMtd. 

fin. Fuudlyi the genus Oimalius {fig. 899, a) includes three small 

I fiabc9 froni the last-named formation characterised by the short fin- 

*pincs and the presence of a scries of accessory spines between the 

pectoral and pelvic fins. There aic two dorsal fins, of which the 

VOL. II. r 


second is behind the anal, the figure being incorrect in the 
respect Fin-spines from the Devonian of Canada, describ 
Oenacantkus and Homacanthus, are referred by Mr Woodwi 

IncertjE Sedis. — Here may be noticed the family Tam 
proposed by Dr Traquair for the imperfectly known genus Tan 
from the Scottish Carboniferous, which is regarded as indicati 
aberrant type. 

B. EuGANoiD Series. — The remaining members of the 
include its typical representatives, and may be collectively I 
as the Euganoid series or the Ganoidei Veri. 

Suborder 4. Crossopterygea. — In this group the pectora 
sometimes the pelvic, iins consist of a central lobe surroundet 
fringe ; there is an infraclavicular in the pectoral girdle ; the i 
the dorsal and anal fins are often more numerous than the su 
ing interspinous bones ; the preopercular extends forwards ( 
cheek ; branchjostegal rays are replaced by jugular plates ; t1 
velopment of the vertebral column varies ; the tail may be 
diphy- or heterocercal, and the scales cycloidal or rhomboidal 
suborder is the most primitive of the true Ganoids, among 
it holds a position somewhat analogous to that occupied I 
Ichthyotomi in the Elasmobranchei. 

Family Holoptychiid*. — In the type family the pector 
are acutely, and the pelvic subacutely, lobate ; the skeleton 
former, according to Dr Traquair, being a biserial archipter 
like that of Ceratodus. The teeth are of the so-called dtnd 
type, the dentine of the base being in 
in an extremely complex manner, with 
branchings which form an intricate n< 
within the crown. The scales are eye 
thick, and sculptured ; and there ai 
dorsal fins, and a heterocercal tail in 
the inferior rays are much longer th 
superior. Before noticing the tjrpical 
we may briefly mention the genus Onyi 
from the Devonian of North Amerit 
diii%^Bidei: from thToe- Europc, which Dr Newberry considei 
(AftwNcwb.^")' *"'™' be allied to this family. The scale 
901) have distinct ridges, and the ma 
has a presymphysial production furnished with teeth arranged 
what like an old-fashioned cavalry spur. The type genus 
ptychius (fig. 902) comprises fishes, which are often of larg 
from the Devonian of Europe and North America. The two 
fins are placed in the hinder part of the body directly over tl 
vie and anal. We may probably place here the imperfectly 1 



.^larrptaihvs, fram the Devomsn of ScoUatuI and Ruuia ; It&dut, 
of The Irish Caibonircrom ; and PtpiorktHa, from the Trias of 
11;h!i-«4, In ClyptflejHf (fig. 903Xfi'<>in the European and Canndian 
I V> onion, we have a wdl-knuwn form in which the dorsal tins 
-irrtl more anteriorly than in HoJopfychms, wliile [he anal is 
■A bdow tbe 6nt dorsal, and the pelvic arc consequently much 

■ u J uLu J. Thtdon*! and wul flm ibwU b* BOn ftantA. 

led to the pectoral fins. PkylhlepU, rounded on 

scales which arc someiinnes smooth, from the Devonian 

1 Carboniferous of Scotland, should probably be included in this 

in which Dr Traqualr also places the imperfocily known 

DfdrvAit, q( the IJcvoniao of Russia and Scotland. The 

of the laiter carry a small row of marginal teeth of conical 

rhile in ihe mandible there is a second row of much larger 

■Hlce teeth, each of which has a distinct soclccL Thcw leclh 

(AlWr Kuky.) 

somewhat compressed, with trenchant foreand-aft edges, and 
have the internal structure already noticed, CtrtenoduSt of the 
Brkoh Carboniferous, and Sigmcdux, of the Upper Paleozoic of 
nonbem India, should perhaps find a place in this family. 

Familt RiiizoDOXTiii*. — In the Rhizodonts the pectoral fins 
are subacutcly or obtusely lobate, the pelvic pair being usually non- 





lob«te ; while iheir internal skeleton forms a shortened tu^ 
ftrchipierj-gium. Ttic teeth are Inbyrinthodont in structure, !■ 
the complex internal network found in the preceding family, 
scales are cyclotdal and sculptured ; the median 6ns are numei 
the same as in Hohpt)ic!uus ; Init the tail ap 
mates lo the truly diphycercal type. The 
jaw has an inner scries of tuiik-likc teeth 
905). As an imperfectly knon-n foim we 
first mention CricoJus {Po/yp/ocoditt) froR 
European llevonian. In the well-knowm \ 
ckopttrnst of the Scottish Devonian, the be 
much elongated, the tail distinctly heteroc 
the posterior fins are placed near the tail 
opercular bancs are unusually large. an( 
ecalcs thin and striated. An apjMrently 
form is Eusthtnopterau of the Dc\'onian of 
ada, in which the vertebral centra appear to 
been unossificd, while there arc slight difler 
in the interspinous hones, tail, and teeth, 
cording to the views of DrTraquair, Gjnvfit^ 
(fij;. 904), from the Devonian of Scotlar 
allied to Tnstufu'pttrtUy having scales 
same general type (although some of thea 
formerly thought to be rhumboidal)i but a 
rhombuidal tail, and the exposed poitioa 1 
scales relatively larger. From the Cartx 
ous of both Europe and North .\merica wi 
the type genus JihiioJus {MegtUitAiAyj in 
of which the typical sjtccics attained a very 
si^e. The larger teeth (tig. 905), on the evi 
of which the genus was onginally founded 
the upper half of the crown smooth, whi 
lower half is longiliidinalty fluted. The tc 
t.trt;e individuals (which may attain a len 
nine feet) are nearly two inches in heigh 
Jihis'^d/'psit, of the same deposits, the ■ 
Iwncs form a well-developed :^cld on the 
rior surface resembling that of the Ositoie^ 
although the mandible agrees tn general str 
with that of Jifiizodus, and consists of an 
angular, and dcaiary elements ; the latter 
n series of ittfra-deniary pieces on its inn* 
which carry the targe lusk-ltkc teeth, 'i'he inferior space bctwc 
mandibular rami is occupied by a series of jugular plates, of 
there is a large median pair, together with a i^niall unpaired a 


•"■S' 904. —Cyrrpif 
<Aitu a/tciltri I frocD 
llii Dtvficiuui of ficoc- 
lud. RwlucsL (ATin 



, md a row of small lateral onc3 on cither side. In the craniuin 
■t striking feature is the anterior position of the orbit, which 
I three Urge suborbital bottes behind it. Other more impcrfcctlj' 



. >»»■— A iMiOwli r nmmt ^JtAjtnliu /Irttni: (VmathcCwboolfcroatorSimUixI. 

genera of this family from the European Carhonifcrous are 
MyeAiut, Slrt^sodus, Khomboptyihius, and Ari-hichlhys. 
PajauTOsTEOLEPiDm^r. (Rhourodivtebid^.). — In this family the 
hsve the same general structure as in the preceding ; hut the 
lolei are of the rhombotdal form typical of the order. There are 
a numler of bteral jagular plates ; the teeth are numerous and 
ihuply-pointed ; and the 0!»ificution of the vertebral column is 

FkE. gafc ^F*ifptttia tkkirl ftta ib* Uh"' KUc «, OiUMMii mttnlttidUtu; nam 
*» D iiMib««fKtiM». (AJWPtadv.) BMb Iwiw mtunJ. ■. Ptciotvl ; ^ P^ric ; <. Anal I 
i, f, Donal tat, la th« l«irw tew* ila iar^ Cm ibonld be ulai.'a] aion in advuicc ol [be 

■npcrfcct. The family is divided into two subfamilies; the first, 
QtftwErjhV/tM {SaurodifiUrini) being characterised by the smooth 
Kales, and presence of a median jugular plate. 'I'hc type genus 
OstwUpis (fig. 906, b) has a long and slender body, with the two 
dwsaU ptaced respectively in advance of the pelvic and anal fins. 
The mM tpecie> u of considerable size. In Ihe allied Thurtim, of 


which an immature indnidtial is shown in fig. 907, il 

placed iiiimedi.iifly over llic pelvic and ann) fiiis. Boitj 

in the Devontnn of Scolhiid and Russix Closely allied is the | 

Difhptents, from the Scottish Carlioniferous, in which the 

fins arc also situ.itccl posteriorly, iht first bein^ directly mtil 

pcU-ic. Mcgalichtkys, from the same dcpoaitii, comprises two . 

of large siice ; and is characterised by the presence of large fulai 

at the roots of the pectoral, pelvic, and anal fins ; and also b; tic 




Fit- $07.— A ytiout Walivitlul tl Tkuniti mtinrbfUtfia: fion it* I>fK>niiui (tfScMlBi 
Lcllenuinnt- two. Keduoed. (ATtu StdEwidi and Murcbhoo.) 

small ivic of the anterior dorsal fin, which is placed above ili> 
pelvic fin. Edosttorhaehis, from tl>e Permian of North Ainenc% 
is nearly related, and appears to be the last survivor of the fai&ilf 
The second subfamily, or Ghptohcmina, iu represented only by I 
genera Glyptolamut (lij^. 90*) and Glyptopomm, of the 
Devonian ; and differs from the last by tlic sculptured scales, 
the absence of a middle jugular plate. Tlie body is much do* 
gated ; there is a long interval between the pectoral and tbe poatoiff 

rig. ^f£.^^{ftttlmm»a A'iimainti : fnun ilir P(vnni:in of Soxlind. Keduocd. x. Sola. 

(ins ; and the tail is truly diphyccrcal ; each genus is represented 
by a single species. 

Familv C(el\canthid«. — With this family we come upon 1 
group of fishes not occurring in the older Palaeozoic, bat extrnding 
from the Carboniferous lo the Upper Cretaceous, and thus affording 
a link between the preceding extinct and the following cxi&ting 
family, The vcrlcbral column is unossified; there are two dor^ 
fins, each of which is supported by n single forked intcrspinous 
bone ; the pectoral fins are obtusely lobate ; the caudal fin is un- 



f; iMge, and completely diphjcercal ; ihe swim-bladdcr u 

ied a«d die wales arc cycloidal. The members of this family 

^ ki^Kt cbanKtemed by the full development of the cranuil 

I,- uid the airaogeineM of the ouifications in the orbil .ip> 




I ^UaJittm ttmtdiltta : (ron Ihc KiriMtiilciui <•! naiatu^ onc-ruiuth 

whes that found in the Labyrinlliodoniia. The type genus 
'JaiiimtAlu ranges from the CarbonifeiDus of Euiopc und Xurth 
Berica to the Upper Trias of the fonncr area, and is well charac- 
bed by the great ihkkncss of iu scales. 
More or lest nearly allied arc Difiurns from tlie Trias of New 



$»«.— SUctM J U*ertf*wt* MnaufK: fram itic Chalk of SuM«i. Radund. 

Grvfkaints from the IJpptr Tria^ of Camiola, and Ut/lti- 

*a ranging from the Middle Trias lo ihc Upper Jurawic of the 

ttincnt. In Vndina (fig. 909) the expo&cd parts of the stales 

marked by ridges, n!>iiig in some parts into spine^t, and the 


rays of the dorsal and caudal fins are divided and furnished 
spines. In the English Liasstc Hohphagus the spines oo 
scales are more numerous, while in Libyi, from the Ki: 
of Bavaria, they are less numerous, the division of the fi: 
extends more deeply than in Undina, and the dorsal and 
scales have a row of tubercles. Coceoderma, from the Kim< 
of Europe, is allied to the last. Finally, MacropoMa (fig. 
from the Chalk, comprises several lai^e species readily characteised 
by the notochord not extending to the extremity of the tail 

Family Polvpterid*. — The last family of the suborder is knon 
only by the existing African genera Polypterus (fig. 906, a) and 
Calamoiehthys, each of which is represented by a single specici 
I'he vertebral column is ossified ; the dorsal fin broken up into 1 
number of small finlets \ the pectorals are obtusely lobate ; die 
caudal fin is diphycercal, with a very short body-axis ; and the scak> 
are rhomboidal. I 

Suborder 5. Acipenseroidea. — According to the views of Dt ■■ 
Traquair, the Acipenseroids, represented typically by the Sturgeon 
and their allies, and forming the Chondrostet of many authori^ Vt 
also taken to include the Heterocerci, as represented by the extinct 
Palmoniscida. The following are some of the leading features of 
the group as thus defined. The paired fins are non-lobate; tbe 
pectoral girdle, which in the typical forms retains its primitive 
endoskeletal cartilages, develops dermal bones, among whidi the 
infraclavicular (fig. 913) is characteristic; the dermal rays of the 
dorsal and anal fins are more numerous than their supporting carti- 
lages, or interspinals, of the endoskeleton ; while in the paired fins 
these dermal rays have to a great extent replaced the original carti- 
lages. In the skull the cartilaginous cranium persists in the typical 
forms, but in all cases it is overlain by a series of dermal bones ; the 
preopercular, when present, tends to extend on to the cheek ; bian- 
chiostegal rays are generally present ; but there are never large 
jugular plates. The notochord is persistent, but there are either 
cartilaginous or bony neural and haemal arches ; the tail is heteio- 
cercal, and the skin may be either naked, or dotted over with bony 
scutes, or covered with rhomboidal scales. 

Mr Smith Woodward remarks that the typical forms of this sub- 
order constitute a link connecting the cartilaginous Ganoids with 
these fishes in which the bones are fully developed, and that their 
paired fins are more specialised than the median ones, which have 
not yet attained a numerical equality between the fin-ra>-s and their 
supporting interspinals. 

Family Acipenserid^ — The well-known Sturgeons (fig. 911), 
which form the typical family, are large fresh-water fishes, charac- 
terised by their elongated body, produced snout, toothless jaire of 



Isdult, the pre«enoc of five rovs of scutes formed of true bone 
bodft and the sculptured cranial bones. The repreMntativeit 
cxiKing genen An'fienser (fig. 911) and Seapfurhynehm are 
in many of the lar^jer rivers of the NoTthern Hemisphere, 
rdmcd to Aiiffttstr occur in the l^ndon Clay, and in some 
Tertiary beds ; while spines from the English Upper Eocene 

Ttf^ 9*t.— Tlw SlarivBa (^cylcmu'r lAiri*). Much ikIuckI. 

the Pliocene of Montpellier probably also liclong to the same 
A feingle srute has also been described from the Miocene 

i^irginta under the same name;. 

fAMiLY PoLVODOKTiD-B. — In this family, now represented by 
A^vuEm of the Mississippi and Psephurm of two rivers in China, the 
lltfai is typically nearly or <|uite naked, the mouth is oT enoimous 
width, and the jaws carry n^inutc teeth. In a fossil state this family 
b Te|«csenuxl by Crouopholis, from the Eocene of Wyoming, which 
ditpUys many points of resemblance to Pcfyodon, but is remarkable 
br ibe possession of »mall pectinated scales, which arc nut confinixl 
lo Ae ttpper lobe of the tail. The cranial bones are of the type of 
those O^ PofyoJan, but llie shorter rostrum indicates a resemblance to 
PyfMimtt. The scales are numerous, and are arranged in oblique 
mws, wbich are not quite in con- 
csict with one another. The rcien- 
tun of the scales in this gi-nus in- 
dicates that wc have tu do with a 
moch less .■ipccialiscd member of 
the £iinily than the exiting formi<. 
The genua Maavpelaluhtkys, from 
the Ttevooian of both North Aroer- 
ka and the Eifel, has frequently 
been reftrrrcd 10 the P^yodontiiitt, 
b«I without any sui5cient evidence, 
nd it» serial position must for the 
pfcscnt remain undetermined. The 
oanium (fig. 912) is short and 

Ivosul, with the orbits completely surrounded by bone, and the 
middle line occupied anteriorly by a diamoml-shapccJ ethmoidal 
ihidd, which articulates posteriorly with a process from the squared 


Ply. 911 — Dli^rum ol Ihe («irilj! xpMI 
of Ihc •jwiliun of Moi'efilalitiii*}! Haiti' 
ngmi ; trom iha [.iv^f-TLtin of North Arnar- 
KL MiuJi rvductt). {ATitr Neabcrry.) 



As an Acipenseroid of uncertain position may also be mcnt 
the imperfectly known Pholidurus^ of the English Chalk — the I 
of which presents some resemblance to that of Pi^hurus. A I 
fish from the English Upper Lias, named Gyrvsteus^ is 
also an Acipenseroid agreeing with the type genus in the 
of teeth, but having a naked body. It differs also consi<i 
from the type of the next family, and its position must 

Family CHONDROSTEiDi£. — This family is formed for the 
tion of the one comparatively small fish constituting the 
Chondrosteus from the English Lower Lias. According to 
Traquair, although there is no evidence of a long snout (fig. 913 
this genus resembled Polyodon in the general form of the fins, aolj 
of the nearly naked body ; but in other respects — such as the fml 
of the cranial bones and the absence of teeth — approaches Ae^tnm. 
Certain features also appear to indicate distinct affiruty with tbe 

'"■ ••*• 

rf, Dcnuiry; mx. 

Pig. gi3.~rroli1e view of ihe head of Chtndmtm atiptmereidtil firom (be Lmnr Um rf 
iJoiseishirc. Reduced, f. Frontal ; //, PcBtfroni«l ; », Parietal : ly, Sqiumonl ; t.l. Out d 
ihe supraiemporaU : /./, Pmtlemporal ;, SupraclavicuUr ; cl, Claricular; I'.c^ InfradMC- 

ular: o». Opercular; i.ef, Subopercubr : hr, Branchiosiegali ; c.i, CnatoliTal: u^AnsBla'; 
r. Maxilla ; j, Jugal ; Km, Hyomandibular ; i.a, SnborbiUiL (AncrTnqBafe.) 

Palaaniscida ; and since the latter are evidently allied to the hig^v 
bony Ganoids, Mr Woodward suggests that they or allied ianta 
may have given origin to two series, one of which culminated in the 
Teleosteans, while in the other " the only advance has been in die 
matter of size, and this accompanied by a certain amount of de- 
generation, culminating in Acipenseroids proper." 

Family Pal^oniscid*. — This and the next family constitute the 
suborder Heterocerci of some writers. In the present fomDy the 
body is fusiform, and covered with rhomboidal scales ; the vertebral 
arches are ossified, but there are no bony ribs ; the dorsal fin is 
single and short ; all the fins have large fulcra at their bases ; ikt 



booes are covered with ganoinc, and moiitl)' sculptured ; the 
is large ; the first bnnchiaitegal ra]rs foim jugular plates ; 
\ h s row of me^inn V-shaped scales on the superior surface of 
lity of the bod)r; and the teeth ore small, conical, or cytin- 



, I*— Lift Uml vin> trf Ibc 4l:uU of Pairmxumi miK rsA-mw ; tiain Ihe Permian. A 

;£ Pnntel; «f^ S q u aMowfc l i t,t^ !nprat«nifpcr»l : ^'. t'<miiempoi^\ ; t, EtKmoic! ; a^, 

■I; /Kx, rminIlU : Bx, MaiillB; a, «, Ui Sii)«rliiult . i/, I>tii[ary: i-/, Ot<r- 

■Aifft IH»»|i«ii:iiUr : i^ SubopcrCBUr ; tr, Bnn<ht<HU(ali> : i/, Llariciilu ; ac^ Supra- 

Ik I ^A PMulnnlB i k/, lafiacUricaUti »r,Oibil. (Aftci Traiiuur.) 

il, and rarely plicate at the base The chief characters of the 
■ of the skull and of the sccondar}- pectoral giidlc arc shown in 
14, in which it will be seen that the preopcrcular covere a por- 
bT the face above the large and broad niaxilla. Pr Traquair 
mes that if a Fal.'coniiicid were stripped of its scales and fur- 
d with a long snout, it would be so like Polyodon that there 
■ be no doubt as to their near relationship; and he further 
ks that the oitc row of V-shaped scales found on the iloraal 
if the extremttjr of the body in Fdyodon is identical with the 
kf scales occuin-ing the same position in Palaoniscus. This 
to the conclusion that the existing SturgeonB are the sun'ivors 
Erics of Ganoids, now totally lost, which fomicrly gave oif the 
ymisdia and Piatyiomida as specialised branches, 
is Eajnily embracer such a targe number of genera, that space 
Is of but tittle more than their )>aie enumeration. In the 
btao CMrvlefiis, fournl both in Europe and Canada, the body 
>der , and the fin$ are large, with the rays very finely divided ; 
tf>is (fig. 915) of the Rothiitfjiendes {Lower I'cmiian), 
iins of somewhat similar structure, is distinguished by 
Uld EQore spindle-shaped body. CosmopiycHiut, of the 
Carboniferous, is allied to Rhabdoiepis, but has the anal fin 
longer base. In the type genus Palaoniseui (figs. 914, 916) 
y is much elongated, and the tini> arc very small and widely 
It occurs in the Kupfet-ScUtefer and Magnesian Lime- 



stone (Middle Permian) of Ccrmany and Engbnd ; and one : 
has t)Mn said to occur in the English Trias. That species k, i 
c\'cr, now referred to ihc Dapediidie ; but a true Paiizomi 
recorded from the Hawlccsburj- beds of New South Wales, 
may be of Triassic age. The remaining better known gcnen I 
Ehnkhtkys, IVoni ihe Carboniferous of Europe and America ; 



rig- 9ii.-~lliatititti4 matrt^ira ; from itia Low« P«raiUn of Suoey. KtdMlAi 
(Ahcr Ac«>li'} 

hpis, ranging from the Penniaii to the Carbonirerous, and perh 
to the Pe^-onian, of Europe; Ntmat^fityckius, Cytlofityihiui, Min 
odu$, and Gonatodus, from the British Carlionifcrous ; the 
Amblyptrms, of the <ierman Permian ; ithadiHuhthys, from The I 
boniferous of both Europe and North America; JEurylefiis, fioca' 
the Carboniferous of Ohio ; Hohtrus, Canabmt, and PMaHfrMte«ll^ 
of the Scottish Carboniferous ; Pygopltrui, of the German Permim; 
MyrioUpis, from the Hawkeshury of New South Wale* ; UrKthemn, 
from the underlying Newai>tle group of the latter country; Gyrt- 

I'lg. Bit.-— yj^xmifm UMmjtwMiu : rmin itie MMilIc PcrmUn of Cicrnuny. RadaeaL 

ilir/fcf, from the Middle Trias of the Continent ; and Oxygnathia, 
Cosmfl/eju's, TJiristoUfis, CentroUfiis, and LUsolipU^ from the Eng- 
lish Lias. Here aho may be mentioned the genera ^/utrolepit, of 
the Permian of Bohemia, and Ccefahph^ of the Kimeridgiflti of Bavaha, 
which, althuufjh having cycloidal imbricating scales, appear to be 
allied to the present (iamily. 



ivr PuTTsoMtDjV. — The l^atysomidec agree wiih the preced- 

in the character? of the vertebral column, fins, scaJcs, and 

main cranial siniaure, but difftrr by the body Ijeconiing deep 

I ibon. with an ovoid or rhoinbuidal contour, and also by minor 

JOS in the cranial struaurc I'hc iccth nrc small, nnd 

<: eitber rfmrp or obtusc The chief diiTerence in ihc struc- 

Ihc skull consists in the deflection or its axis below thai of 

tcbnU cclumn, instead of being continuous; while the hyn- 

r, itutcad of l>cing very oblique, becomes nearly vertical, 

: the eUunoid is elongated. These changes cause the moiirh to 

: KparUcd widely from the orl>it, the jaws to become " prognath- 

xnd the gape of the mouth itself to be tuu(.-h rcUucc-d ; tliey 

readily apparent by contrasting the figuic of Palaoniscus *itii 

of CkirodMS. Dr Traquair regards this family as a group of 

' ilised forms descendtd from the Paiaenhci4a ; their external 

nhUnce to the Dapcdiidn being probably due merely to adap- 

I to itmiUr conditions f£ life, and not indicating a real aftinit)-. 

:£urymotu5 (5g. 917) the contour of the body h not iio greatly 

r«.vtT— < 

(Alto Tnaiuir.) 

from the PaJseoniscid t>-pe ; the pectoral fin is large, the 
doml also large, and beginning above the pelvic, while the base of 
the anal la ihwi. This genus occurs in the Carboniferous of Scot- 
bnd and Belgium. In Jienedenius, of the Ilelgian Carboniferous, 
the body becomes more oval, and the dorsal fin is placed more pob- 
totioriy. More or Icia nearly allied 10 this grouj.) arc Afesokpis, of 
the British Caiboniferous ; Eurysomus, of the Middle Pennian of 
id and Belgium ; and IVardich/kys, of the ^coilisli Carbon- 
In Chirodas (fig. 918), of the British Carboniferous, the 
is rhomboidal ; the dorsal and anal fins liave a lung base, 
lays, and an anterior spine : while the pelvic fin is unltnown, 
and the pectoral ^niall. Closely allied, again, is Chirodopiis^ from 




the same horizon. The last and type genus Platysomta (figii 9I 
and 930) is represented by a large number of species, ranging fa 
the Carboniferous of England and North America to the MBdd 

Fig. giB- — CkiradMi gnaiulBtiu : from the Corbooif emn* of Scotlaad. Rcdacad. 
(After Traquur.) 

Permian of England and the Continent The contour of the bcx 
is less rhomboidal than in Chirodus ; the dorsal and anal fins he 
no anterior spines ; the pelvic fin is very small, and but seldcKo pf 
served ; while the pectoral is of medium size. 

FiK' 919-— /'/o'/nwiiu tiriatta; from the Middle Pcimlui of EosUad. Rednoad. 
(After Tnquur.) 

Suborder 6. Lepidosteoidea. — In the Lepidosteoids the paii 
fins are non-lobatej there is no infraclavicular bone; the rays 



and anal fins com-spond in nunib*r wiih their suppnning 

IS bones ; Uit opcri:ular bones arc like those of tlie 'I'cle- 

tbcre is frequently 3 median jugular plate on the tirst pair of 

uoscegsis; the dettlopnaent of the vertebral column '\^ries; 

is of the masked hcieroccrrcal t)~pc ; and the ica.\eH arc rhom- 

or may be occasionally replaced by angular scutes. This 

f .ry.i'iill"""*'*' 

Jii! IS 

TIm pclvtc lln 1* nix ihom. 


^■BnSer is represented at the present day by the somewhat abet- 
jK L^d»s/t£dx, and also by a large number of Me^ozoic (onra 
anting from tbc Trias upwards. As already mentioned, the exter- 
na rescDlblance of the Dap4diida and PytnodonHda to the Ptaty- 
wdim b not iq^ded by Dr Traqunir as indicative of real afHnity. 
Fakilv DArciiiiD^.. — In this family, which by some writers is 
mbdirided and known as the SauriJa: and Slylodantida, the body 
i< riUser fmifonn or ovste ; the opercular bones present certain 

chancteintic features ; the upper lobe of the body-axis of the tail, 
lod usually lite anterior borders of the oiher fins, have well-dijvcl- 
oped fukta ; the \-omer and jaws carry sc\-i:raJ rows of small teeth, 
rf which the ouiennost are curved and resemble claws {griffehahnt) ; 
lad the ossification of the vertebral column is imperfect. Among 
iie genera with fusiform bodies, orw of the best known is Stmionohis 



(fig. gai), which is widely spread wcr £im^. .According loj 
I)ccclcc this genus is rcprcNcnEcd in the BuntCT and Miischdk 
it is ccimmon in the Kttiper, or Upper Trias, and thence 
the Kimcridgian, or Up|h;r Jurassic. It has also been 
from ihe Scormherg beds in ihe upper part of the Karoo sysufal 
South Africa ; and it a))pL'firs to be aho represented in the 
of North America, where it has received the name of luk 
In these fishes tlie dorsal fin is ainiJl, the inc<juabty of the 
[x^rtion of the caudal »(lrungty marked, and there is a row of 
bpinc'like scales on the middle line of the h.nck. Other mon^ 
less nearly allit-d forms, of which some were formerly included i 
Palacniseus, are Aienfro/^horus, from the British Permian ; 
pttrus, fro]n the North American Trias ; and Dictyopygt, from I 
Trias of both Kngland and North America, the type species <A ' 
latter havi[ig I)ccn originally named Palironistus supersttt. 
HiltrfUpidotui, typically from the I^wer Lias of England, 
HfiemstrDfi/uis, from the Kimeridgian of Bavaria, we ha« 
in which the contour of ihe body is more like thai of the type | 
H(leraUpidt>tm, according to Dr DeecWe, is also represented in ths] 
Muschelkalk, and surx'ived to the Kimeridgian. The name ASt- 

kpidolus is applied to allied typca also represented in the Muschcl- 
kalk. Dapedim (fig. i)7i), in which Aiihrnodm may be inchidcd, 
comprises b number of medium-siied fishes with broadly avatc 
bodies ranging in Europe from the Keuper to the Lower Jurassk, 
btit aUo occurring in the Kota t>eds of the Indian (londwana^, whkh 
are higher than the Malcri beds from which Crratodut is 
obtained. The dorsal and anal fiiis have elongated ba»es, and the 
inequality liclwccn the upper and lower loI)cs of the scaled part of 


[caudal fin is oomporathvljr slight In tyjMcal species the teeth 
ItBD^ilc, while in others (on which vfrAwtWcr/ was founded) they 
[ibriied at their summit ; since, however, both types .ire oocn- 
'ibaiid in a single species, ttiey do not afford grounds for 
: dkdnction. The u>-c.alled Amifyurut has been founded on 
I liisk spcdmcns of DapeJiut. Allied fomis aiu I'ieuroUph 
\ibmnnltftSt from the Upper Lias of ^\'^;Tl^.-IIll)L■^j•. In Tttrn- 
w, of the European Lias and llic Koia beds of the Indian 
the liody resembles iJiat o<" the type genus, but the form 
donal and anal fins is different, the caudal An is nearly sj-m- 
ADd the vcncbral centra develop rinfp> of Ihirc ; all these 
I approximating to the next family, in which Sir P. Egcrton 
Ixtth this and the following gciius. Ch't/int/efits \s an allied 
trjMcally from the Wianumaiiu and Hawkesbur)' beds of New 
W'alcs, but also occurring in the Stormberg bedn of the South 
Karoo system, and not improbably in the Indian Kota beds ; 
stnta being of approximately equivalent age. 
KuiiLV P^CNOix)xiii>«. — ^The Fycnodonis form a compact 
ranging from the Lias to the Eocene, in regard 1o the serial 




ns iMj. TT^Ii'iiMi. tatBt, and nrlibnl (ctumn oT Pycnodnnu : m, Jawt, and i, vomerine 

iTl - II- ** ' -*■ -^ "---■-• - Da. of CrA^ki .- J. Venctnl eolunn at PairJiat/jl^m : 

A Dot 4( ft mil* I ; /. Sota «f Gyrtiiu. The dtsiIiSM ii. itiutly ibIuibI. 

poatton of which very di%-ergent view's have been entertained. Dr 
Traquair is however, dii^iosed to regard them as specialised forms 
connected with the Dapedudtc and Ltpidotida^ and they are aecord- 
ii^y here placed bctvreen those families. Tliis family ha^ been 
tcccirdnl from Europe, Asia Minor, North Ameriea, and Australia, 
and pfesents the following characteristics. The body is of a rhomb- 
VOI. uu c 



oitlal shape, presenting a striking resembUnce in this rcs|i 
ihe Platysomdit ; the cnudal fin is of a coinpleielj- ina;^kMl 
certa] typt;, md there are no fulcra to the fins. The notod 
pcrsislciit, but the neural arches and ribs arc ossiAcd, and 
later forms the heads of the latter (fig. ^i^ itrc cnUr]gcd, at 
simulate portions of vertebra! centra. The prcmaxilLe arc too 
the conjoint palatines and vomer form a triangular bone (fig, 
carrying five longitudinal rows of o\-al or round molariform 
while in the nundiblc the dentary hones, which form the exi 
of the symphysis, have two or four chisel-like tevth, and the S{l 
are enlarged and carry on cither side from three to fiv< «( 
rows of mobriforni teeth (fig. 923) opposed 10 those of the! 

i Commencing nith the typical genus Pym&dm, ure find this, I 
defined, restricted in Europe to tlic Lower Eocene of Shqipi 
the Middle Eocene of Monie Bolca in Italy. There are onlj 
rows of teeth on either side of the mandible, of which the inn 
is the largest ; tii 
m ^-1--*'''^?^^^^—-. '''* vomer tlie U 

1 ^^0^"^^^ ^ ^^^^^ •'^^ \\vKe. inner ro 

■ M jMB^^^^- \ round, and tho 

■ ^K .;^MHptf9^^Ps^^>i^^^ I larger and oval. 
I ^P^j^j^g^ Sr^^ irf ^^^^c j] * characters are foi 
I M TfcfteJ^StJ^^c&^SS^B^ ^^ position o 

I M ^' iKfeai^^^j * - r]^^ ' "'''''^ ^'^ ^"'8' * 

vJB^3^pl^R^^^^jf^^^^\ '^'^ tnoulh, aM 

I JUPC ^^S ^S^ t^^^'jr —^^^ . 1 supcrioriiyin ihel 
r^*' iyM^^X^^^^^^^^S^^'x ! °^ *^* dorsal M 

pared with the afl 
.■\ fish from Ihe 
CC0U8 of Brazil tia 
referred by Ph 
Cope to this t 
The allied /'•lAs 
/*« (fig. 923, . 
which all ibe 
teeth are sab 
occurs in the 
of the Lcbanoi 
Cretaceous Pisol 
Mont Aim£ in F 
and also in the Middle Eocene of Monte Bofca. The | 
number of the Cretaceous forms are, however, referable I 
genus AaviemitH} CorloJus,^ which comprises soioe species atl 

* The natna CnMui being preoccupied by Ca!od»n, the term linvttmn, 
powd tiy AgsMii for detaclwd teelh, it adopted. ' 


Titf^A.—ynrwaiAcnltmnmBrrriftiiil rnxii the 



sitt. The vomerine xeetli (fig. 9*4) of ihe middle row are 
and moch larger than either of the others, while there 
rmaOjr three rows on either side of the mandible, although 
occAfiOTUiIIy increased to four. The dorsal fin extends 
of the anai This genus ranges from the (lault to the 
and is widely spread ihrough Europe. In MetoJim we 
genus ranging from the English Lias to ihc Ixiwcr Kimer- 
<rf Bavaria, and not improhahly also represented in the 
Oeensaod. Here the Tomcrine teeth of the middle row 
I Itfger Ouii the teeth of the other rows, which are exceedingly 
and in the ouicrmoal Hnc have a w,-in-likc surface. .Allied 
■K Alnhirui and Athrodou from the Ktmen'dgian of the 

(fig. 935), whieh comprises two small species from 
CrcOccous of Italy, is tJiaractcriscd by the great Icngih 

td the dorsal and anal fins, and a]»0 by the concav-e surfaces of the 
aobnfann teeth, which are of »<ubcqua] sixe. Aficrodim is snother 
■Died genua, ranging in Europe from the Kiineridge to the Puibcck. 
The arrangement, uf the leeth, and the peculiar siruciure of the 
beads of the ribs arc shown in fig. 923. In Cyr^us we sccra to 
ba«e the most specialised members of the entire family, all of them 
Seitig characterised by the sculptured crowns of the motariform 
teeth. TIk vomer is very narrow, and has the Iccth subcirtular, 
tlioae of the middle row beit^ much the lar^^e^t : while in ihc man- 
<Eble tfaeie are four rows on cither side, of which the first and third 
are the larger. This genus is especially abundant in the Lower 
KinKiidgian lithographic limestone of KaYarin, hut it also ranges 
tipwatds into the Chalk of Sussex, and downwards into the l^wer 
[ Junnic Stonesficld Slate. Finally, Co^odus is founded on an im- 
L perteel !qKCtmen froni the Chalk of the Lebanon rcilly belonging 



to this family, but which has been rcTened to the Tele 

Familv I,epii)otii)-« (SpH^BonoNTin^,), — In Ihc Ltpidotia 
body is more or less rusirorm, ihe upper soled portion of the i 
longer than the lower, and the Tulcm of the Bna are well devt 
The palatine, vomer, mnxillt, and dentary cairj- rovrs of knc 
teeth (fig. 926), while the prcmaxilla is furnished with teeth 
^isel-like fonn. The type geiiu<; Lepitiotui (fig. 9i6>, with 
Spharodtis is in pfin identical, has a wide distribution 
time and space. Thus, in Europe it ranges from the Muscht 
or Middle Trias (where its scales have been described under ' 
names of Dactylahpis and Thallodus) 10 the Chalk, and it 
occurs in the Kota group of India, and the Cretaceous of 
and of North America. The large button-like teeih of Z. 
are abundant in the Kimeridge Clay, and «pecimen$ have 


ece £ 

vicnof tcMkuiilaKali mtivcainclK conKn. 

found comprising nearly the whole palate, and exhibiting the curious 1 
maimer in which the replacing teeth gradually turn t«-cr a:s ihqr 
come into use. The names Ntphrotus, Ctntrcdus, Omphaiodm, 
Htmihput, and Asteroden, have been applied to molariform and 
chiscMike teeth belonging to members of thi^ family from the Trias 
of Sik-sia and Thuringia ; while teeth of the latter type from the 
bone-btid of the same period, in both Wiirtemberg and England, 
described under the name of Sargodim, should, perhaps, be likewise 
sed here ; and NtorhomboUpis, of the English Chalk, appears to lie 

1 allied Form. 

Familv Eugvatmid* (Sauroiiostid*). — The body in this 
family is long and slender ; the snout short ; the fins hai*e fukas, 
the caudal being of a partially or completdy m.iskird hctcroccrcal 
type i the vertebral centra may be either imperfectly or fully ossified, 
and the teeth arc pointed. The range in time of this family extends 
from the Upper Trias to the Ncocomiar of Europe, but may, pa- 
haps, also include the Chalk. In the cjpical group we have E^t' 



ranging from the Keupcr to the Kiintritdguin, while in the 
find PlatytiiMgum and PiyikoUfii, and PMiJo/Uyrus and 
KJ in Ibc Kcupcf. A second group is represented by 
•i-^h-^hs. of ihe Kimcridgian of Bavaria, and also hy Th-frafo- 
\i :v.;<\ Pterygitf^tents *i\ (he Kciipcr. ihc latlcr heing disiingiiishird 
die ib*cnce of the pelvic fin*. The genu» Phtjiidopkerus in- 
.tnull Fishes aomewhai resembling a Carp in form, which 
(mm the Musrhelkalk to ihe PurbecV. A fish from the 
clkalk, originally deacrihed as PhaiiJi'Phortts porro, has been 
ihc tjfjic of the genus Prohitledttt, on accouni of peculiar 
in ils aqtiaznation. I.argci forms are ranked under the 
lAijAlnOff of the Lias anil Kimcrid[>ian. and Ophmpsis, which 
from the Lias to the Purlieck. Of the Tcmainin^ genera 
Euumiut and I*ropterus in the Kimeridgian of llavaria; 
from the latter deposits, and also in the Lower Creeiuand 
Hiitiomstui in the Ita^-arian Kimcridgian and the English 
xbeck ; Maa-osimius ranging from the Stonesfield Slate to the 
Kineridpan of Bavaria; and J^gnonotus from ihc English Lias. 

from the Chalk of Sussex, may perhapi he placed 
Ixrc. although its &ku11 approximates 10 ilut of Atniti, The scales 
;ted, juid hAvc a pectinated posterior hordcr. 
iiY AsriDOKHvwcHiO;« (Rhvnchodo.vtid*). — In this family 
'IS much elongated, and covered with scales of unequal 

ni- gajfA, Ttm C*r-pllia iLtpilmlnt uttta). Rcoinl. Korlti Anwrka. t^ AtfU*- 
rJ^X^tAiu; trom Ibc JunoMC' Bo(b niuh nriiwaL 

; ; tbe skull is produced into n short rostrum: the caudal fin is of 
Ik mnhed beteroccrcat ijrpc ; the fins carT>- fulcra ; the vcnebra* 
late ossified rings ; and the teeth arc either blunt or sharp. This 
amtly is rciiresented only by the Mesozoic genera Aipidorhynchm 
ind SttMioslamM, in both of which the dorsal fin is placed above 
he ttal. In the former (fvj^. 917, b), which ranges from the Lias to 


the Purbeck, the length of the upper jaw exceeds that of the 1 
and in advance of the mandibular symphysis there is 
bone (not shown in the figure) apparently corresponding to the) 
found in certain Dinosaurs. In Belonostomus, which, as DO* I 
striated, ranges from the Lower Kimeridgian of Bavaiu to 
Chalk, the predentary bone is so much elongated as to make i 
upper and lower jaws of nearly equal length, and the predenV 
carries a median row of large conical teeth, flanked by two roM 
minute teeth ; the teeth of the normal bones of the jaws hasi 
mammilated crowns adapted for crushing. 

Family BELONORHYNCHtDiC:. — This family name has beCD p 
posed by Mr S. Woodward for the remarkable genus Beiemorkfudk 
typically occurring in the Upper Trias of Carinthia, but also fiM 
in the Lower Lias of Dorsetshire, where the specimens had be 
originally referred to Belonosiomus. According to the writer qooll 
Belonorkynckus was allied to the latter genus, having a similar ki 
and slender body, with the same general position and structnR 
the fins, and probably furnished with a predentary bone. Thefidl 
of the fins were, however, either absent or very minute ; and, wi 
the exception of a median dorsal and ventral series of scutes, i 
another series on the lateral line, the body was naked. Tlie i 
perfectly known Saurichtkys, from the Rhaettc of Bristol, is a doa 
allied, if not generically identical, type. Specimens of the npf 
jaw show that (as in BelonorkyncHus) the bone was covered with i 
tubercles, and that the maxillae gave off horizontal palatal plates, li 
those found in Amphibians and Reptiles. 

Family Lepidosteid*. — The Gar-pikes of the genus Le^doA 
(fig. 927, a), inhabiting the freshwaters of Northern and Cent 
America and Cuba, agree with the Aspidorkynchida in the gene 
contour of the body and the arrangement of the fins ; but t 
rostrum of the skull is much longer, and the tail distinctly hete 
cereal. The scales are lozenge-shaped. The existing genus appe 
to be represented in the Lower Eocene of France by a species ( 
Maximiliani) formerly referred to Leptdotus ; and it may also oci 
in the Upper Eocene (Oligocene), where a species described uni 
the name of Naisia has been referred to it In North America 
have also Pneumatosteus, from the Miocene, and Ciasies, from 1 
I^wer Eocene, both being freshwater forms, and the latter a 
occurring in the Eocene of Rheims. 

SunORDER 7. Amioidea. — According to Dr Traquair's class 
cation, the last and most specialised suborder of the Ganoidf 
typically represented by the existing Amia and a series of Mesoz 
genera approximating more or less closely to the Teleostei. 
these forms the paired fins are non-Iobate ; the infraclavicular bon 
absent ; the operculars are Teleostean ; the branchiost^als hav 


jtifiulw pU(« ; the vcrtebrse arc more or less completely 

1 : the toil is masked hctcrocerca} ; and the scales are thin and 

Dt cydoidal. In aII cases the teeth arc small and pointed. It 

obatile, indeed, that in the Mcsozoic rcprcsenutives of this 

I we have Tonns closely allied to the atKre^ors of the Teleostei, 

lit H more th^in likely thai future discot-cnes will show a com- 

!pts»age between the Onnoids, as represented by this 8v1>order, 


lav Pacrvcormidjr (MiCROLEPtixyn). — According to Pro- 

■ Ztrtd, Paikyfprmtu and its allies should be placed in this sub- 

allhovgfa other writers have regarded them as mure nc-arly 

I to the Daptdiida. In ihcise forms ihc .scaler, alihou^'h thin 

ilcUiD^ ate subrhomlKjidal, and the vertebral column is 

^faiooiiipJctelr ossiBcd. 1'hc chief genera, which arc European, 

Vackywrmvs, ranging from the l.iaii to the Oxford Clay 

Jun»ic); SHiitHtis, from the Lower Lias; £uthyHvtm, of 

horiion : and ify/tiavmus, Saarofitis, and Agasst'iiay of 

*er Kimerid^nan of Ba^-aiia. 

iiuv CAirmm*, — 'llie Ctiutoids are a family of Salmon* 
. fishes, Taryint; greatly in :&i2c, and ranging in time from the 
I lo tlic Chalk, but especially common in the Ktincndgijin litho* 
jjpipbjc limestones. The vertebral column may be cither very imper- 
$esAy or completely osstlicd, the tail is more or less deqily forked, and 
liie liu have fulcra. Caiurus itself comprises a number of species, 
: of which arc of large size, and two of which arc represented in 
I jiS^KC The scales are \^ery like those of the Teleosteans, 

tC*twnit/mrtafiti ; from ihc KMuridgika al BkvarU. Reduced. 
1,- r, Pditii.; a, Aiiai ; f, Cmudtl ; S, DhmI iId. 

pg flin letain the Ganoid character of an inferior layer of bone and 
10 oppcr one of enamel. The dorsal fin is placed immediately above 
be pelvic This genus ranges from the Lias to the Kiincridgiaii, 
7, mcximtt! attaining a length of three feet. Stnthi/odus, from the 
Cioiendgian uf Bngland and the Continent, is a closely allied if not 
ientical genu^ Other genera are JsMofum, from the Dorsetshire 
xmer Uas ; and the Jurassic Lindesmui, EMrycormus, OUgifUurus, 
EjMWgj^WJ {AUakM^tis), Matrorhipit, and jEthalion. Oli^opltums 



has the dorsal lin placed behind the peK-ic ; And in thi$ Ani 
of The other genera the vertcbne consist of the upfMrr 

f,^ horse-shoe-shaix-d elements i 
-^ 'T ' in the prcliininnr)- notice of tl 

<£L\ ' Family Leptolepidid^- 

-j[KH|aL ing similarity in external coot 

j KeI ^^. the CatUToitls U the genus /A 

(fig. 929), which is placed ; 
(■iinther next to that family, aA 
some palieonlolugisis regard i 
TeleoMean. The vertebra! col 
fully ossified, and the fins ai( 
out fulcra. In this genus^j 
ranges from the Lias to the ) 
idgiiui. the dursa) fin is plocf 
mcdiatcl)' over the pelvic ; 1 
Thrissops, of the Kimeridgia^ 
Above the anal. In external i 
tcrs these Fishes cannot bej 
guished from Teleosieins; aaj. 
proliable that they are ind 
conneried with that order, C 
ihey should not be placed iol 
Family AMnn.f- — In the !% 
the vertebral column is fully ot 
.tnd in the caudal region has 1 
of centra la-aring the ncuif 
lixmat arches, which alternaH 
iiUcrrcntra devoid of such a| 
ages. I'ulcra may or not b( 
cnt ; and the caudal fin 13 e 
with the extremity of the V6 
column bent sliarply up int 
upper lobe. In Aftgafums. 1 
I'urbeck and Kimeridgian, fuk 
present, and the dorsal fin is 
and does not extend in advai 
the pelvic. Lophiuntt, fror 
same horizon, and Opsi^mis and Amiofsis, of the Lower i 
arc other genera, of which the first two arc nearly relsl 
Mtgalurus. The existing Amia, of Ihc freshwaters of the 50 
United States, has no fulcra, and the dorsal fin occupiesi 
quarters of the length of the body. It appears probable thi 
genus is represented in the Upper Eocene (Lower OligoceiM 
the Lower Miocene of the Continent, where its remains havt 

Fia- 9>8J«. — Skelelon af CalnrT,, 
f/finjptt^t ■' frtHn lUe K imcrifl^jjifi of 
Fiane*. Oiw-tuJ*' n^iunl uic. (After 



ioribcd under the names of Cydurus and Xotaits ; and it also 
in ih« Eocene of Colomdo and VS'yoming, where it has been 
as Pntamia and Ilypamia. The Eocenes of the laitcr 
and of Kheims have aJso jrielded the allied PappUhikys, 

dtere « but a single row of teeth ; and niemljers of this 
uaj occur in the Chalk of Ihe Lebanon. 
rXM. Seuisl — The remarlcable genus Darypttnu, of the Pcr- 
i of Hcssc and Durham, may hi; convcart-ntly noticed here, since 
ipcKttion is T«Ty pToblemiiticaL In thi^ remarkable lish the body 
riiqied somewhat like that of a Sunfish, and the anterior part of 
K doml fin IS taller than the whole depth of the body. There 
% ooreovei', the Canokl characters of fulcra to the fins, and the 
olodtordal condiiion of the rertctiral column ; hut, on the oiher 
led, there arc no ganoid scales, and the pelvic hns are placed in 
fnoce of the pectorals, as in some of the Tcleosici. On account 
' ihtt curious comhination of characters, I'Tofetisor Cope has pro- 
xedto make Doryptenn the type of a special order, the Dory- 
with the family Dorypttrida. 



CLASS PISCES—continmd. 

Order Teleostei. 




Order VL Teleostei, — The last, and in many respect! 
most highly organised, order of Fishes is the Teleostei, whic 
eludes the greater number of existing forms, and (unless soi 
the genera here placed in the Amioidea belong to it) does nol 
back beyond the Cretaceous. The Teleostei are in all proba 
descended from the Ganoids, and occupy in the class a somi 
analogous position to that held by the Squamata among the Re 
and the Fasseres among Birds ; all traces of Amphibian afiS 
having been entirely lost in this order. 

It is impossible to give a definition of the order by whidi i 
be sharply separated from the Ganoids, but the following a 

most characterisd 
-^ e^s tures from a pa 

'^\ -J ,'^ tological point of 

'y.'3-'.---j%, The body is it 
covered with thin 
tic, cycloid, or ct 
scales (tigs. 833, 
but bony scutes 01 
oid scales are occ 
ally present The 
of the endoskelei 
well ossified ; an 
gills are freely sus 
ed in a gill-cavity 1 
ed by a well-deve 
operculum (tig. 
The caudal fin c 
adult is of that completely masked heterocercal type usually t< 
homocercal (fig, 931). The pelvic fins may be either abdo 


Fig. 930.— Skull of Troui(5ii/mfl). Reduced. Letters 
as ia fig. St], p. 917. 



ibccd in advance of th« p«ctoniI«. The fin-r3.>-& nu.y or may 
artjculaicd ; and thtn are never fulcra on the fins. There 
pairs of nasal openings on the top of the head. Other 
are fbtind in the soft ports which arc not usually avail- 
ibe case of fosidls. 

[fiinher be observed that Tclco-iicans agree wiih OnnoJds 

tylic structure of the &)cull, and that cranial bones and a 

pectoral girdle (fig. 844) are alwa)-8 tievclopcd. The 

ent of the cranial bones in a typical Teletistean in shown 

930, where it will be seen that the preopercular (TV) does not 

|af Um PtKhlftrcMjtta'idtUijX Xxhifwl. lAtltn m la tg. Ejfl, {i. gij. 

on to the face as it docs in sonic Oanoids. In ceitain fomi*, 
•r (Si/uriifj'i, the cranium deveIo[>s large bony piates, artictJ- 
with othcn in the cennco-dorsal region, and presenting a 
»g resemblance to some of ihe cartila^inoiisi Ganoids. In the 
group Mrong npirK.-^, arliciiLiting with tlic urKlerlyin}; bones, are 
Dped tn tiw pectoral and dor?ia] fms ; while the .Salmonutds, miiny 
ids, and others develop behind the rayed dorvtl an ndditinnal 
ilhout raj's, to which the name adtpoit ot fatty fin is applii'd. 
icordii^ to Or Giinthcr's cbusifiralion, the 1'ctcostei are divided 
six suborders, but other writers would group the last five of 
toother under the nanic of I*h>-M>cly&ti, as of equivalent value 
e first, or Physostomi. The former arrangemtnt will he fol- 
Rd tiet«, but only such families as are of importance to the 
1*1 can be noticed. 

R I. PHVSOSTom,— In this suborder the swim- bladder, 

t, is connected by a duct with the pharynx ; ihc pelvic 

generally abdominal in position, and have no spine ; while 

lin-rays are articulated, but .sometimes only the first rays of 

and pectoral 6ns arc ossified. I'his suborder comprises 



the most generalised Teleosteans, and those most nearly 
with the bony Ganoids. 

Family Saimohidx^ — The members of this and the next : 
are so intimately connected by fossil forms, that it is very 
to draw any distinction between them, and it has accordingly I 
proposed by some writers that they should be united under 4e? 
name Hakcida. The fossil genus Hake is, however, very imps- 
fectty known, and if it be eventually found advisable to merge the 
two families, it would seem preferable to employ the name &/« 
iV«r in this wider sense. Existing Salmonoids are characterised bf 
the presence of an adipose dorsal fin, and by the premaxilla anl 
maxilla forming the borders of the mouth, and both bearing te«t^ 
There are no scales on the head, and no barbels to the moudu 
Recent Salmonoids are either marine, or inhabitants of the fredi- 
waters of the Northern Hemisphere. Remains of the existiiig 
marine genus Osmerus (Smelt) occur in the Upper Greensand of 
Germany, the Lower Eocene of Claras, in Switzerland, and the 
Miocene of Licata, in Sicily ; while nodules of unknown age iwsA 
in Greenland and Canada enclose a species oiMaihtus indistinguisb- 
able from the existing M. villosus of the former region. As fonns 
connecting the Salmonoids with the Clupeoids we may notice A^^ 
hpis, Acrognathus, and Osmeroides, from the English Chalk. The 
first has a rounded body and a depressed head, with minute teeth, 
and the pelvic fins abdominal. In the second the orbits are of 
enormous dimensions, and the teeth extremely minute Tlie tluni 
genus (fig. 932) is also abundantly represented in the Chalk of the 
Lebanon, and appears to have no adipose fin ; its skeleton in many 

Fig. g3a.~OtmttviJtt Utvetitiuh : Ironi the Chalk ofSuiMX. Rednced. 

respects resembles that of the Clupeoids, but the ribs have not the 
peculiar structure of that group. Sardinius, from the Chalk of the 
Lebanon, and Sardinoides from the same locality, and also from the 
Chalk of Westphalia, are more or less closely allied forms ; while 
Opisthopteryx, of the Lebanon Chalk, may also be provisionally 
placed here. 

Family Clupeid*. — The existing members of the Clupeoid or 
Herring family differ from the Salmonoids by the absence of the 



fin. And tbc presence of peculiar dermal ossifications at the 
i of the ribs, which form bony plates on the sides of the thorax. 
the numerous forms arc nuirine, and usually occur near the 
The type genus Clupea (Herring) occurs in the Miocene of 
iberg, the Upper Eocene of the Isle of Wight, the Eocene 
and Gtanis. and the Chalk of the Lebanon. The 

nt til — nc Cmbrmi Htfifnf iCtmt*^ Imra^aiy Rcdutcd. 

existing Herring i& shown in fig. 93J. The Lcfianon 
[dull hu aUo yielded a species of the li\-ing genus Engraulis ; 
\ ibc extinct genera Stomhvciupea, Lejitoiomtts,^ ChiroaKlrites (also 
t*B ff'enphaliA), ftnd Spaniadoa, which are more or less closely allied 
(•euting tj'pca. In the Eocene of the Conrincni wc have repre- 
KMUlves of the existing genera Engraulis (.Anchovy) and Chanos, 
ud the extinct Platinx. Ca-logasier, and CrasiognathuS — the latter 
idr^ allied to the existing AffgttM/ft. 

In this place we may conveniently notice a number of extinct 
geoen more or less allied to the ClupeoidH, but of which the family 



is produced into a rostram, occasionally equal in lengtli to 
entire body ; this genus occurs in the Chalk of the Lebanon, i 
is considered to show affinity with Optstfuipteryx. Another 
seems to be related to the existing Clupeoid genus Elt^s, but 
presents characters connecting it with the American 
family Characinida. Among these we may notice Rhaa^epiSt \ 
the Cretaceous of Brazil ; and in Europe Elopides, from the 
Eocene of Oarus ; Miopopsis, from the Cretaceous of Bohemia 
IstFta ; Hemielopopsisy in which the borders of the mouth ^ip^iea to^ 
have been devoid of teeth, from that of Lesina, on the DaknilflB^ 
coast ; and Protehpis, of the Bohemian Cretaceous, chaiactenMi': 
by the short jaws and the presence of crushing-teeth on the patalb 
Thrissopater, again, from the Folkestone Gault, differs from Ae 
modern Elopine type by its compressed body ; while Halec, of tbe 
European Chalk, is still very imperfectly known. Alosa^ of Ibe 

Fig. 935. — Skcleion of Diflamjiitia trrviitimia ; from the Qulk of ibc Lcbmnon. 

Eocene of Algeria, and Diplomystus {fig. 935), originally described 
from the Eocene of Wyoming, but subsequently found in the Creta- 
ceous of Brazil and of the Lebanon, are allied forms. The latter 
genus has a series of dorsal scutes, which are very characteristic 
Hemitrickias, from the Tertiary of Northern Italy, differs from all 
existing Clupeoids in having two dorsal fins. 

Family Protosphyrjenid^ — With this family we come to the 
first of a group comprising several families of extinct marine Fishes 
characterised by their large spear-like teeth, and hence termed 
Saurodonts. They should evidently be placed near the Clupeoids, 
although their division into families must be regarded as provisional. 
The present family is characterised by the production of the ethmoid 
in advance of the maxilla, to form a long cylindrical rostrum ; by 
the loose connection of the premaxillee with the maxillx ; and pro- 
bably also by the complexity of the mandible. The teeth were 
implanted in distinct sockets. The type genus Protosphyrana 
{Erisickthe) occurs in the Upper Cretaceous of both Europe and 
North America ; and the large spear-like teeth of P. ftrox, which 

nme distriliution as ihe genus, are very common in tlie 
lie b*ds of the Cambridge Creensand, and were for a long 
fctml la the American Saurvetpha/us land/ormts, which was 
bought to be a Reptile. I'hese teeth arc compress^, and 
t maiipnal senations. The 6ns were prOTidcd with large 
I spines, which were at one time referred to the Sctnchian 
f^hedtts, and subsequently were made the type of anotlicr 
under ihe tnme of Peteeopitrus. 

ilLV Dercetid* (Hoplopleuridje). — The members of the 
I family of SaurodonU are charaeterised by their elongated 

their powerful dentition, and the presence of several scries 
je triangular scutes along the side* of ihe body; there is 
te dorul fin in those fomu in which the 6nit are known ; 
nh are rwt implanted in sockets ; and the skull i& frec|uentiy 
Wd into a roftttum. This family probably passes impcrccp- 
iioihc next 

1 typical genus Dercttti (in which Leptotrathtlus may be in- 
I) occurs in the Chalk of England, Bohemia, and the [.ehanon, 
v> in the Upper Cretaceous of >\'estphalia ; it is characterised 
I kngth of the upper jaw exceeding thai of the lower, and by 
escnoe of five rows of scutes, of which the middle one hears 
eral lir>e: The allied As/ndopiairus is confined to the Lebanon 
' Other members of this family are Blaekiut, from the Middle 
e of Mnntc Boica; PHnthophorus, of the English Chalk; 
rhynehus, from the Upper Cretaceous of Westphalia; and 
•AamfAus, from that of Istria. Plinthophants has two rows of 

hut i)i otherwKe naked. 

BLV Enchoi>ontid«. — ^Thc genera which may be provision- 
ouped under this name are diKtinguLshed hy the moderate 
^ion of the body, which may be cither naked or 

K^ f j l st il l W of gwi j/ i» tit B*lnifr1i from ih* Chalk of ih* l^lanon. 
R^DCcd. (AfurPtcMiMidltumbiri.) 

9 with Ecuies, and by the elongated premaxillie and maxillae, 
carr^- small teeth andiyloscd (like those of the mandible) to 
tie. A considerable portion of the maxilla is excluded from 
Ugin of the jaw by ihe preraaxilla ; teeth occur on the pala- 


tines and ectopterygoids ; and the dentary bone of the 
has one series of large teeth, with one or more inner rows trf i 
ones. The type genus Enchedus, with which Eurygnathus of \ 
Lebanon beds is identical, occurs in the Upper Cretaceoa 
Europe, North America, Brazil, and perhaps India- Closdy i 
to this genus is Eurypholis (fig. 936), of the Lebanon Chalk, 
is merely distinguished by the presence of a few dennal 
Cimolichthysy from the Upper Cretaceous of Europe and the Un 
States, and Pomognatkus {Pkylactoaphalus), from the Chalk 
Europe and the Lebanon, are nearly related genera, mainly didii>i 
guished by their dentition. Here also may be placed Ischy 
tus, of the Upper Cretaceous of Westphalia, which may 
have had two dorsal fins, and appears to connect Etuhodus with ttt^ 

Family Pachyrhizodomtid,*;. — Allied to the preceding, butwift ■ 
the body more compressed, and covered with either scutes or scak^ 
or both together, is a group of Fishes which may be provisiona&f 
placed in a distinct family. The premaxillse and maxilUe are Utg^ 
and carry powerful teeth, which may form one or more rows, ud 
art; set in incomplete sockets, and anchylosed to the bone ; while 
the abdominal vertebrae are characterised by their longitudinri 
striation and the absence of deep pits. The type genus Ptukfrid^ 
odus {Hyfsodon in part) ranges in Europe from the Chalk to the 
London Clay, and also occurs in the Cretaceous of North America; 
its teeth are so like those of Reptiles that a lower jaw was described 
as belonging to a species of Mosasaurus. Mmpo, of the North 
American Cretaceous, may be certainly placed in this family; in 
which we may also probably include Siratodus, of the Upper Cre- 
taceous of both Europe and North America. An allied type, fiom 
the Lower Miocene of Belgium, has been described under the [m- 
occupied name of Amphodon. 

Family Saurocephalid* (Saurodontid<e). — The last family of 
the Saurodonts are laterally compressed Fishes, in which the maTrilltf 
and premaxillae are large, and carry powerful teeth, which are usually 
implanted in distinct sockets. The dentary bone of the mandible 
has but a single row of similar teeth, and there are no teeth on the 
palatines and ectopterygoids. The vertebrae, with the excepticm of 
those of the cervical region, carry two deep grooves and pits wi 
their lateral surfaces. The type genus SaurocepAalus is represented 
in the Cretaceous of North AJnerica, and also by a single species in 
the topmost Cretaceous of Maastricht, in Holland ; the teeth are 
subequal and closely approximated. Icktkyodeetes and PoHheus 
(fig. 937) also occur in the Upper Cretaceous of both Europe and 
North America ; the latter, which extends down to the Gault, attain- 
ing large dimensions, and being characterised by the great crest in 

the lypi'cit American one wiih /fhthyodtda. Saurodon 
the Upper Orccnsand of New Jeisey ; wbU« wn may 


~n* aBttrier poni^i «f ihe ihctttM of Ptrtktitt mtUaui : Fnm ihc CnUMOut 
afHotthABvtcb Grail)- n.hKvd, (AfWr Copt.) 

Ily place in this faintly the impeifcctly known Tom^gnaiAus, 

;lish Chnlk, in which the teeth were anchyloscd to ihc 

at sockets, and had subc)-lindrical crowns enamelled 

sbalili: Saurodonl, of which the family position cannot be 
niay be mentioned Giganlicfifhys {Titttnichtfryi), founded 
of very large sim, from Ihe Cretaceous of Eg>'pt. 
GoNOR)iVNCtlu*«. — Wc may now briefly niciuion five 
jittjng families related to the Salmonoid^ and Clupeoids. 
I of these is now Feprescnted only by a single species of 
w^inu found on the coasts of South Mricii, Australia, aiid 
JCKacly allied, however, is A'otcgtyiKUS, from Ihc Eocene of 
|, which is mainly distinjjuished by its di>niition. 

OsTEOCiXisslDyE. — Thc second family is now represented 
^mm of South Africa, Sumatra, and Queensland ; the 
llimt of the Brazilian rivers ; and HeUrotU of several of 
iith Africa : thus nresenline a sirikins examtilc of discon- 


that it should not rather be placed in the allied Americsn 

Family Esocid£. — ^The£iC(7</«arenow represented bytbel 
(EsQx) of the rivers of the Northern Hemisphere. They are i 
tensed by the margins of the upper jaws being formed by ^ | 
maxiltse and maxillae ; by the presence of small conical teeth i 
palate ; and the absence of an adipose 6n, and the positi<»i of] 
dorsal in the hinder part of the body. Species of Ea>x \ 
the Upper Miocene of CEningen in Switzerland, and the Pie 
of Silesia ; while the extinct Sphenolepis, of the Upper 
Paris and the Eocene of Aix in Provence, characterised hfj 
wedge-shaped scales, is considered to be allied. Recent writeni 
place in this family the marine Istiaus of the Chalk of We 
and the Lebanon, in which there is a long dorsal fin occupyiif ' 
greater part of the back. 

Family ScoMBRESOCiDiE. — The members of this family 
inhabit tropical and temperate seas, and are best known by the i 
pike {Belone) and the Flying-fish {Exocaius). The jaws are I 
as in the last family ; the dorsal fin is placed above the anal in fli 
caudal region, and there is no adipose fin. Belone is found in tk 
Miocene of Licata, in Sicily ; while Hohsteus of the Middle Eooa 
of Monte Boica is an allied genus. The living genus Ext)eatm\ 
characterised by the enormous development of the pectoral fins,ai 
was preceded in the Chalk of the Lebanon by the nearly relatt 

Family Cyprinodontid,e. — The Cyprinodonts are mostly smil 
carp-like fishes, inhabiting the fresh, brackish, or salt waters of 
considerable part of the world; and readily characterised by the {«> 
sence of scales on the head, and the absence of barbels. Spedi 
of the type genus Cyprinodon {Lxbias) occur in the Middle ar 
Lower Miocene of the Continent, while the CEningen beds hai 
>nelded a species referred to the South American genus Pted/ia, 
comparatively large Cyprinodont has been described from the PB 
cene of India. 

Family Cyprinid.e. — The important family of Carps is ve 
numerously represented in the freshwaters of the Old World ai 
North America. There are no scales on the head ; the mai^ 
the upper jaw is formed by the premaxillse ; there are no teeth 
the jaws ; there is no adipose fin ; the lower pharyngeal bones cai 
one or more rows of teeth ; and the mouth frequently has barbe 
The body is more or less compressed, and is often comparativf 
deep. In the Miocene of the Continent we have representatives 
the following genera now living in Europe — viz., Cyprinus (Caq 
Gohio ((Judgeon), Ltuciscus (Roach and Dace), Tinea (Tencl 
Rhodeus, which is mainly Asiatic, Aspius, and Cobttis. Acanthopi 

^^^^^^^^^■IBiua) miiuu luc piuuuuijr uiuic ur less 

to those now inhabiting the lianic region. 

ScopcUDAL — The Scopeloids are marine fishes allied lo 

(with «hich tbey agree in the structure of the jaws), in 

tx»dy may he naked, and there are neither barbels nor 

They are represented in past epochs by Htmhaurida^ 

lUK of I&trio, which it allied to the living Saurui, and 

r'ut and Anaf^rus from the Miocene of Sicily, of which 
is related to Parairph now found in the «ame region. 

Sit.cttiD,e. — The Siluroids or Cat-hshcs form a large 

fresihwatCT fiihcs of not less importance than the Carps, 

all temperate and tropical regions, and in some ca»cs 

,tbe sea. The skin is cither naked or covered with bony 

■re alwaj-s harl>cls, which freciucntly have a bony axis ; 

of the upper jaw is formed by the picinaxilla; ; there 

liar ; and there may or may not be an adipnite fin. 

of the Siluroids is often remarkable for the great develop- 

the supmocctpital (fig. 93S), and the presence of derninl 

MM in the region of the neck, which spread over the nape, 

ulate with the bones of the secondary pectoral girdle. The 

second intcrspinous bonra of the neck frctjucntly also 

large bony buckler, bdiind which ihe long dorsal spine 

by means of a ring with the first interspinal ; and this 

be fixed in an upright position by a curiou;; mechanism 

witJi the secortd interspinal. 'ITic " helmet " of the nape 

poniinuous wiih the "Injckler," and these hones, together 

kc of the cranium proper, arc frequently ornamented with a 

Kulpture. The pectoral fins frequently carry a spine as 

that of the do«aL The>-ngeal teeth arc generally 

points in which the Siluroids resemble the Plncoder- 
idi, I'rofcssof Huxley has suggested that we may rc- 

falAIAfl rn ih* «n^*ctnm nf iKft ^vicH'nT 



Eoceoe of Barroc we hzve SC^zoids icfened to tbe existing ' 
genos Jrinj ; uie pec^iliir. sixneviui bean-shaped, otoliths ( 
dihfs beinz doc cscomsoa ia the Bancn beds. It is inti 
to notice the aasociuioa in tbe English Eocene of ^uroii 
Oocodiles aruf Trknvchocii Ovlocustts ; the three grot^ 
found together a: the present dij io India and Airica. The 
of Smnaoa ais yielded renuins of exdoct species of th( 
Oriental genen PstrnJctr^miu and Sa^rini,- the last-oamec 
being also represen^d in the Ptiooene Sivaliks of India 
gigantic B. Yarrtiii^ now inhabit 
Ganges. The same beds have also 
renuins of Ciarimi, now found in b 
Oriental and Ethiopian regions^ crf'a 
belonging to tbe genus JSe ^ r^ramt 
93S). wbicfa is now ccxifined to tt 
and not improbably also of Ckrysiti 
trofHcal Africa. The »^«ring Ji 
aor of the Indian and Biinnese rii 
also left its remains in tbe Siwaliks 
tbe cluuacierisdc Oriental genus 
likewise rqiresmted in the same d 
Finally, of the widely-distributed 
Anus, which we bare already me 
ftom the English Eocene, there is e 
of two Siwalik species ; one bein^ 
rently neariy allied to a laige exialii 
Afticin form. In the Eocene of 
America there occurs the genus iUi 
which has \'omerine teeth, and i 
allied either to Ariia or to JH* 
from the same deposits have been referred 

Fi(. ajS.— Upw rieir of tbe 
llnill of HltmtTWJKkiu inUr- 
mtdiMi; fiam ike Si!*. Qm:- 
half ruiDT^ lire- m, Suprvxa- 
piul. *BAi. Fttjatii -ncailK*: 
ett, Eihmoid ; /r, Fnmial; tr, 
Orini ; n/. Sapnoccipilal: I- 


Tbe sculpture 

while spines 
latter genus. 

Family Mur.«:md^ — The last family that we have to nc 
this suborder is that of the Eels. In these fishes the body ii 
elongated, and either naked, or covered with rudimental scali 
toothed maxillae form part of the border of the upper ja 
there are no pelvic fins. Of the freshwater forms, Ai^idh 
ranges from the present date to the Chalk of the Lebanon, 
abundantly represented in the Miocene of CEningen and the 
Eocene of Monte Bolca. The Marine forms, or Congers, an 
sented by species of the existing genus Opktchthys in the 
Bolca beds ; and by the extinct Spha^branchtis of Uie latter d 
and Rhynchorhinm of the I^ndon Clay. Peculiar larval fo 
the type known as Leptocepha/t occur in the Continental Tei 
some of which are probably referable to this family. 



BCKMincM 7. AwACAXTHmi. — In this and the four rollowing su1>- 
fKn, icf ether forming the Physoclyrtti of Mime writers, thv snim- 
Iddcr (when pcesem) has no connection with the pharynx, and 
i pelt-ic fins arc nearly al«-2>'s thoracic or jugular in position — in 

ibtter case being in advance of the pectorals. The present sub- 
s is further characterised t)y the rays of the dorsal fin being soft 
jointed, and the pelvic fin never abdominaL 
iiLT Gadii^c — In the Cod family the body (fig. 939) is sym- 
■nd covered with smaJI scales ; there tna.y lie three dors:i1 

, tad the pelvic fins are jiij;ular. All the genera are marine, 
:the family i« unknown Wore the Eocene. The extinct AVww>- 
^yx and Paiao^pidiii have been described from the Eocene of 
■^; while in the London Clay of Slicppey we luve swedes 

H to GoAts (Cod), Mcrhidus (KaltL'), and Pkysis. Other 
^Bds occar in the Mio<:Gne of Hicily. 

■^ilULV Plbvkonectid£. — The Klal-fishcs arc characterised by 
le laiecal compression of the body (fig. 940), of which the 


fore part and the head are not bilaiLTilIy it)* 
with one side upwards and the other downward, the bcsd 
Twisted round so as to bring the two eyes upon that side ^ 
becomes the upper one, and which alone is coloured. Theft 
swim-bbdder, and the dori;.-!! and anal finK occupy almost the 4 
length of the body. Remains of a species of /^AoweAworT 
{fig. 940) ate fownd in the Middle Eocene of Monte Bote* 
those of a Jif/m (Sole) in the Miocene of \V'urteinbcrg. 

Suborder 3. Pkar^iigognatki. — In thi» ittnall subon 
portion of the rays of the dorsal, anal, and pelvic fins a fd 
by non-articuLated siiincs. The lower pharyngeal bones art tU 
and the &w ini-bladdcr has no ducL 

Famii-v Pomacextkiu*. — Of this large family the only Eon 
fosfti) representative is Odonffus from the Middle Eocene of I 
noica, which is allied lo the cxi.sting Ileliastcs : but Ptofciisor 
thinks that Priscaiara from the Kocene of North America 
perhaps be referable to it, although vomerine tccih arc preseoi 
the members of this family have ctenoid scalei. 

Family LAUKiD'fi — The Wrasses are a large family of I 
fixhes most abundant in tropical and temperate regions, and 
acicriacd by their cycloid scales, the single dunso] fin, the tb 
position of the pelvic fins, and the absence of teeth on the | 
Many of them have the lips greatly thickened ; and the phar] 
bones bear molariform teeth. The cxistitig genus Labms is rec 
from the Middle Bnccnc of Monte Doica and the Miocc 
Switzerland ; while SaurinkAthys of the Miocene of France a] 
allied to the living Odaana, Frctn 
of the Eocene of North America i 
ancestral form of Uie Black-fish (TUi 
of the same country. 

Family Pharvncodomud/E. — Am 
of extinct fishes more or less nearly 
lo the Wrasses, but differing in several j 
very markedly from that family are reg 
as forming a group by themselves. 
t)Tie genus Pharyn^dopUus (Numm 
tHs) occun; in the French Miocerte, a 
the Tcniary of the Canaries ; closely 
to which i.* Fhyllcdus from the Creta 
of Ocxniany and the London Clay of 
ITcy, derived leeth being also found f 
SuiTolk Crag. The pharyngeal teeth of Phyihdui (fig. 941 
remarkable for their thin and leaMike i>lructure, and oko It! 
rapid manner in which they arc succeeded from below by 
ones. The imperfectly known Egcrionia from the Lower E 

r nbwAi Irani tlie Ij>ii1iiii 



and PlatyUtmus from the Middle Eocene of Brack- 
may be prorisionally included in this family. 
litr Chromid^ — The Chromids are a family of iimall fresh- 
f'Fisbcs from Palestine, tropical Africa, and Aint;rica ; and alM 
by one genus in India. The scilcs arc usually ctenoid, 
liw is intcmipled, and the teeth of the jaws are very 
To this bmily ht referred the large genus Pyctuisterinx 
the Cretaceous of the Le1>anon ; and with lcs» certainty 
of the same deposits. 
DElt 4. AcAKTHOPTERVCii. — The Acflnihopteryjii form a 
series characterised by part of the ravA of the dorsal, anal, 
: fins beii^ non-articuUied and forming strong spiia-^. The 
[pharyngeal bones are usually separate; and there is no duct 
swim-bladder. Tlie scales are very generally ctenoid. 
ILV OniioCEfMAUUA. — The Qphioaphalida arc fnahwalcr 
almost confined to the OrientaJ region, in which the long 
^■nd body arc entirely covered with scales, and the dorsal and 
filK are loilg and devoid of spines. These fishes arc in the 
hbl of btirving themselves in the mud during drou(;his. The type 
|miK C^hiottpMalus is represented in the Pliocene of India by 
peoa clgoely allied to, if not identical with ^omc of those now 
|ilbil»g the &an>e area. 

' Fauuis FtSTUt-ARiiDX AND Cestrircii)^.— The first of these 
knfis comprises the marine " Flute-mouths," characterised by the 
Mf body and the production of the anterior bones of the skull 
Uoa bog tube, terminating in the mouth, and by the absence or 
MB kite of the scales. The existing genera Fislularia and Aula- 
has, Duw found on the bordcrn of the troi»cal Atlantic and Indian 
IM>m, are represented in the Eocene of Monte Uoica and Cilarus ; 
hSe AmJiteafs, now confined to the Pacific coast of North .^mcTic.^, 
Imtiul in the Eocene of Sumatra. Extinct genem from Monte 
Blcaaic Vroiphtn and Rhampfu}sui — the former characterised by 
Bwcdge-likc caudal fin, and the latter by a large spiny ray on the 
dk. A Fistularian, from the Lebanon Cretaceous, has been named 
ifaK^M/Awj, but the name is preoccupied. The Cetilriicidtr, which 
ree with the ftstulariida in the structure of the mouth, but difTcr 
the form of the body, arc known in a fossil state by a species of 
klivittg genus AmphhiU, from Monte Bolca. 
Bmilv ML'c:ii.injK. — ^'rhis and the two next fazniiies include 
B chatacterisied by the presence of l«'o distinct dorsal lins, of 
Bch the first is either low, or ha& weak spines ; and by the ab- 
■moal positioo of the pelvic (ins, which have ^ve rayii and one 
bie The Grey Mullets inhabit the coasts of tropical and tcm- 
rate s«as, and have cydoid scales with no lateral line. The recent 
Mu^l occurs in the Upper Eocene of Aix; and Calamo' 


plmrus, of the European Chalk, with which the North 
genus Syllamus is probably identical, may be placed 

Family ATHERiNiDiC — These fishes, which differ from 
Mullets by the presence of an indistinct lateral line and the] 
number of the vertebra, are represented in the Middle 
Monte Bolca by two minute species of the existing genus Ati 
and also by the extinct Mesogaster. 

Family Sphyr/enida:, — The Barracudas, in which the 
line is continuous and the vertebrae are not numerous, are 
at the present day only by Sphyrana, of which some spades 
a length of eight feet. That genus occurs in the Middle Eoccocl 
Monte Bolca, and has also been recorded from the Uppcf 
ccous of the Lebanon ; but it is probable that the latter fonn I 
to one of the Saurodont Phj-sostomi. Cladocyclus, from the U| 
Cretaceous of England and Brazil, is an extinct genus. 

Family Blenniid^g. — The Blennies form one of four families if 
but little palfeontological importance, in which there is a long danrf 
fin, which may be entirely spinous ; and the ventrals, if present, M 
either thoracic or jugular. It is probable that Pterygoaphaius, foM 
the Middle flocene of Monte Bolca, should be referred to tbt! 

Family Gobiid^. — The Gobies belong to another group of fl« 
suborder comprising two families, into the characters of whidi il 
will not be necessary to enter. They are represented in a foaa 
state by the existing genus Gobius, from the Monte Bolca Eocene 
and the extinct Chirotkrix, from the Upper Cretaceous of thi 

Family Dactyloptf.rid* {Cataphracti). — This and the nextsb 
families form a group of considerably more interest to the paheoo 
tologist than the preceding. They are collectively known as tb 
Cottoscombriform section, and are characterised by the presence o 
spines in at least one of the fins ; by the dorsal fins being eithe 
continuous or close together ; by the spinous dorsal, when present 
being short ; and when the latter is absent by the length of the sol 
dorsal. The pelvic fins are always jugal or thoracic In the preset 
family the body is cylindrical and elongate, with a coat of bon 
scutes ; the dentition is weak ; and the pelvic fins are thoradi 
Petalopteryx, from the Middle Eocene of Monte Bolca, is considere 
to be related to the existing Dactylopterus, in which the pectora 
are lengthened to an enormous extent. 

Family Cottid.*;. — This family is best known by the freshn-ab 
Bull-heads {Colfus) and the marine Gurnards {Trig/a), The bod 
i^ more or less oblong ; the dentition weak ; the dorsal fin is usual) 
divided, with the soft portion the larger ; and the pelvic fins ai 



rfc. aix) goictally have five soft raj-s. Cothts itself occun: in 
Upper Miocene of (Eninf^en ; while ihc Upper Eocene of Aix 
ibc extincl Lepidocattus, cli^m)^ishcd by its acnoid scales. 
oocun in the Europc-in Tertiaries. 

iiLv Traciiimd*. — This family comprises a numlwr of genera 

r«inill marine Fishts distributed over ihe greater pan of the world, 

i which the body is long and slender, and may or may nol have 

The dorul nn may be either single or divided, but its soft 

is always much longer than the spinous. To iht» family 

probahljr be referred the naked Callipreryx, from ihe Middle 

of Monte Boica; while TrtuAiiiopiij, of the Upper Tcrliar>- 

Sfwin. i< ronsidervd to be allied lo the existing TretMnus — a 

cLtiDLt genus bving Pitujvth'gimis, of the Sicilian Miocene. 
Fi^iLV SCOMBRIUX. — The Scom^riJit, typically represented by 
ckercb {S^m^r), but also compitsing other pcUgic Fishes, 
by the oblong and scarcely compressed body, ihc 
jpcd dentition, the two dorsal fins, the general presence of 
in (he poslcrioi pari of the body, and the thoracic position of 
I pelvic fins, which have one spine and five rays. In the Miocene 
[ Eocene of Europe, we meet with species of the existing genera 
(Mackerel), Thynnm (Tunny)! i»nd CyNum, one Lower 
species of die latter having been described under the 
of Sa>mi>avJfin, Curiously enough, the specialised genus 
■ (Sacking-fishes) is found in the Lower Eocene of CMariis. 
{S^i^amodtts) is an extinct genus from the Lower Eocene 
and the I-ower Miocene of Belgium, characterised by 
: development of its dentition, and iu single row of conical 
Iceth. Other cilinct genera are Paiimphyes and /surus, 
the T'ower Eocene of GlnriLS and Ortynus, from the Middle 
of Monte Boica. The exiting Pelamys bos been recorded 
the Lower Miocene of Belgium. 
Famiuy CoRVrH.¥:.Nit).e.— Another pelagic family allied lo the 
preceding t& represented by the well-known CoryphKnaa^ popularly 
knowD by the misnomer of Dolphins. The body is compressed; 
the toeth, if present, an* small and conical ; and there is a long un- 
divided dorsal 6n, without a distinct spinous portion. IIk existing 
genu Afrm {Gaitnxntmus) is found in the Middle Eocene of Monte 
Boloi ; while the family is also represented by the extinct Cetihgna- 
tims, of the Ijondon Clay. 

Kamilv C\TTlu.-E, — I'he Dories have the body very deep and 
iBiich compressed, with the dorsal fin divided, and its spinous por- 
tioa taller than the soft part, atvd the pelvic fins thoracic. 'Ihey 
ate repnnentcd by a species of the existing genus Zeus, in the Mio- 
c«»c of .Sicily, and also by the extinct Cytloides^ of the Lower Eocene 
of Glanis. 




Family Carancid.k. — Of gical intcrcsi to the paloeortoiogiitj 
the so-called Horse- Mackerels, on account of the beautiful 
lion of some uf the fossil forms in the Middle Eocene of 
Bolca. 'fhe body ik more or less compressed, and may be ob 
or deep, and cither with or witUout scales. The dorsal lin, 

Fig' ^^~-^*ftit/k*rMt TvlrtMnt : froai ihc Middle lieccAc ot MiHiM &al^ a, Ao^ 
C. Cauilkt ; u, DirTHl : r. FeckimI ; T, PeIvIc <iii> | 


may be tingle or divided, has the spinous smaller than the 
part ; and the pelvic fins, if present, are tlioracie. One of the I 
retnarlcahle extinct genera is Semiopharus (fig. 941), in whiclll 
dorsal fin is of enormous height, while the pelvic 6ru are* 
greatly produced. This genus is knoum only frona the 14 
Bolca Eocene. Other extinct Teniar>- genera are FfaidffBi 




Caranx {Carangopsis), Argyriosvs {Vomer), Liehia, and 
and Equula, from the Miocene of Sicily. 

Family Acronurid-e. — The last family of the Cot 
form section comprises tropical marine Fishes, popularly known I 
Surgeons, which are readily recognised by the sharp spine 
ing each side of the tail. The body is compressed, and oblong i 
ovate, with small scales ; the front of the jaws has cbisel-Iike < 
pointed teeth ; and the dorsal fin is undivided, with the spioawl 
portion less than the soft. This family is represented in a fodl 
state by species of the existing genera Naseus and AcaHt/ama bm,\ 
the Middle Ek)cene of Monte Bolca. 

Family TRicHiURiDiC. — The next section of the subaFdcrii| 
represented by the existing Scabbard-fishes and the extinct J'aJm- \ 
rhynchida. These Fishes are characterised by their elongated, ccid- ■ 
pressed, or band-like bodies, furnished with long dorsal and uil 
fins, of which the former may be divided into a number of fink)^ 
somewhat after the manner of the Ganoid genus Fofypterus. Al 
these Bshes are marine, and are found in tropical and subtFopkd 
seas. Species of the existing genus Lepidopus (Scabbard-lish) occ« 
in the Miocene of Sicily, while Hemitkyrsites and TViehiuritktkft, 
of the same deposits, are forms allied to the living Thyrsites uid 
Triehiurus, but differing by having the body scaled. Xiph^trjt 
is another extinct genus from the European Eocene ; while Am»- 
ckylum, of the Lower Eocene of Glarus, resembles Lepidopus excqit 
for the presence of some long rays in the pelvic fins, and the two 
are probably identical. 

Family Pal^orhynchid*. — The members of this family differ 
from the last by the production of the jaws into a long rostrum, 
which is either edentulous or provided with very small teeth. The 
dorsal fin occupies the whole length of the body, and the anal is 
also elongated and reaches nearly to the forked caudal. This 
family is known only by Hemtrhynckus from the Eocene of the Paris 
basin, and Pahforhynehus from the Lower Eocene of Glarus. 

Family Xiphiid*. — The Sword-fishes, which are of pelagic habits, 
and generally attain very large dimensions, are characterised by the 
production of the upper jaw into a long spear-like rostrum. They 
are represented at the present day by Xiphias (fig. 944), in which 
pelvic fins are wanting ; and Hisfiophorus, in which these organs are 
long and filiform, and the dorsal fin may be of great length and 
height. Fossil Sword-fishes from the London Clay have been 
referred to Histiophorus {Tetrapturus), although it is not certain 
that they may not prove generically distinct The genus CaJorkyn- 
chus, which was formerly referred to this family, is noticed among 
the Chimeroidei. 

Family Berycid/e. — The Berycidte are characterised by the pres- 



101 1 

of mucous-bearing cavities in the head, and by the pelvic fins 

(except io Morwcfntrii) a single spine, and more than five 

This bmil)- is one oC the oldest of the suborder, lieing nbun- 

tvpmoitcd in the Chalk. The Tossil gcner», which may he 

in olphabetioil order, are a» follows — viz., Acragaster, from 

Upper CreUoeoiu of Westphalia ; Bfrya^iis, from the CreU- 

oT England ; Seryx, in which there is one dorsal tin with 

tftiocs, is represented by a single species in tlie Kurojjcan 

,and by two species in the Chalk of the Lebanon, while it is 

at the present day; J/olotenirum, from the Middle 

of Monte Eolca, the Miocene of Malta, and tropical seas 

! present day; HamoHotui, fioni ihe Cretaceous of both Kng- 

ihc Lebanon ; Ho^opuryx (fij(. 9.(5), in which the spinous 

kA the donal fin is grently devclupcd, And there are aUo four 

Fif-fAj.— JVMIy«r<9< J/Bfxinui^.' Inim ib« Upper CluUlt o( &iuau. lUdaM)!. 
(Afta hLini*!!.} 

;wnes in advance of the nnal, is known from ihe Upper Cre- 
of twlh Europe and the I.«-b;inon ; Mynprislis, from the 
Middle Eocene of Monte Boica, and now found in tropicnl seas ; 
PristigrHytt from Monte BoIca ; Pseudoberyx, from the Chalk of the 
I^b.'tnon, characterised by the almost atidominal position of the 
pdvic 5ns ; SpAtiuxtphaJut, from the Upper Cretaceous of West- 



phalia ; and S/tnrs/i^ma, frnm the EngUsli Cbalk, of irhtdi 
affinity is somewhat doubtful. 

Pekcikorm Section. — The remaining faniiliea of this sub 
are characterised by their more or less compressed bodjr; bjr( 
dorsal fin, or 6n9, occupying the greater portion of the bock ; bqrt 
strong developinent of the spinous part of the dorsal fin, 
at least as long as tlic soft portion ; and by the sofl anaJ 
irg lo the soft donu). The [wlvic fin* are thoracic. 

Family Scorp.enid.^ — This family, which is allied to 
lowing, but h.ts villifomi tccih. is known in a fuiisil state only! 
species of the type genus Si6rf^na, from the Eocene of AJgcril. ; 

Family Sfakid*. — Tlic Sea-Breams resemble the 
which we shall notice immediatoly, in general appearance, bol 
mouth is cilhcr provided in from with chiscl-lilcc teeth, nr on 
sides with molariform ones. All of them inhabit tropical .indt 
pcrate seas. Among existing genera the sphnerciidal palatal teeth* 
Chrysof<hrys occur in the Red Crag of Suffolk, the Miocene 
Malta, and in beds in the Canaries, which are probably refenbkl 
the same epoch ; while Sarg;Hi is recorded from the Miocene 
France and Wiirtembcrg, the Upper Eocene of Algeria, and 
older Tertiarj- of New Zealand ; and Pa^llus occurs in (he Chafti 
the Lebanon. It appc.its, moreover, that many of the 
palatal teeth of Chrn<*phrys have been described as SfiMarodus, ■ 
the anterior cutting-tccth of the same genus, together with 
gcal teeth of some of the Carps, have been described as Capita 

Re. 9fS. — Skeltuo oiX/'tt»nltu tnicriuati*--.!; frefn ilw MUdl< Hooawt 
Munic Rain. Jtcdocnl 

CuttinR-tceth of this type, described under the latter name, ocair ■ 
the Miocene of Austria and Silesia, the Pliocene of Italy, and the 
Eocene of Northern India. Sf'amodui {fig. 946) is an e^ttinct genus 
from the Midcilc Kocene of Monte Boica, while the name Stepkam- 
adut has been applied to a genus from the Upper Cludk of the 
Sahara, characterised Viy the brtvidlh -ind denticulated edges of Ibe 
cutting-teeth. It may also be obsen-ed that teeth from the European 




jhe» have been described tinder ihe names of Sargodou, Sorui' 
and Atima. 
fKutuY Chxtodontida. — The Coral-fishes, or Chatodotils, 
from the Pcrcoids in the greater vertical depth of the body, 
' llie CDOtinmtion of the scales over the median firu, aiid nUo by 
Utcral tine Mopping »hon of the caudal fin. 'l*heir teeth are 
timtle-lilte. These marine tropica) fishes (which are generally de- 
pcnbetl under the name of S^tiatttificnnes) are remarkahle for the 
fcnmne govgeou»Re»s of their colouring, and are of cotiiparattvely 
ItanQ aui& They are represented in the Middle £ocen<: of Monte 
Bofea by tbe existing genera Holatanthus, Pomacanthus, Efikippns, 
^dmttpkagm, and T^xota ; the biter being now confined to the 
C)rieni:il and Australian reyions. The earliest Chaitodont is Plaiy- 
porwmt, of the Upper Cretaceous of U'cslphalia. 

F.i«ii.> Pkkcii>:C — With this and the next family of highly 
ppecialisefl >lshes we cotnc to the end of the existing representa- 
•nes of ihc present subonJcr. These Fishes arc characterised by 
[Ae eootinuous lateral line (fig. 947), the general absence of scales 
[from the median fins, the conical teeth, and ihc ab»«nce of barbcb. 
Tbey are all cimivorotis, and inhabit the frc<;hwntcr$ and couts of 

^B liQpical and temperate regions. The existing genus Perfa (Perch) 

OXun to the Upper Miocene of (Eningen ; while in the Upper 

Eowoe of .\ix we have the allied but extinct J'araperca. In the 

MjJdIr Eocene of Monte Boica, we find species referable to ihe 

outiflg genera Lahrax (Bass) ; Lalti, now inhabiting the Nile and 

Uages ; Duks, of the Indo-Pncific ; Semtnus (Sea-Fcrch) ; Af^sm^ 

of the Kediteirancan and Atlantic ; and also Tfumpon, nf the Indo- 

Plcific The extinct Cr(h/^rm and Smtrdis (fig. 948) likuwiae 

otcor in Ihe same deposits ; the latter being also fwind at Aix 

lad in the Mmcene of Wiirtcmbcrg. Atanus and Podoey't, from 

the I^orer Eoosie of Glartis, are memljcrs of this faJiiily, which 

Wre formerly refcned to the Beryeidie. \\\ ihc Eocene of North 

Aawica we have Afiof'/otus, presenting characters cynimoii to 

nc-MT— TbCoaBMpeKh(/'/tT«>r>t«'<Mi'tfi). RtOunO. 


Aitdstrodon, founded upon detached teeth from the Qulkj 
France ; similar teeth also occurring in the Cretaceous of Tdl 
These teeth are generally regarded as pharyngeal teeth of TdeoM 
although it has been suggested that they may prove to be 
teeth of Pycnodont Ganoids. 


1. Agassiz (L.)— "Recherches sur les Poissons Fossiles." NeacU 


2. "Monographie des Poissons fossiles du Vieux Gics Roa| 

Neuchatel. 1 844. 

3. Cope (E. D.) — " The Vertebrata of the Cretaceous Formatiaai 

the West." 'Rep. U.S. GeoL Survey of the Tenitories,' vot 


4. Davis (J. W.)— " On the Fossil Fishes of the Carboniferous Lii 

stone Series of Great Britain." ' Trans. Roy. Dublin Soc' 
vol. i. (1887). 

S- " The Fossil Fishes of the Chalk of Mount Lebanon in Syii 

Ibid.,yo\. iii. (1887). 

6. Egerton {P. DE M.)— "The Nomenclature of the Fossil Chimaa 

Fishes." ' Quart. Joum. GeoL Soc.' vol iii. (1847). 

7. GOnther (A.)— "Description of Ceratodus." ' PhiL Trans.' tl 

8. " An Introduction to the Study of Fishes." Edinbuirh. il 

9. Hancock (A.) and Atthev (J.)--" On Diptenis and Ctenodi 

'Ann. and Mag. NaL Hist," ser. 4, vol. viL (1871). 
la Hasse (C.) — " Das Naturliche System des Elasmobranchier 1 
Grundlage des Baues und der Entwickelung ihrer Wirbelsauli 
Jena. 1879. 

1 1. Huxley (T. H.) — " Essay upon the Systematic Arrangement of 

Fishes of the Devonian Epoch." ' Mem. GeoL Survey of Gi 
Britain.' Decade X. (i 861). 

12. "Structure of Crossopterygian Ganoids." Ibid. Decade) 


13. " Stniclure of Ceratodus." ' Proc. Zool. Soc' 1876, 

14. Koken (E.)— " Uebcr Pleuracantkus." 'Sitz. Ges. Nat Ber 

1889. Na 3. 

15. MlALL {C. J.)— "On the Genus Ceratodus, with Special Refi^rt 

to the Fossil Teeth found at Malcdi, Central India." ' Palieoi 
logia Indica' (Mem. Gcol. Surv. Ind.), ser. 4. vol. i. {1878). 

16. ■ — — "Sirenoid and Crossopterygian Ganoids." ' Palieontograpb 

Society.' 1878. 

17. Newberry (J.)~" Palaeontology of Ohio" (Carboniferous and 

vonian Fishes). 1873 and 1875, 

18. Newton {E. T.)— " The Chimasroid Fishes of the British CreUci 

Rocks," ' Mem. Geol. Sur%'ey of Great Britain.' Monograph 

19. Owen (R.) — "Odontography." London. 1840-45, 

20. Pander (C. H.>— " Die I'lacodermen." 1857. 

21. " Die Ctenodipterinen des Devonischen Systems." 1858, 

22. " Die Saurodipterinen, Dendrodonten, Glyptolepiden, 

Cheirolcpiden des Devonischen Systems." 186a 






Sit {C H.)— " Fossilen Kisclie dcs Silurischcn Systems." ifis**- 
B (I.) AND LanklsteriE. R.)—" Monograph of ihe Fi*he» 
ibc OM Red Sandstone of Untain (CcfiJia/iixpiaay ' PaLconio- 
phical Swwiy.' 1868-7* 

CAiR (R. H.>— " pescription of Pygcpicntt Crteiwchii? &c 
Trans. Roy. Soc. Edinb.,' vol. niv. (iSfi?). 

" Mooogiaph of the Gaaoid Fi^cs of the Briii^h CArboniferous 
'onDUions l.Palaam'sa'dir).'' ' ralzrontographical Society.' 1877, 
On the Genus Dipterus' 4c ' Ann. and Mag. Nat Hisi.,* 
S, «oL iL <i878). 

On the Stniciure and Affiiiiiiei of Triitiih>pierus ahtut." 
Trani- Roy, Soc Edinb,' vol, xm ii. ( 1875). 

— "Oo the CraaUl Osicolog>- of RhUodopiii* Ibiti., vol xxx. 

'** Report on Foi?ul Fishes colleclcd in Eskdalc and Ltddes- 

" " PL I. Ciinoicki. /*»«;, vol. x»]t. (18S1). 

** Strocture and Syitcin.iiic Position of CluiroUfiis." Also 

'Oa Some Fouil Ktihcs from the Neighbourhood of Edinbuigh." 

* AniL and M»g. Nat Ilin.,' xr. 4. >'ol. xv. Cid75). 

-^ " On the Structure and Affinities of ihc Platysonuda." JMd., 

*t>l xxvL. iiSto). 
^— " On the NontencUture of ilie Fi»he« of ilte Old Red Sanditone 

oTGrvat ltni»in." ' Gei>L Mai;.; dixad. 3, vol. v. (1S8S). 
^ - " On Ibe .Structure and Clas&ilication of the AsUrolepida^ 

- Ann- Mag. Nat. Hi*t.' ser. 6, vol iL (iSSS). 
" /AMHMuJnu coDopared with Cea-osltus." ' Geol. Mag.,' decad. 3, 

*ot vi (1889)- 
WmrCAVES (J. F.>— ■• lllujiraiions of the Fossil Fiihe* of the De- 
vonian Rocl» of Canad.!." Pta. 1. and IJ. 'Trans. Roy. Soc 
Canada,' (oU. iv. and vi. (18S7 and 1SS9). 
L Woodward (A. S.)— *' On some New Species of Hc^ocentrvm, from 
the Miocene of Malta, with a List of Fossil lierycidtr." ' (Jcol. 
Mag..' decad. 3. voL iv. (tlUi7X 

"On Soni« Remains of the Extinct Selachian Asttranmlhus. 

from the Oxford Clay ci Peteiborough." 'Ann. Mag. Nat. Hist.,' 
nrr. & vol ii. ( r888>. 

—— ""On lh« genus A\'i>limniu" 

" Syivopst* of the Vertebrate Fossils of the English Chalk." 

■ Proc. GtxA. A»«oc.' vol. t. (1887^ 

"CatatOftue of Fossil Fishes in rhe British Museum.'' Pt. I. 

Ebumobninchei. 81,-0, London (i8!t9l. 

- " On the Occurrence of Onychajus in the Lower Old Red of 
Herefotdshire." ' Geol. Mag.,' decad. 3, vol. v. (iKXS). 

. -On Ihe Paljconlology dif Sturgeons." 'Proc. Geol. Assoc.,' 

ZTTTkl ( K. a.!—" Handboch der Pafaeontolo^c." Abth. !., vol. iii., 

Sts. 1.:. Svo, Munich (1887-88)- Critically reviewed by A. .Smiih- 
roodvrard in ili« ' Geol. Mag.' for 1889. 

— — " On the Fossil Fish-spines named Ctxlorhymhus^ Hid. 
** On the genus fiJiMPlfpit" 

Proc. Zool. .Soc." 1S87. 
' Geol. Mat;.,' decad. 3, 

vol. iii. 





The Amphibia, which comprise the Frogs and Toads, SaUr 
Csecilians, and the extinct I^byrinthodonts, and are 
known as the Batrachia, agree in so many points of structure ' 
more generalised Fishes, that Professor Huxley groups the two < 
together under the common name of Ichthyopsida. Recent ( 
tions on fossil forms, tend, however, to show in the other ■ 
a transition from the Amphibia to the more generalised 
The Amphibia as a whole differ from Fishes mainly by the ' 
stance that when median fins are present they are devoid of I 
rays ; and that the limbs, when present, contain the same : 
elements as those of the higher classes. They agree with Fishai 
having branchise in their earlier stages of life, but these very f 
quently disappear in the adult, when respiration is carried on enticdr 
by means of the lungs. An epidermal exoskeleton is genen^ 
wanting. There is no amnion, and at best but an imperfect alUiADii 
in the embryo. In existing forms the cranium always artictiUtdlD 
the vertebral column by two distinct exoccipital condyles, but in a fat 
I-abyrinthodonts these were not ossified. The mandible articuUto 
to the cranium without the intervention of a suspensorium ; so Atf 
the skull, like that of the Dipnoid Fishes, is autostylic. A la^ pn* 
sphenoid is always present ; and cranial bones are largely developed 
although their number is generally less than in Fishes. The extttml 
nares are terminal in nearly all cases. The vertebral column i 
more or less completely ossified, and can generally be differentiate 
into cervical, dorso-lumbar, sacral, and caudal regions ; the saciun 
but rarely comprising more than a single vertebra. The infraneun 
segments of the vertebral column are frequently amphiccelous, an 
in recent forms each bears its own arch ; but in certain Labyrinthc 
donts, as we shall see below, the neural arches are carried by altei 



eliral sc^Trtnts and from circumstances to he detailed in 
ocl it n inferred )))' Professor Cope and othcni that the vertc- 
or tsiisting AmphilMans are realty inierrentra, currying 
airhcs which have been shifted to thcni fruiu tlic luat 
The ribs may articula[e to the veneUra by .1 )>in^le (uppt^r) 
process; but in Salamandtoids and I^byrinthaduni^ the 
alto cniry an inferior rib-facet, when the head* of the ribs 
[Uerttly double. As a rule no riblcss (lumbar) vcftcbnc 
and Ueriul ribk are wanting. The lilernum, which is never 
m Ftshcs, appears in the Amphibia in the middle line of 
and may be extended anteriorly as an omo- and cpi- 
CijC- 97+1 ")• '^^' pt^toral and pelvic girdles arc well 
; the former {t^id.) in recent forms ha\'ing, however, no 
or intcrclavick, although it is nearly certain thai these bone» 
mcd in ihc thoracic buckler of the IjibyrinihtKionts. A 
rod-like precoracoid {fig. 974, b) ocrurs in r«ccnl forms. In 
the pubJi b frequently uiKMsificd, and if ossified is much 
than the ischium. In most cases the ischium and the car- 
or lx>ny pubis of eiihi-r side unite tu form a continuous 
in which the obturator foramen is not represented, although 
■ bammll pettunition ; and the ischia meet in a ventral sjTnphy- 
The body of the iliura may be almost wholly in advance of the 
liiu. The limbs vaiy greatly in their proportionate length 
ifiSerent orders, and tnav be entirely absent ; while thccarpnis 
tu^aU may \k utiinssificd. '["he limb bones of ihc [.abyrinth- 
approximale 10 ihiiM; of the Anomodont Reptiles. The 
and taisiu alwa)-» have a centFale, and in soaie of the primil- 
Labpinthodonts there may b« at legist three ceniralia in the 
TI1C fourth arul fifth tarsalia always remain distinct from 
other in those forms which have five digits. The latter number 
(omd in the I^bjrinthodonis, but in some ci>iatin){ formti the 
,CBtt> Bjay \k TcdtKcd to three or two. The number of the 
fhbngeals in the d^its of penledactytale forms, counting from 
tic first to the lUth, docs not exceed 3, 2, 3, 3, 3, and thi.'i num- 
hcr may be reduced in M>tnc exiatin); forms. M a rule the tail 
ii romporaiivelj short. 

Teeth are usually present on the premaxilla, maxilla, vomer, and 
dv dentary bone of the mandible : but are generally wanting on 
the paUline and ptmyoid, although pre^-m on the latter, and also 
Qo the parasphenoid, in many Labyiintliodonts. These teeth arc 
onaUy anchylosed to the bone, and in existing forms are of simple 
ttmcturc. In the l.^hjiinthodonu the Atnictute of the leeth may, 
bovevcT, became very complex by foldings of the dentine, this 
KTUCtnic being an catrcmc development of that met with in ceruin 
CUMids; tnd in some of theiie form& the usually l;uge pulp-cavity 



may be greatly reduced. In existing forms there is gc 
exoslcclcton, and if scutes or scales are developed they arr 
in th« skin: but in the I^Ahninthodonts bony scutes were' 
generally present, although frequently restricted lo the venDal ; 
of the body. 

A marked, although not universal, feature in the elm it 
cKinge from a respirntion liy gilU lo one by lungx; this 
bdng accomiJiintcd by ulher structural alterations, and 
nutnrnorphusis. In some instances only cxtema] gills are 6cn 
which form a plume on cither side of the neck ; and it is ttiae] 
which persist in such forms as do not undergo a ractamc 
In those groups, however, in which a metamorphosis takes 
internal gills may be developed for a short period. 

'I'h.ii the Amphibia have taken their origin from primitnc] 
allied (o the Dipnoi and Ganoidei is pretty e%'ident. Eviij 


in F 










Pit UD-^Eolarnd ilEW of the umia niriiir< of ihe cnnium at FrvlHtrm. wiik ihr* 
I W h U e IjAyrtMkcaant bam >luutcJ ; trrm iha pMimsn of Bohcnb. A', Niul ; F, Fn 
Am. ParitOl: S.O, Snprunipilal : £.f. Epialicl S.T.. (iupraleopwal: 6j, S* 
P.t/. PoMfmtal: 711. Ja^kt: ^.r.'. P«iorhitBl : <m, 1V*bii(1U: M.S, Uwj 

afiinily mih the primitive Ganoids is indeed very clearly shown 
the so-called labyrinlhic structure of the teeth of nearly alt ihc 
Palieozoic Amphibia, since we find a similar typ« of dental simciure 
obtaining in many of the early Ganoids, and nowhere else in the 



ifumil kiogdom. The similarity in the structure of the 
column of the earlier Eksmohranch and Clanoid Fi»hc9 
that otf the Ijibyrimhodont Amphibians is also important 
pointing in the same direction, 
class may be divided inlo the four orders, I^abyrinlhodontia, 
Ecaudata, and Caudaix The ftnl is totally ektiiict, the 
is at prvsem unknown before the existing epoch, uhile we 
i> certiun record of the accurrcnce of the Llmd hcfore the 
and of the fourth previously to the Tertiary. 
I. Labvkimthouontia. — Since the name of this order is 
i orictly applicable to alt its members it has l>een proposed to 
jte the tcim Stcgoccj^mla ; but, as the same objection might 
t Hken to a Li^^ nunil>er of temu in use, such a cliangc seems 
Using, then, the labyrinihodontia as including the 
ala and Mtcrotauria of Mime wiiccra, its members may be 
by the following features. The body Ls more or less 
led, and funiished with a tail ; tlie skull has paired supraocci- 
(fig. 950, SO)t and its postero-loteral re^ons are roofed over 
iposiorbttal (P.f.o) anteriorly, and a supratempora! {S.T) poste- 
There i&, moreover, very generally an epiotic ' (■£■/);. 'he 
ariMts (icquenily have a bony ring in the sclerotic ; and there 'a a 


Ifa yi^VaMnl np«i of iki ibnclc bocUtr iA Aitim*J»a FmiarJi: fram llu liannijui 
dfimoti \w94iMa n»utal tin. n(. Inioctavick; '/■ Clavicle; !>u|inclivjcle, <>. 

pwicial fonunen. Palatine and vonTCrinc Iccth are very generally 
{xsenl. and the dentine of the teeth is frequently more or leii^ 
I-iUoi. or plicated, from tlie sides. The centra of the vcrtebrM, 
vbcfaare amphicoelous, may be imperfectly os^ifitd, and frequently 
nuia a notocbordal canal in the middle. Usually there is a buckler 
on the inferior surface of the thorax, consisting of one median, and 
tw lateral flattened hones, probably representing the intcicU^Hclc 
•ad dincles ; the relatiotw of these hones being shown in fig. 95 1. 

' Dr BMir regartb tliik bone as \\tt opisihctic ; Rnil alwi conudcn that the 
bat Wk tcnned the Mjiunoui U the utpntctrtpoiat, and nV« vtrtS. 



Posteriorly to this thoracic buckler an armour of dcnnal scud 
generally developed on the ventral surface of the body; and in \ 
cases this annour may cover the entire body, ihc form of the 
or scalas, then var)'ing considerably in the different regions. 
(Mired pent«daml.-ite limbs were tisunlly present. 

'n>c (itef^goids are always separated from one another id 
m«dt:in line. The p^.^lvt•;^ of the more typical forms is ren^a 
like that of the l*an<i3iiun;i n Aiiotnudoni Kepiiles, presenting 
same absence of an oUiinHor foramen. 

The paired suprnoecjpilal ossifications constitute a feature I 
elsewhere only amun^' the Ganoid Fishes ; and ihe frequent 
cation nf the artirular hone of the mandible is also a charicto'i 
found elsewhere in the class. In many cases the external 
of the loncb of the skull (fig. 953}, and of the thoracic buckla;1 

"Fit. f $1.— Laurd irkw 0/ iha ImptTrect Irft niniu nf ihi tn«ndihl( of r^iy^rmU iWa 'i rfi; 
fmiTi ilw La«W MwBioit of tncJix l~hi kiun inilk^c inuium oiiuli. (Aftrf HadtJ.) 

butOptured by a aeries of irtcgulni' grooves and rtdges ; tra%-cn«l IB 
the former instance by a number of mucous canals, l-'rom the 
occurrence of a similar sculpture in the Pariasaurian AnoMO- 
donts and the Crocodilians, Professor Secley regards the«!« gnwfi 
as directly descended from Labyrinthodonts ; and it is evident tU 
the passage from the I jbyrinthodonu lo the former group of Rtf 
tiles is almost a complete one. The gills (fig. 953) of the >-ogn| 
arc generally lost in the adult ; but in one group the external g9b 

I-ibyrintliodonis range in Europe generally from the CarbonifenxB 
to the Trias, and are especially abundant in the Permian ; but ooe 
genus {Rk'iKosaunfs) pcrsisU'd to the Lower Jurassic, In North 
.\nierica and Indi:i this order is abundantly represented in smta 
mainly representing the period from the Carboniferous to the Tria*. 
According to the views of Professor Cope and l>r Kritsch, this order 
is to be regarded as one presenting generalised characters, some of ' 
which approximate to those of modern .^nipliibin, white others are 

' li ihoulil be olaenrcil that In the fi£ure af the pelvii of Erjppt pvtn in ' 
Zitld's Tatximlutut^e,' abili. i.. vol. [it,, pt, ii., p. 364, fig, Jji, tbcbindctnd { 
ulthe Milium U mt.^tnkcn for I he pubu. 

ce^reRAL structuke and orders. 


I ; and wc may probably regard ihc Aiioniodnnt Reptiles 

[ taken their origin (mm a group closely allied to the l^aby- 

lt», if not actually from this order. As regards the !»ub- 

of Uie order there is still considerable unccriainty, and all 

risiotu mint conaequcntly Iw regarded as more or lc*» pro- 
r>r Fritsch has, indeed, proposed to range the familit-s 

Ur Mrries or sulKirdcni, according to the external contour of 
r and the nature of the venebral column ; thiti grouping 
>vision4lly adopted in the present work, with some cmcnda- 


tI>KR I. BRANCKiOS.vuiti,«. — In Iht^ isubordLT the external 

tee approadies of the modern Salainamiers. The ver- 

tve centra composed of a single piece, which retains traces 

otocbonl : the ribs are short and straight; and the neural 
dibted in the middle of each vertebra. The teeth are 
I itruciurc. and internal gills were developed in the young. 
Ihoriiics include the next two groups in the present sub- 
do the name of le'jiospondyli- 

» pRormio-viu*- — The ifraHfkUsaurida of Ur Fritsch, 
Doded on a name which is apparently a synonym, may be 
nown as the Prolrilomda. They are by the 

eadih of the ^uU (fig. 950), and the smooih teeth. Typi- 

e palatines, according to Dr Credner, are small irnnsversely 

id bones tying in the 
pan or the skulL, and 

BDg the maxillcc ; this 

nent Iwtng similar to 

tad in the Ecaudaix 
pe genus Protriton 
iMmrut or I'ieuron- 
found alHindamly in 
DUO of the Continent ; 

iSevelofxnent has been 

rked out by Ur Crcd- 

n the study of a large 
of specimens of all 
specimen shown 

1J3 exhibits the neatly 

' ddon of a small irvdividual, while the details of the era- 

Kture are exemplified in the greatly enlarged skull shown 


liat> to ihc obser^niions of Dr II. Credncr, u appcats thai 
llfesi spedmens known were 25 mm. in knjijth, in which stage 
turc was aquatic, and brcailica by k''l'^> which were supponcd 
airs ci^ arches. By the time they atuiincd a length uf (rum 6a 






of I'turilim 
'cimuii of fmoti. (Afttr 


to 70 mm., these larvae cast their gills, and became ur-breaAen; 
development bein^ thus analogous to that of the existing Salt 
The aaults measure from too to 160 mm. In the course of 1 
morphosis the skull decreases somewhat in width, and the 
buckler grows much more rapidly than the scapula and coraooi^i 
curiously enough, the pelvis shifts its position, and thus inc 
number of presacral vertebne from 20 to 26. In the larva 
side of the body is naked (fig. 953), but in the adult there is a 1 
armour of bony scutes on this aspect 

Other genera of which some may perhaps be included in 
family are Ampkibamus, from the Carboniferous of IllincMs ; 
from that of Ohio; Batrachiderpeton, from the Carbonif 
Britain ; Hykrptton, from that of Nova Scotia ; Damsoma^ 
the Permian of Bohemia ; and Sparodus, from the latter 
Batrackiderpeton is remarkable for the absence of maxillary ' 
and the clustering of the palalo-vomerine teeth ; ^arodus \ 
ing the latter feature, but retaining the maxillary teeth. The 
tines of the latter genus are splint-like bones interposed between 1 
vomers and maxillEe. 

Family ApateoniD:*. — In this family the skull (fig. 954) b'l 
angular and comparatively narrow, while the teeth are maiked 1 
small grooves at their summits. The type genus Apatam b : 
in the Carboniferous rocks of Germany, while the allied Melanerptl^^ 
is from the Permian of Bohemia. An enlarged view of the dcml 
aspect of the skull of the latter genus is shown in fig. 954, the reten- 
tion of the internal gills indicating that it belonged to an imtnannc 

Suborder 2. Aistofoda. — In this remarkable group the bodf 
has a snake-like form, with well-developed ribs, but probably wiA- 
out either pectoral or pelvic girdles or limbs. The teeth are not 
folded ; and Dr Fritsch considers that the external gills persisted 
throughout life. 

Family DoLicHosoMATiDit. — This family, which is equivalent to 
both the Plegothonidte and Molgopkida of Professor Cope, is repre- 
sented in the Carboniferous of Britain and the Permian of Bohonit 
by Doli€kosoma and Ophidcrpelon. In the former the skull is lonj 
and narrow, with no sculpture on the bones, and it is probable thit 
the body was entirely naked ; but in the latter the skull may haie 
been shorter, and there was an armour on both surfaces of the 
body, the scutes on the ventral side being long oat-like splintti 
while those on the back were rounded and shagreen-like. The 
ossified gill-supports were furnished with small enamel-like denO- 
cules. One of the species of Dolichosoma attained a length of about 
two feet Pkgethontia and Molgophis, from the Carboniferous ot 
Ohio, appear to be nearly related. Faiaosiren, from the Penniar 
of Bohemia, is a gigantic form provisionally included in this faraity 

h'iih the length is eslim-ited at fott)'-five feet ; whilt: AJfnodtnna 
tK ^iJttc deposits tnay inilicatc a distinct family. 

the many reseinblanct^ presented by the Dolichosomatida 
erirting Apoda, Dt Fritsch considers it prolublc that they 
tfa* fCgvded as nearly Klated lo the ancctitral forin,s from which 
fpvup ha£ been derived. 

>kLiEK J. MiCHtMAL'HiA. — Thc Labyrinth odonts included in 

i flidtonler resemble Lizards in outward appearance, and have 

of the Tcrtcbnc more or less elongated, and long, curved 

LT Urooordyud*. — This family — the Ntttridia of I^x)fes8o^ 
and U>e Pty&HiiJa of Professor Cope— -comprises stout, long- 





kjM— L'vpn tBT&e* «( lb* dtud of Mti»tr^t»n /•^ilUm: fna ihc Pmnun af I 
*■■ ^HH •MHal NW- A BnxKUai O^., win-wcipluili otbu Itiun ■■ in fia- 



Wed (iEffnis, in which the cpiolic cooiua of the sltull are much pro- 

' teed; the cranial bones arc pitted ; thc ncumi ^ines and chevrons 

rf Ibe caudAl rertcbra: are much dilated at their extremities and 

I pOxmed; and caudal ribs are wanting. The lyjie genus U^nh 

r iirhhti, as wcU as Ctrattrptton, occurs in the Carboniferous of 

and Ohio, aod also in the Permian of Kohemia ; one sjiecies 

lonner attaining a length of about twenty inches. Ltplerpeton 

ncteristie of the Carboniferous of Britain and Ohio ; other 

gen<.Ta fiom the latler deposits being J'tyenitis, (Esfoct/A 
j>fiiiima, .ind probably Sauropieura, 

I-'amilv LiMN^nrLTiD^. — In ihc one genus Zi««f//f/iwi, 

Permian of Bolicmia, ihe naked body is more elanyalcd and tfM 

shorter than in the preceding family; the skull being broail 

FIr. US'- SuUya ftaUU i (nun the Permian td BoAtHnia. *. Dndw mrftcc «kb na 
iDorctl on the Iffi udc ; and B, Rlihl tUenl view of lh( (IwktMi «l the cauM nfi«a. Sil 
iMitt I'nucb.) 

Frog-like, with smooth lx>ncs ; and the upper and lower pcoo 
of the caudal vcnebra* simple. The teelh arc small, with theii 
miLs cither smooth or folded. 



ii,v H^ix}Pt.e5i(>Mi>.«. — This famil)' is readily recognised hy 

mewhai lurrow head, the smooth cmnial bon«s, and the en- 

m of ihe whale body (fig. 955, a) in an armour of scutes. 

Buidal ribs (fig. 955) are w«ll developed. It comprises the 

HyUpitsion, StvUya (fig. 95s), Rtm^on, ai>d Or/hoci>Ua, from 

ferniJAn of RoheniU. Hyionomu! and Smi'/er/vton, from the 

tr.!iir^ruus of Nova Scotis, are imperfeclly known fornis vrhich 

-J the same bmlly, in which the name Hyhiwmidir 

^w. -v .iii'^pted. .Ml the species are of small size. 

Family MickobrackiU-C. — This family tntrludcs smalt slender 

with short pectoral limhs, Mrongly sculptur«U cranial bones, 

scutes coveiing the entire liody. The type genus MinvhraMs 

in the Pennian of Bohemia, and has a long nanow skull, 

speris are known. Another representative of this family i.s 

tttJuu, of the Corhoniferoiis of Ohio, characterised by its 

apamSed skull. Cttyttnus, of the same deposits, may be pro- 

iruU^ placed in this family. 

iBCnntx 4. L.^BVRiNTHOtiONTiA Vp.R.*. — Thc genera included 
Ifats group arc choraaerised by their Crocodtle-like bodies; the 
like centra of thc vcrtclMie, when ihcse arc fuUy ossified ; by 
Tertebral column beinj;, at least in the young, veiy generally of 
types ktfown as rhackUameus and emMomenms : by the teeth 
more or less folded ; and by thc outer surface of the skull 
iog a more or less strongly-marked .sculpture, frequently arcom- 
by the presence of the so-called mucous canals. Some 
ties divide this group into the Ttmncisfwndyii and Stent'- 
hit, according to the incomplete or complete ossification of the 
oenlnt ; but Dr Fritsth regards the whole series us con- 
a single group — a view which is >u|)ported by the cir- 
e that in many of tho« form* in which the vertcbrfc are 
OSBided in thc adult, in thc yuung sdge ihcir ossification is 
Before proceeding further it will tie advisable to briefly consider 
luiure of the above-mcnlioncd ly-pes of vertebral structure. In 
genera like Dfphspotidyiut^ (DipliWtrtebron) and Crkotus 
OiMliil vertebra consists of an anterior centnim carr)-ing the 
aich, and a posterior intercentruni In which thc chevrons are 
led. Thew iniercentra, according to the views of I'rofessor Cope, 
id with ihc chevron - ((earing interrcnlra of C/r/iydra/'s 
the Anomodoot Repiilia, and the wedge -bones of .S/>Ae/t- 
among the Rhynchocephallans ; this type of structure being 
as the e/nffffiomfrous. In the trunk vertebrae of other genera 
TrimerarhiKAis (fig, 957) and ArfAegosaurus each vertebra' (fig. 

* Thi) naoM bki been yv)^*e^ in livw oftbe hjlrid i>iftntrt4ltvH. 




956} consists of four portions — viz., a basal intercentrum (h 
cuiinim), a pair of pleuroccntra, and a neunl arch. In this 
ihfleiHous tyjx: Profcaor Cope regards the plcuroccntra as 

suiiting the centnim of the 
/i Iwlomcrous type, since they 
carry the arch ; and as he 
that the functional centra in ■ 
fomis like Cfiffydosaitnu, 
entiy correspond 10 the inter 
of Are/ugasaunu, while the pi 
centra are small and Ti\ 
about lo disappear, it is 
that in other Ainphihia tlic 
, ™ .... centra arc totally wantiiur, 

vMcliia; fipom ih. ifoni tnA )A .Ltie. i the ^'crtcbral bodics, which in 
•(HO., wfteffriuch.) caudal region have the chc 

united to them, are really inter 
centra, to which the neural arches have been shifted. ProresHX I 
Cope regnrds Ihe rhachitoitious and embolomerous structures as char- 1 
actcrs of at least family value ; but Dr Frilsch considers that tbc two] 
types occur in diCTerent regions of one and the same species, as ' 

-■■M-' 'A 


Fig. ^i;.— Pi.ii> al •liiill anil wtatnal onjuinn ef Tiimrrvrlkai Ui imrigmlm! frtm At J^itm'mt 
of Hortli Amciici. a. Ban- and •xocdplult: t, t, lAinl anil pgMcrior new at aula ^ 
MMdiblv: </. r. Ponioni of vanabral column itfnmid b/prMtun; i, iMueuiu*:/, n u ita 
ecotn. (Aftcf Cope.) 

know to 1k! the case in some of the Ganoid Fishes (sufirot p. 959), 
where we find in Eutycarm*i! and Aipid'trhynchus a rhachiiomous 
type of vertcbnc in the cervical and dorsal regions, ortd an eml»l- 



«)cr-:u>. \yi>e in the t^udal. An excellent enrapic of the thachitom- 
1< of vertebra is ^own jn fig. 960, the pkuroccntra bcinj; 

hould be obMSved thai this intorprctation of the homology or 
inhitomous vcndxa is not accc|)lcd hy Mr Hulke. 
'iL^ ABCHeflOSAtJRin.e. — The well known genus Arefte^ 
J. ranging in Europe from ihc Cirtioniferous to the Permiiin, 
w uken oji the type of a family, which for the present niay 
mo^t of tboK forms in which Uie dorsal \-eTtebn£ are of the 

I.— upper orbfT «r <ht «ian 

'^•cAMMsnu; (rcB the C>t- 

Bcdw-xl. /'".v. Pninu- 

~Jtf>. MuIIb: St. Naul. L». 

, pMiiokl : PiF. pDUftiMul ; 
>kt, pMdiU>j -.7. jBB]«d- 
iiHM«it: ^. fa— ra wJ : ST, Supra- 
I I ■• ^.E3Mk::ir(l,S>|inoedp. 

rialH Otpw ^Rrft^: rrarr ih« 
Pcnnhaof Konli Aoitrica. R«- 

LliMie; fi, /«. f*, CnintU: 

rbAchiiotnoos type throughout life. Its members have, indeed, 
been iplil up into ihc Melosimrida, Trimtrorhtukida, and Erya- 
fida ; Imt the observations of Dr Fritsch indicate that for the 
pRKDt at least such divisions are not definable. According to the 
ImI iwmrrl auihoriiy this family may be choiacterised as including 
t^byrinthodants of medium dimensions ; having cytintlrical teeth 
of raxying size, in which the folding of the dentine is cumpamtivcly 
ftlj)[bt ; the upper sur£ice of the skull beit^ pitied ; the supraocdp- 
itati ridged ; and the tninlc v-ertebiic rhachilomouK, and the caudal 
asuaiDy crabolomcfous. A ring of l>oncs is usually developed in the 
cclerotic; the rentral surface of the body is always covered with 




scutes ; and the pabttnes form long Bplims lying on the 
of the muilUc. This famil)- is ct-idcntl}- the most pcii 
of the entire order; the vertebral column displaying an 
development at a stage which is iram.itional in higher for 
funher remarkable for the small size of the coraooid, in whic 
thi^ group agrees with the Salamanders and some of the 
dont Reptiles, The huriterus has distinct condyles, as in 
group ; and in (he pc)%-is the pubiK is ossified, and, like lliat < 
Anomodonis, unites with the ischium without the intervemii 
an obturator foramen. The tarsus and carpus {6g. 95^ 



rig. gfio.— If ri luitf-i! upEiit uT ilic irch ind tntMMnlnitn, and pcMahv 
ufadoni*! vctEcbra of AK^^fviifwrHj JCivAei; ftan iIh I.ow>tr P«ni>ijin«r 
ii)iuiE. irith U'rrd ci|uniiiini, a/; >, Suture Mvttn apine »ni atdb; *-m, _ . . 
upih)«l>: J, Tamaitna fraeut : <, Rib-fMai; t.r, N«uml caiul! i.t, InMccmm 
tram Gauidry.) 

characterised by the number of ceniralia, there being, aooc 
to the interpretation here followed, four of these bones ii 
fonner and ihree in the latter ; and the first centrale articu 
respectively with the tibia in the tarsus and with the radi 
the carpus. This ty[xr of taival and carpal structure is e^ 
the niDiit primitive yet observed, f 

Iti Arrhf^stiurus, and also in /.y/^saurus of the Kurope&n 
niian, and Trimtrorhathii of the reputed equivalent strata 
America, the occipital condyles were not ossified ; but tl 

a* f<irawii at HaA hmtna. Oiw-lulT laonl tin. (Aftn Cope ) 

fentian of Dresdfn, but it is preoccupied in the fiaurnpterygia. 
Po rtMH Bi of a vcriclira of £itcfiir^saitms, from Autun in Kriuice, are 
dunra in Ag. 960, (lie neural spine being remarkable for the great 
hScnl cxputeion of its suDimil : according to l>r Fritsch's restora- 
C^- 95<^) tbc ])lfurix'cntra bclongiiig to tlii^ vcriclita would be 
antcnoT itd<, but I>r Zittel would mther regnrd those on the 
ispccl 33 rcfcTal>lc to this segment of the colmnn. In 
Bi)ori lUge of the Lower Condwanos of IndiA thJi; group is 


represented by Gondwattosaurus ; while, if we may judge bfJ 
detached intercentnim which may belong to it, Rhytidostats, of i 
Karoo system of South Africa, should also find a place here. II 
American Permian, in addition to Trimerorhaehis, has also yidd 
Zatrachys, Eryops^ Acheloma, and Anisodexis. Rrygps includes w 
large species, in which the nares are widely separated and i 
placed at the extremity of the snout, and the thoracic plates : 
not sculptured. 

Family Diplospondylid*. — This family* is proposed by 
Fritsch for the genus Diphspondylus, from the Permian of Bohea 
characterised by the embolomerous structure of the entire ve 
bral column, and the absence of pits on the skull. Cricotus ( 
961), from the Permian of Illinois and Texas, appears to be 
allied form, which Professor Cope makes the type of the- fan 

Finally, it may be mentioned here that the genus lehtkyacamil 
from the Carboniferous of Ohio, which is frequently placed in 
Microsauria, is described as having rhachitomous vertebre. 

Family Nyraniid*. — The genus Nyrania, from the Permian 
Bohemia, of which the skull is shown on an enlarged scale in fi 
824, 825 (pp. go2, 903), differs from Archegostmrus in that the pi 
tines, in place of forming splints on the inner side of the maxil 
are situated near the middle line, internally to the vomers a 
pterygoids, and would therefore seem to represent a distinct fami 
This arrangement of the bones of the palate is similar to d 
obtaining in the existing Caudata. Some of the genera notk 
among the Arcfugosaurida, in which the skull is unknown, n 
belong to this family. 

Family Dendrerpetid*. — This family, which may be takai 
include the Brachiopina of Professor Miall, contains several gent 
of which the precise serial position and full affinities are at presi 
somewhat uncertain. The skull is parabolic, and marked by d( 
pits ; the parasphenoid in the type genus has a short stem ; and 1 
teeth have irregular foldings at the base. The vertebrae were ft 
ossified, and may have been of the embolomerous type. The t; 
genus Dendrerpeton is a medium-sized form occurring in the ( 
boniferous of Nova Scotia and the Permian of Bohemia, and cl 
acterised by the orbits being placed near the centre of the sk 
Another group of genera, constituting the above-mentioned Bracl 
pina, appears to agree so closely with the type in cranial characti 
that it may at least provisionally be included in the same fam 
The orbits are generally placed somewhat anteriorly. This gn 
comprises Srachyops, known by a single skull from the Mangli st 
of the Upper Gondwana .system of India ; Microphelis {Petrophryx 
^ DiphotruMda, see note, p. 1027. 

he Kjitoo system of South Africa ; Bothria^t, fiom the Lower 
wc of Australia; and Rkinosaurui, from the Jurassic of the 
Mountains. Tbcre is frequently a fyra ar\ the skull, hut this 

tmting in Micrvphofis. as in Dendrrrptton. It is pioltalile that 
Xtt^ slcdcton from the Lover MesoeoJc Havrkcsbut)? beda of 
t Soutli Wales, dcicribcil under 

preoccupied name of P/atyeps, 
POIP to BiUhriicft. 
fuoLT A-vnmAcosAL'Biae. — In 
I £unily the veriolvnl cotumn is 
If OBSiticd in the adult ; the teeth 
\ deeply infolded ; the mucous 
Hb between the orbits and the 
U fonn a lyic-shapcd pattern 
nni as tbc fyra ; and the ventral 
fne of the body typii»IIy has a 
(■ring of bony scutes. 'Hie skuti 
If lie {Btabolic, but is usually tri- 
{obr. This ramily may be divid- 
ioto three mbfamtlies. The first, 
Btf Attiia, it represented solely 
the genus Bapheta^ of the Car- 
lificrous of Nova Scotia, which 

only be provisionally placed in 
i family. It is only known by 
imperfect skull (fig- 963), which 
broiad, and rounded anteriorly. 

(he lax^mmatima^ChauHtd^ntia), the members of the one genus 
vmma atuin a lar^ size, and are characterised hy ihc triangular 
U (^ 9^.S)r which has large potiterior projertions, with the lyra 
ling iwo Mraight grtwves, continued posteriorly as ridges. The 

b«re compnessed, large, and irregular, with the foldings deeper 

rig. gA].— Vfifwr v1*w of tha Lkutl ■/ 
/. *jv* ni Aihnittti I from lite Cri^^v^l' 
ifrruut «if Durtiam. RDrtvi:cil. 1.4iL«n 


than in the Ardlfgosatiriia. The vertebra) centra are di^U 
and may perhaps be emlxdomerous. laxomma {fig. 96^) 
ia England in the Carboniferous, but has also been 

Fl|. 9IS4-— I>i>rul ttirfac* of ihe tlmll of jiMtinu»iaiitnt KiuielB : (Ron its CarbetOaem 
orEn|liuid. OiK-almh iuiuidI (IK. (Aflef A4ihv|'.) 

(rota tfie Pcnnian of Bohemia. The large size of the orbJO 
well ^own in the figure ; another charocicTistic feature being 
absence of ]>osiglfnoidiiI process to the mandible, llie 
scutes are unknown, and if it be eventually found thnt these 

wanting, thi& genus should perhaps form the tyft 

^ of a distinct family. In the typical subfamily 

W^ AHihrMositun'na the skull (fig. 964) is triangular, 

Mb and characterised by the small Kite of ibe pali- 

H ■ tal vacuities ; while the teeth arc b-ubcylindricaL 

^1 ■ The type genus jinihratosaums occurs, as its 

^m ■ name implies, in the Carboniferous, ranging frooi 

^HH Britain to ihc Continent. One of the denul 

V^W ventral scutes is shown in fig. 965. The sklrfl 

^^^ was covered with scales or tcutcs ; and it 

. ^r* ^"^^'..'^ appears from one specimen that these »cut«*4)d 

Aitbcy.) not extend over the pancial foramen, which ib- 

duces l)r Credner to consider that the PoUeomc 

Lahyrinthodonts were provided with a functional parietal eye, d 

which an aboned rudiment persists in ^fumnhn. The imperfectly 



Id J^ttsyaft, from th« English Pcnnisn, is allied in cranial 
to AmiJiriuosaunis. Nforeorer, some writers place in this 
the imperfectly known genera P/afyeft, from the Pcnnijm 
toasii, and Mofntmrion, from the conesponding sir»ta of 

ciLV MASTODoysAURiDjE. — The members of ihJs family, to- 

with some of the Amhratosaurida, constitute the Eiigtypia 

NfiuH's classtiicatioo, and muy be regnided as the 

reprtscntativ'cs of (he order. They are distinguished 

the Uacr family by the still more complex structure of the 

; the stronger sculpture of the skull ; and the absence of 

on the ventral ^iirfaix of the body. Kirgc palatovamciine 

(fi|f. 966) OK pbceti on the inner side of the maxillary 

axid there b a oorrespooding inner series of smail teeth in 

fei :t> 


FIb. ^Ml— Fdu»l HMCt of ttM truluin 

'1§ MtmHi d i' mmta'm #*/iHI/fflH i ttt^ the 

KMf td Warluwl a%. RcdiKcd. (Artct 

Pin- ^. -' MmilBdimiaiirft 
giftnttfi. », Uonal upcct 
dl >kut!, Rrculir nducnl ; i, 
loo-ili on ■ Iwgci Ksle. 

muicbble ; while there is no Ijony rinp in the sclerotic. The 
features may Ik; also noticed, although some of tht-m arc 
fiomiimn to the typical Anthraceiaurida:. The m.iiidiblc has a 
Inge pOKtgienotdal procesit, and the crowns of the teeth are conical. 
Tltt palatines have the &ame position as in the ArrA(gf)sattrida. In 
the pdns of the type genu< the piibes are separate from the t.schia, 
and do not enter into the formation uf the acetabulum ; and the 
luial ribs form kidney-like disks. The centra or bodies of the ver- 
ichac is the adult form disks which arc fully o:>eil)cd ; but in the 



young, as we infer from MasiaJonsaurus, in which alone 
has liL'cn observed, they were rluehilomous, as in the aduM 
tion of the AriAegt»iiurnia. The pubtaJ vacuities weze lai 
approximated, as is well shown in fig. 966. This familyj 
divided into tvru groups, according to the preiicnce or aba 
an inner nniculnr buttress at the proximal extreinit}' of the ni 
The gruup in which this buttress Js present is reprcsentedj 
type genus Mastodonsaums, best Icnuwii by llie huge AfaslodM 
g)gante»s (fiys. 966, 967) of the Tnas of Europe, which pf 
attained a length of seven or eight feet, and ranged from tl) 
chclkaJk to the Khfctic Besides other European Triassic j 
this genus is also represented in the l^owcr Mcsozoic (Ma]a 
of India by a form closely allied to M. gigatiUut, and by anj 
the Hawlcesbury beds uf New South Walcj. Figure 968 ^ 


I ■ 



of M-nt/Jtrnmatta ^tf^aJtut. (^really cnliugcd. (Alter 

Fie. *j._ 

the i-'ffa O 
of Indik a \ 
Mcdoa of a la 

Structure of a traiuiTrse section of a segment of a tooth 
enus ; ihe mode of attachment of the teeth to the jaw b4 
'^hibitcd in fig. 969. In the transverse section it will be oi 
that there is one set of sinuous linear interspaces coinmun 
-with the exterior, and a corresponding scries (scixaraicd froi 
other by the dental wall) of sinuous procesiies from the centra 
cavity. Other genera of UiJs ^roup arc CapUosaums (int 
CydolosaurHs), frotn tlie Keuper of Germany ; Trtmalosovnti 
the Bunter of the some countr>', distinguished by its more t 



; tnd the aptarenUjr allied Goniogiyptus, from the Panclict 
of Ute Lo«rrr GurKlwunns of Indix PiuMyj^nia (fig. 953), of 
latter dcposU, laay oUa be pfxivisionally included in this grou]>. 
the- Mideri staijc of the Upper Clondwatus we have also a 
.apjnrenUjr cloacly allied to Capitosaurus, and thus 
precise parallelism in the evotutinn of the group in the 
knd European horizons. Here aUo wc may perhaps place 
hntall ronn dctchlNrd under ihc proviiiional name of GfyJUo- 
M, frvm the Indian Pancbcts. Mtt"pia$, of the Continental 
and Rhxtic, is diminguitthed fioiii the preceding genera by 
matt *nterior position of the orbit* ; ZabyrtntJuufcn, of the 
Kcupcr, beitig probably allied Tlic sctund group i* reprc- 
-b^ DioAtc/piatitu, of the Wanickshirc Trias, in tvhtch the 
i»le has no inner articular buttress, 

UJrtZtRTAiM Kamilv. — Here may be mentioned the genus 
s, foDndcd upon large vcrtclval centra (fig. 970), from the 

> l(. ana-TMn M>ut«ml Mam gf Emmmm aiaJiattnt : bun l>i( CMbDnifcrvus 

CarbonifeTOiu of Nova Scotia, which were regarded by I'rofessor 
Mxnti as belonging tu an Ichthyosauroid Reptile, but which really 
indicate a large Labyrinthodont, perhaps rererable to the Mai/o- 

Leat-ing out some ill-defined ger»era, mention must l)c made of 
Purepias, of the British Carlioniferous, wliicli was formerly classed 
octt to BittrathidtrpHm. 'Yha skull is eloiij^aicd, and reniarkahle 
lor the incomplete orbits; while it appeals lluit many of the ordi- 
nary bo»es are wanting. The cranial Ixines have a pitied sculpture : 
and the vertebral centra arc thick and fully ossified. 


fare foot, some of tho«e of the hind foot having a length 
i n cbe*. llic al»cncc of any knou-n Dinosaurians from 
M stroi^y in faTour of thi; Labyriiithutlont nature of 
; and it has been .si^jigrstcd that those described as 
Sarfki mvtc made liy Tremelmanrus Jiraum of the 
r, the bkuU of nhich has a length of ci^tht inches. If this 
prove to be the case, the name Trtmatosaunts would have 
to tbe earlier Cfiirojaurus. The laige^t traclu fri>in the 
of Cheshire have been described aft C. Htratlis. Oihcr 
lis of feet from the rcmuan and Tria^, which may be of Laby- 
podoiu ongtn, have been described under distinct names which 
liOl be untK<«ssary to quote. 

Ow'tK IL Atoi>-*- — The remarkable limbless Cajcilians being 
fcnnwn in a fossil state require no further mention. 
llCNtUfJt III. Cai;data. — in this order the body is elongate, and 
hccmform or anguiform, with a tail, and usually with two, 
I o cc asio n a ll y only one, pairs of limbs. The cranium laclis the 
sapratemporal, and supraoccipital bones of the ijtbyrin- 



M. . 


ami the palatines are approximated to the middle line, 
, d immially to the vomers and pterygoids. The ribs are 
|uid the bodies of the veitebrse arc either amphi- or opistho- 
». The reiemblance in llie contour of the skull and the 
Mrt ribs to the Protriiomda suggests an affinity between the two 
'nps: bm the po^ilio^ of the pabtines is rather indicative of a 
MwMhit* with the Xyraniidtt. 


104 1 

bf lite ftwnee of a maxilla, and of 6vc digits to the 

lUEs Sacxiae akd PnoTErD*. — Sire/i (fig. 971, a) and 

tagOba viih AltnefiroHt^us (ibid., c) arc characterised by 

I p cHHifil jib, ihe sbs«nce or inaiilla:, the amphicuslous t'cnc- 

aiuf the ittJBCtion of the number of the di^tits hclow five ; Stmt 

-r.z fmn ibe oihcr two by the aljscnee of pelvic limbs. No 

■- arc known. 
— \~..iAUi\>Jf- — In this family the gilk are shed, and 
•.ate pmtent ; but it agieea with the two Inst in the aniphj- 
kxftriuv, the canilnginous carpus and tardus, and the ah- 
:of f^'cjidi. In tlic ty|>ieji) North American genus Ampkiuma 
.9;!, •)lhe body is inui^-h elongated, and the limbs :ire very 
j\sadi(r American genus is Mtiwpoma^ allied to which is 
vtnthu (Ciypiobranefivs or SieMdia), typically represented 
'(^i^itic Salamander (J/l maximits) of China and Japan, and 
inh DC may prolialily include the large Salamander (tig. 97a) 
ibc Upper Mtocene of Switzerland, originally regarded as 
and »utnefiuently described under the name of Andn'as. 
'FuitlV SAU\MANUKtL>^ — The true Salamanders loac their gills, 
there are iruianceti, as in AtnblysUima {SiruUn), wliere 
Ity periirt in some indi>-idual*. Eyelid* are present ; the vertebne 
ke (tnoiFly opisthoctelous ; a.nd the carpus and tusus mure or less 
pabi I'hii ffl^mily is now represented in Europe by the S.tla- 
■uidcn {Saiammmdra) and N'ewts {Alv/ge or Triton). In a fo.<isil 
the eaiuing iieij^ eristala (tig. 973) occurs in the Norfulk 


i%. en-— Tte Cncrad >*«ivt (,V«4r>r <iTr/aU>. 

Hmco, and reprecnlatives of lhi» genus have also been recorded 
Rhe Middle and Lower Miocene of ihe Continent. TIic latter 
poiits have also yielded remains referred to Saiamandra, while the 
IK CiuiofriUm has been applied to an imperfectly known form from 
t Lower Miocene of St G^nd-le-Puy, in AHier. He/iarehon, frotn 
(corresponding strata of Rott. near IJonn, is allied to Saiamandra^ 
aay not improbably be identical with CMotriion. Mtgalo- 



triton is a large form from the Upper Eocene PhosphoriM 
Central France. 

Order IV. Ecaudata. — The Frogs and Toads form a U 
specialised order of comparatively late origin. In the adidl 
body is short, destitute of a tail, and furnished with four Safe 

Fig. 974.— A, RkFteion or the Tiag^Rmalim^rmria): tr, TnnivciH procoMfoT ttti 
tOt Sacrum; «r^ Urotiylc ; f, lUum; j, SupnucapuU ; #, Kumcnu ; mA^ lUidiiu and ol 
Cirpii^ ; /, Po^cx ; y. Femur ; fr, T^bla and fibulA ; cd, Calcuieo-utragalui ; im^ Tuw 
Prohillux. n, Petioral arch and Mernamaf Frog {after Gegenbaur). The dulled parani 
cariilaifes. t, EpisternuiD : e, Omoitemum ; /, Body of the memtun ; x, Xiphuieniu 
Supraxcapula ; tc, ScapuU ; d, Precotacoid : ee, CoTKCoid ; t, Epiconcoid. 

which the pelvic pair is the longer, and adapted for leaping. 1 
are no gills. The skull is short and wide, with enormous o 
and the parietals confluent with the frontals. A peculiar ossific 
known as the girdie-lione encircles the skull in the ethmoidal re 
and there is a predentary ossification in the mandible. Thi 



arc few in number, and generally procoelous ; there 
vertebra in the sacrum ; the \'cnebral column terminates 
urostyk (fig. 974, »r). and there are usually no separate 
iUa. (r") arc proloi^cd backwards !>o as lo throw the 
um far behind the sacrum ; the radius and ulnn (a^), nnd 
»nd fibula (tr) arc respectively fused together ; and the inl- 
and nstragnlus (ra) greatly elongated. There are five digits 
icfa foot, with an additional ossicle (sp) io the pes which 
rcntly represents a prohaDux. 

Dconjing to the presence ur nbseoce of the tongue this order is 
fed into the snbofders I'harterogluesa and Aglossa ; the latter 
tuning only tbc two families DaityUtkrida and lipids. I'hc 
bnoglosBa are subdivided into the Firmistermne and Ardferim 
pi; the former chaiactcnscd by the cpicoracoid.s Coiming a band 
Hexing the cotacoids (fig. 974, b), and the Liiier by the over- 
bioc of tlic cpicoracoids. Since fossil forms ore l>ut very imper- 
R known, only brief mention will be made of tiiose families 
dt have fossil rcprcMrntativcs, 

I^UfiLv l>iscocLossinje. — Commencing with tlie v^rr/mw series 
the PhancrogloBsa the present family is distinKui^bcd from those 
t fbUow, and ifacreby^ apptoximatcs to the Newts, in having opis- 
CEBkHa vertebra: and rudimentar>- nb&. The European genua 
^akimmtar \^ probably represented in the Upper Miocene of ^wir- 
lind ; although some writer; h^ve referred the lo»»iI species to a 
tioct genus uodcr the name of Pelnphilus. Opi>.thoccelous ^-er- 
me from the Middle Miocene of Sausan, in (.iers, may belong 
^ci to Bomhima/cr m to the other existing Euro|>ean genus Alyies. 
tffbrK', from the Middle Tertiary of It^ily, is said to present many 
biities tu tbe present family, but in the absence of ribs approxi- 
}fta. to the one that follows. 

;FA]ni.T Peloimtid*. — This small family is characterised by the 
of teeth in the upper jaw ; the absence of ribs ; and the un- 
extrcmity of the sacral nix The vcttclinc are usually pro- 
although occasionally opi&thooEluus. The existing gunus 
occurs in the Miocene of Sausan, while the imperfectly 
Protaf<ti^*at4S, from the Miocene of Bohemia, m.iy belong 
to thb or the next family. 

Lv FAi.-»:oiiATRACHin.*L — ^This extinct family has teeth in the 
>aw ; no ribs ; cxpaiided ribs to the sacral vertebra ; and pro- 
toAoos vertebrx. The winyle genus PaltrolKitr<i(Aits {PrabatratAitt) 
9 flow known to ha^-e been widely distributed over the Continent 
daring the Lower Miocene ; more tiian a doicn species having been 

Fahilv BiinixiD*. — The true Toads are characterised by the 
galahtCDce of teeth and dor>al ribs; the expanded extremities of 


the ribs of the sacral vertebra; and the procoelous vertebne.' 
the type genus Bufo existing species occur in the Europeaj 
Indian Pleistocene. S. Gessneri, of the Upper Miocene of i 
land, appears closely allied to the living B. viridis, althon^j 
gether with another species from the same beds, it has been : 
the type of Palaophrynus. Dr Filhol records the type gentH I 
the Upper Eocene Phosphorites of France. 

Family Cystignathid*. — This family represents the Fn 
Tropical America and Australia. The huge Ceratophrys . 
Horned Frog of Brazil, occurs in the cave-deposits of that < 
while Laionia, of the Swiss Miocene, appears to be a dosdy i 
if not identical genus, characterised by the smaller head, 
slender pelvis, shorter manus, and longer pes. A somewhat 
instance of distribution occurs in the case of the Chetoniaa 
dra. The cave-deposits of Brazil have also yielded remains of I 
existing Lepiodadytus pentadadylus. 

Family Ranid*. — In the Firmisitrmne series the True 
have teeth in the upper jaw, and the extremities of the sacral 
are not expanded. Rana is represented by existing species in I 
Norfolk Forest-bed, and probably in the Pleistocene of Sardiiua; 
probably also occurs in the Miocene of Sausan and other places i 
the Continent ; and has been described from the Upper 
Phosphorites of Central France, where the one known species i 
to be allied to the Indian R. tigrina. The Oriental genus 
sus, in which there are no vomerine teeth, is found in the Eocene* 
Bombay, Ranetvus, from the Middle Tertiary of Italy, may ] 
belong to this family, although it is said to show affinity to the 
balidiE ; and we may here mendon the imperfectly known Aw^ \ 
rana, Batrachus, and Protopkrynus of the Lower Miocene of FnnO^ 
although their family position may be doubtful. 


1. Atthey (T.)— "On Anthracosaurus Russelli." 'Ann. Mag. NU 

Hist.,' sen 4, vol. xviii. (1876). 

2. Bacr(G.)— "Die al teste Tarsus (y4rrA«iw<i«r»j).'' ' ZooL Aiudga.' 

No. 216. (1886). 

3. BuRMEiSTER (H.) — " Die LabyrimHodonten aus dem Bunten Sud- 

stein im Bemburg." 1849. 

4. Cope (E. D.)— " Description of Extinct Batrachia and Reptilia from 

the Permian of Texas." 'Proc Amer. PhiL Soc,' vol iv«- 

5. " On the Intercentnim of the Terrestrial Vertebrata." 'Trans. 

Amer. Phil. .Soc.,' vol. xvi. (1886). 

6. " Synopsis of Extinct Batrachia from the Coal - Measures." 

' Geol. Surv. of Ohio, vol. ii.. Palaeontology * (1875). 



'" D.) — "The Batrachia of the Pemjiflin ol North America." 
i-rican Naiunilist,' vol xviii. (1S84). 

e (H.) — "Dst Stenoccptialcn aus dcm Rnthliegendcn dea 
rn'fichen Grund^^ bei Dresden." 'ZeitKhr. dcutsch. gcoL 

vol«, xxxiii.-xxxviil (1881-86). 
- (\V.)— "AcadUnGeolftey." 2d cd. 1867. 
r — ■• Air-breAtheri of lh« Coal-Period." 1863. 
Idllo /L.V— " Now 9ur le Balrachier de Bcmissart" 'Bull Mus. 
" Mist, Kat. B<Ik-." voL iii. (1884). 

-IN f E-1 and Atthey (A.K" On the Skull .nnd some other 
-f LfitrcmMt AllmaMni." 'Aqd. Mag. Nat. Hisl.,' scr. 4, 
,*..^v ' >... - "Die Labyrinlhodanten dcr sctiwabiachcn Trias," — 
■ * PaLrocitogrspbicii,' vol. xxK>-t. f 1S89). 

tn-s-rM 'A.''— " FauiKi der Oaskoblc undder Kalkstcinedcr Pcnn- 
. BOIimcns." PraRuc Ct883-86!. 
. '—"LesEnchaincmcntsdu Monde Animal, &c. — Fossiles 
r-nmaircs." Paris f 1883). 
** \jc* Reptiles dc I'Epoque Pcrntienne aux Environs d'Autiui." 
t* Boll. Soc. Ctol. France,' icr. 3. vol. vii. (1878). 

" I.'Actin<>don," 'Archiv. Mus. Hist. NaL, Paris,' toL x. 

UJET fT. H.) — " New Labyrinthmlonts frnni the Edinburgh Coal- 
"eW." '<3uaiT- Joom. Ceol. Snc..' vol. xviii. r863. 

** Description of Vertebrate Kcmains from the Jarrow Colliery, 
Kilkciuiy. 'Trans. Rny. Irish Academy,' vol. xxiv, (1867). 
— " Vcnel>ra(c FiwmU fmm the Panchct Ri)cl«," ' Pahmntotogia 

Irdira (MeiiL Geo!, Sun-. Ind.),' scr. 4. vol. i. (1865). 
Lvt'iKKKK <R.>~"Tbc Biiori LabyrinihoJoni." ' PalKonioloi^ia 

Iiitlica.' scr. 4. toL L (1885). 
MlALL (L C.>— " Report on the Labyrinthodonts of the Coal-Mca- 
•tnT*.' ' Repi Brit. A»»oc.' 1873. 

** Rcporl on tlie Structure and Clauification of the Labyrimho- 
dootB.* Uid. 1 874. 
0*r»:M(R.) — "Fowil Repu'lia discovered in the Coal-Mcasures of 
South Jogxins Nova Scotia." 'Qu^t- Joum. Gcol. Sot,' vol. 
Kviii. (1862). 
■ "Ooa Ijibyrintbodont Amphibian (*A>AV(j//«*/j from the Cape 
of Good Hope- md., vol xl. (1884)- 
. PoBTis (A.) — '■Resti di Batraci Fossili Italian!." 'Appunti Paleonto- 

logi," pan 3. Turin ^885). 
'. Zrmu. (K- A)—" Handbuch dcr Pala;ontologic." Abth. i., vol iii., 
part J. Munkh (1S68). 




General Structure. 

With the Reptiles we enter upon the consideration of the Gn 
two classes which, from the possession of many common chuid 
have been brigaded together by Professor Huxtey under the 
of Sauropsida. The name Monocondylia had, however, bees 
viously proposed by Hasckel for these two classes, and some 
consequently prefer to use this term. These two classes lie 
Reptitia and the Aves, or Birds ; and although recent researdi 
shown the existence of a close affinity between the more _ 
Reptilia and the Amphibia, and thus with the Mammalia, yet it 
in no wise tended to interfere with this association. It should, 
ever, be observed that since it is probable the Reptiles have 
origin from forms more or less nearly allied to some of the eariicr 
phibia, with which we are at present acquainted, it is obvious 
there must once have existed animals in which the characteristic fa-' 
tures of the true Amphibia and the Reptilia were more or less Uaidei 
and that the practicability of drawing a distinction between the ti* 
classes is thus (as in other cases) more or less due to the impeifec 
tion of our knowledge. With this proviso, and bearing in mind thil 
some of the more generalised forms with which we are even no« 
acquainted may not conform in every detail with the undennenlioneC 
characters, the Reptilia as a whole, together with the Birds, may b< 
distinguished from the preceding classes on the one hand, and from 
the Mammalia on the other, by the following features. 

Epidermal structures in the form of scales or feathers are gener- 
ally present, but there are never hairs. The vertebrse, which art 
ossified) usually have no epiphyses.' The basioccipital, with on< 

> These are present in some of the SAuropter^ia among Reptiles, uid b 
I'ariols among Birds. 



I exceptions, is completely ossified ; arxl, in conjunction with 

npitaU, forms the single occipital condyle hy vthich the 

aniculatcK with thv atlas vertebra. In the adult there is, 

t, no diaiincl paiasiihcnoid ' on the base of ihc skull. 'I'he 

rami respectively cottsist of an articular caniLige-hone and 

atembrane-bcnes ; the articular bone being connected with 

of the cranium by a quadrate. The apparent ankit;- 

tn all exiatiiiig fonns, is Mtuated between the piuxlmol and 

rows of the tarsus; and not, as in the Mamiiulia, between 

luid astiaKalus. Gills are ttct'cr developed during any 

o( life ; the embryo is prcn-ided with an amnion and an 

and there are no mamotary gland:>. A» the palaeontologist 

have to deal with the other distinctive reature.s derived from 

parts, it will not be necessary to refer to them in this work. 

ling the features of Reptiles as distinct from Dirds, the 

iaaution between the two classes is so close that it is difficult 

any very clear diagnosis. In the present class, however, the 

anicio ree take th< fwm of overlapping homy scales 

ita). or of shickla with their edges in apposition (CWlonia) ; 

1 dennal bony scutes are verj- frequently developed. The rcr- 

may be omphiccclous, opisthocurlou-s or proctclous ; but the 

of the cervicala do not have cyhndroidal and aadcllc-shaped 

surfaces. The sacral vertebra:, when prcacnt, have brnad 

ribs for articulation with the iha. The sternum in cxi.<<t- 

is thotnboidal ; and the libs may be attached to it by a 

median process, or jiroceises. The tnlerrla^icle is never 

mth the clavicles. Tlicie are more lliaii three digit:* in the 

; ; and never less than three in the pes. Except in the Thero- 

brattcbi the three elements of the pelvis as a rule 

I dtttinct ; * and there is apparently no known instance, except 

! which may probably lie regarded as a pathological peculiarity, 

! bwion of the inetatarsal&, or of their union nith the d»tal row 

:ta»us. In all living Kcplilcs there is both a right and a left 

: arch ; the arterial and venous circulalions arc at best hut im> 

separated -, and llie blood is cold. 

the various orden of Reptiles differ so greatly from one 
in structure It will be advisable to make most of our obscr- 
I on their oMcoIoigy under those >>c«'cnil headings. As a rule, 
T, the bones of the cmnium retain the general arrangement 
ale in the .\mphibia ; there being distinct postorbital or 
psifrontal ossifications ; usually either one or two leinpoial arcades; 
ud distinct post-, supra-, and infratemporal fosste, as defined in the 

' Dr lUiH nasgett* llut thii bone kujt lie pment in PalitaiafUrta. 

' TTicf ate uicbyliMnl in Ttituda oilm of the Indian Slwaliks, nd alM In 



introductory chapter (p. 904). The bones of the skull aie ' 
dense ivory-like structure, and in most cases their sutures 
although in certain groups the premaxillse, frontals, and 
may respectively unite. A new element — the transverse, or 1 
palatine, bone (fig. 1089, 7i) — connecting the maxilla 
pterygoid, appears ; but this is absent in the Chelonia and 1 
Ophidio. An epipterygoid, or columella^ connecting the 
with the parietals, may also be present A parietal foiamen 
may not be present ; and the occipital condyle is usually 
the hinder extremity of the cranium. The apertures of the ( 
nares may be terminal, as in the Amphibia, or approximated 1 
orbits, as in the Birds ; and the bones of the palate may 1 
a floor underlying the narial passage and thus cutting it off I 
mouth. As a general rule the mandibular symphysis is aotj 
chyiosed ; but this takes place in the Chelonia and Omit 
The quadrate may be either loosely or immovably attached UJ 

The dentition is usually well developed, and the teeth of 1 
parts of the jaws are occasionally more or less differentiated ; 
there is no known instance where they are implanted by douUei 
or where their crowns have deep infoldings of enamel. They mijl 
present not only in the jaws, but also upon the palatine, 
and more rarely the vomer. In other instances, however, teeth 1 
be entirely wanting, and the jaws simply ensheathed in hom. 
teeth may be anchylosed to the outer side of the jaws, when I 
dentition is tenned p/eurodoni (fig. 975); or to their summits, «W 

the term a^radont a tf 
plied ; or they may fal 
set in a groove, wiA ■ 
without anchyloai ■ 
the bone ; or, findji 
they may be placed ■ 
distinct sockets, wto 
the dentition is said t 
he thfcodani. The teeth on the palate are generally anchylosed toft 
subjacent bones. There is usually a continuous succession of tert 
developed throughout life ; the new teeth coming up beneath thoi 
in use and absorbing the base of the crown, as is shown in the ted 
of the Ghariai represented in fig. 1090. In shape the teeth 
Reptiles present great variation ; but a very common type, fro 
which many of the variations are derived, consists of a more or le 
laterally<onipressed and recurved cone, with fore-and-aft cuttii 
edges, or carina, which may or may not be serrated (fig. 976). C 
casionaliy, however, the teeth of the jaws or those of the pala 
may have nearly flat crowns adapted for crushing (fig. 986). 

Fig. 97s. —Inner vicvi' or the left ramiu ot Ihc Ruuidible of 



both pairs of limbs are present the vertebral column 

>tiaicd tnio cervical, dorsal, lumbar, sactnl. and caudal 

and Oic iijitbnc themselves are 

Illy os&tlied, although in some prini- 

a small notochordal canal may 

Ihcir centra. In a Urge number 

fornw the inajoriiy of ihc centra 

but they are nmphiccdous 

Ibnil and a lew cxisiing types ; 

the Diaocaaru and Chelonia 

nxtous type b common in parts I'it-jjt. - rranie u.) b>mi 

l«eri«. In all case* there is an in- ^.Zfj'7MZ:::^Jy'^i!'^.t, 

between the skuU and the atlas ; te:?^:S^X"r '"•" = 

may eilber form the inferior ring of 

t, cw, whet) the centrum of the alias is separate (Ichthyoptcry- 

may t»e of the normal wedge-like form. In some groups addi- 

aniculaiionii may be developed on ihe arches of the trunk vcrtc- 

, taking ihe form of a wedge-shaped process, or tygotfikfne, fitting 

» a corpoponding caxHly, ot sx^anfnim{f^ 977). The transverse 

of the dorul vene- 
I way be cither loi^ (fig, 
k) ot Tcry short (fig. 
j) ; and the ribs ouy ar- 
eithrr by a single 
whh the transi'crsc 
or ljy Ivo heads 
It portions of the 
; pjoccs-v or liy one ar- 
Indation to the biter and 
ly nothct to a facet on the arch or the centnim ; there being great 
pfa&m as to the position of the transveriie processeH and nl^faccts 
■ dUerent groups, and also in the differetit parts of the column of 
(ih)^ animaL OccasiotuUy the ribs articulate at the junction of 
In ^ q tcb rifc. Chevron-bones are generally present in the tail ; and 
kttmtunt may be retained. No living Reptile with limbs has less 
tm two sacral rcitcbnc, and in certain extinct forms the numlicr 
■n be increased to five or six. In nearly all Reptiles the tail is 
Idevdoped. The ribs may have uncinate processes. In many 
forms the sternum, which may be ossified, is rhomlioidal, 
have the last pair of ribs attached to a backward median 
It^ structure in many fossil groups U not known, but 
to Ptofesaor Marsh's interpretation some of the Oino- 
had paired stental ossifications, corresponding to the two 
from which Ihe sternum de\'elop« in the Ratitc Birds, 
ribs may be developed in the parictes of the ventral 

H«. 977.— Pedcrlcr {*) wd tmoU (B}a*pcct> of k 
dcnol wmsbn «f > Siukc Th* tatiiiai in a on lb* 
■td^ of Uh near*! ciuuJ an llw ifSAnlra. 



surface of the abdomen, and consist of a median and two! 

In the pectoral girdle the scapula is generally a more ati 
elongated bone, with an expansion at its glenoidal extreniitf:^ 
separately ossified precoracoid exists only in the Anomodonii j 
978 ^/f), this bone being in other cases fused either iridi| 
scapula, as in the Chelonia (fig. 1008), or with the coracoiij, 
Lizards (fig. 830) ; the fontanelle which frequently occurs m 
latter instances apparently marking the original line of 
of the two bones. The coracoid may vary in shape fnwn 
spatulate form (Chelonia) to that of a cheese<:utter (Dine 

The humerus has in many cases no distinct distal 
(trochlese), although these are well developed in the 
Lizards, and Sphenodon. In the more generalised types then ■ 
quently an entepicondylar foramen to this bone, but in the 
and Chelonians the foramen or groove is ectepicondylar ; and il 
few instances both foramina are present The radius and 
always remain distinct. The number of bones in the caipua ' 
considerably in the different orders, but in Sphenodon alone 
existing forms is there more than a single centiale \ the five 1 
bones (carpalia) may be all distinct from one another (5g. 829). 
the pelvis of the majority of reptiles, the ilium is produced 
behind than in front of the acetabulum (fig. 831), and thelaHdl 
more or less completely closed by bone ; while the pubis (fig. Jj^ 
is directed downwards and forwards, and, like the ischium, usa4 
meets its fellow in a ventral symphysis. Among the Dinosaunot 
however, the pelvis may be of a Bird-like type, when the pobcsA 
not form a symphysis. Usually the obturator interval fmnt ■ 
open notch ; but by the ventral union of the pubis and ischium d 
the same side this notch may be converted into a foramen. "Df 
femur among the Dinosaurs may develop an inner trochanter; vA 
except in some Omithosauria, the fibula alwa)'s remains distine 
from the tibia. As in the carpus, the elements of the tarsus va; 
considerably in the different groups, but the centrale (navicular) i 
only present in the Anomodontia, the Proterosaurta (according t 
Professor Seeley), one family of Chelonians, and the Ichthji 
pterygia ; while, with the exception of a few extinct types, and perhaj 
the existing Chelonia, the fourth and fifth tarsalia in all land fom 
coalesce into a single bone, which supports the fourth and fifi 
metatarsals. In certain groups — such as the Anomodontia ai 
many of the Chelonia (fig. 829) — the number of phalangeals in tl 
feet of pentedactylate forms may be the same as in Mammals, 
2i 3. 3i 3) 3 — the digits being reckoned from the first to the fift 
In Lizards, however, and their allies the number of phalangeals 
usually 3, 3, 4, 5, 3 in the manus, and 2, 3, 4, 5, 4 in the pi 



[auDC numbers obtain in seme Dinosauna; but in existing 

in which ihe fifih digit of the pes is aburtcd, the 

are 3, 3, 4, 4, 3 and z, 3, 4, 4. Among the Sauro- 

and in the IchthyoptcT^-gia the number of ph^langcsls is 

increased. In a Ur^'c numbtr of forms the tail is long; 

is gmcrally short in the Anomodontia, Sauropteij'gia, and 

I legaids the ctassiiicalion of Reptiles, scarcely any two WTitcrs 

as to the number of orders into which the class should be 

and still less as 10 ihcir mutual relniion:», and the larger 

onder which these orders may be arranged. There is indeed 

litiic diffiruUy in rt^rd to existing forms, in »hkh the few 

ha\-e Iveeome more or le-w sharjily differentiated ; but when 

back to the early part of the Me^ozuic epoch, we find that 

an the orders into whieh the class hax been divided show 

I Bgns of psu&ing more or less eomplctely into one another, that 

{ quite ifupoesible ID exhibit their true relationship by any system 

daastficBlioD. The best arrangement seems, therefore, tn 

the orders under a series of diverging branches, which will 

Ic to one another more and more as wc recede in time — 

, about the epcKh of the Lower Ferroian or possibly the Carbon- 

, it B probable that, if we knew all the extinct forms, these 

would be seen to originate either from one, or from but very 

It ttans. In regard to the number of these branches, there 

roooi for a considerable amount of discussion ; almost the 

' abtolutely sure ground that we can feel being the aftsocintion of 

Ofders forming the Aicho^aurian branch. To a less extent 

Fcune retruLrk applies to the orders themselves ; and the right to 

distittcdon of the Protcrosauria is not admitted by many 

while there is rtot perfect accord in regard to that of the 

llhalu. The provLsional arrangcmcnl which is adopted 

Idas work Ls a modificaiion of one rtpenlly proposed by Ut (J. 

IT, of New Haven, and may be tabulated as follows, vir.. : — 

TTttromarous Branch. 
Sjma^tMirimi •• 

Areiotaurian ■• 

Order 1. Anomodontia. 

r 1. 2. Sauroplcrygia, 

1_ N 3. Chelonia. 

4- IcUihyo pterygia. 

J. Proierosauna. 

•• 6. Rhynchoccphalia. 

II 7. S(|uamata. 

8. Dinnxaiiria. 

■I 9. Crncodilia. 

» iol Omithosauria. 

b may be added that the close approximation to the Amphibia 
jnemed b; the earlier members of several of these braaches 


suggests the idea that Reptiles may have been derived from 
Amphibians by more than one line of descent. 

The Reptiles in the passage of time have suifered more 
than any other class of the Vertebrata, only four of the abovH 
tioned ordinal groups — viz., the Chelonia, Rhynchocephalia, 
mata, and Crocodilia — now existing ; and the second of these 
represented only by a single genus with two species. There n 
doubt as to the earliest known appearance of the class, since it 
been thought that Mesosaurus (Stereosiemum) may be of OA 
iferous age, but it is more probably Permian. In undoubted 
mian we have the Proterosauria, many of the European 
donts, and the Rhynchocephalian genus Paiaohatterta ; i 
many of the American Anomodontia occur in strata which 
referred by the Transatlantic geologists to that period. \ 
the advent of the Trias we find all the orders, with the enqd 
of the Ornithosauria and Squamata, more or less fully represoA 
And while the former order makes its appearance in the 
ing Lias, we have at present no traces of the latter till the 
Jurassic. The class reached, however, its zenith of devdofn 
in the Jurassic and Cretaceous epochs ; the greatest 
of huge aberrant forms being characteristic of the later p>tt 
the former and the earlier part of the latter epoch. Although 
one existing Rhynchocephalian genus is closely allied to TJnm 
forms, yet we have no instance among Reptiles of the ensteoce^ 
a genus right up from that period to the present day, as we htRbl 
Ctraiodm among the Pisces, thus indicating that the higher i(| 
ascend in the scale of organisation, the more rapid is the ctmp 
of types — the same law being exemplified by the occurrence of • 
isting species of Reptiles among the totally extinct Mammals of Ac 
Indian Siwaliks. 




Anomooohtu, Sauroptekyuu, AKD ChEIjONIA. 

JCS BRAKCit. — The Rcplili-s included in this brnnch 
: may, for the pn»cnt at least, be arranged in a single otdcr, 
some writers would prefer to regard ihe suborders into 
order is here divided as of ordinal iraponanca 'i'hc 
able feaiures found in this order are the rcKemblance on 
to the IdbjTinlhodont Amiibibb, and on itie other to 
I. AsoMODOJrriA. — ^This order, which is equivalent to the 
Iota (TbcromoTpha) of Professor 
|rc»cnta the follouiitg charactcriiUc 
L The body i* laccniform, and 
M art adapted for walking. The 
I OMBpatatirely short, with n fixed 
p, t pArieUl foramen, either one 
jlempocal arcades, and large nasals ; 
(alate the pterygoids meet together 
H of the basisphcnoid, which they 
b, bat diverge anteriorly ; while the 
b are generally small, and plaecd 
By to the pterygoids, as in Mammals, 
ttie temporal arcade consi:iis of only 
^ chain of bones, ii is a squamoso- 
ly one (p. 904). The dentition is 
^t. but the teeih may be anchy- 
!0 the lionc The vertebras have 
Dcluus and in lionic cases noto- 
I centra; the dorsah canning long 

ise processes, and the ribs aniculatin^ by double heads in 
trior region of the trunk. As a rule abdominal nhs appear 

tS ilic irVt humrnii of ItftitA 

lyuvni at Souib Africa. Half 
nManil hi*, s. Kn'epirandirlu 

ay. CAdei Hiulcy 




to be absent. An interclavicle, clavicles, and precoracotds a 
ent, and a sternum was probably always developed. The 
(fig. 978 bis) has an acromial process with which the prec 
articulates. The humerus (figs. 978, 982), is characterisei 
well-developed distal condyles, and the invariable presena 
entepicondylar foramen ; while its delto-pectoral crest is gi 
much developed. In the pelvis the pubis is placed ent 

advance of the ischium, to whii 
completely united, with the {wes 
some forms of a small fontanellt 
senting the obturator foramea 
ilium may have almost its whol 
in advance of the acetabulum, 
tarsus has one centrale ; and tti 
angeals of the manus and pes a 
caliy 2> 3. 3. 3. 3 in number 
Mammals, the whole structure 
foot being likewise of a Man 

This order appears to be coni 
the Permian and Trias. It ha 
considered that the Anomodo 
the parent stock not only of a 
Reptiles (with the possible excel 
the Ichthyopter^a), but also 
Mammals. Later researches < 
however, altogether countenan' 
view, although there can be ik 
Fig. 1,78 w».— Lateral M^t of that they are closely allied 
parent stock of Mammals, 
observations have indeed shown 
conclusively that this order is 
descended from the Labyrin' 
Amphibians, and more especial 
the Archegosaurian family. Thus in the small size or abseno 
obturator foramen in the pelvis the entire order shows most ■ 
affinities to that group ; while in the small size of the Cora 
some forms, in the presence of a distinct precoracoid (epico 
of very distinct condyles to the humerus, of the centiale 
tarsus, and also in the number of phalangeals, it has ch 
common both to the Labyrinthodonts and the Monotreme; 
are not found together in any other group of Reptiles, 
three groups also resemble one another in the non-devel» 
as a general rule, of abdominal ribs ; while signs of affir 
tween them are shown by the shortness of the tail, and th 

the canilagc bonn of ihe righi aie 
of the pectoral girdle of « Uicyno- 
tlont ; frotn the KarOQ riy^tem of 
Africa, k, Si:apula ; a, Acromial 
prtxreu of do. ^ f-^^*'i Precoracoid ; 
cor, coracoid ; gl^ (jLenoLd cavity. 
Half natural ti». 



the exocci;>ilal dements of the occipital condyle in the 
; order. In the presence of an entciJicondjlar foramen tu the 
the Anomodonls agree with Rhynchoccphahans, Sauro- 
ukI Miunmals. In many cases (fig. 978 iis) the piecora- 
a lai^e plate-like bone suturally united with the whole of 
border of the coracoid, and also aniciilaling largely with 
pfocesR of the scapula ; thus eichiliiling a iiarallelisra 
ibc ttruclnre of the pcctoMi and pelvic girdles found in no 
jlcs. The above features, together with certain poinU in 
of [he palate mentioned below, suggest very strongly 
It of the Moootrerne Mammals from the name primitive 
thai which gave rise to the Anomodonts. If, moreover, Dr 
o li^l in cortsideting that lliis order docs nui include the 
. ancestors of Mammals, it would appear that the dcveluptncnt 
lb more specialUcd repiesentatives has follovred a course in some 
parallel to that of Mammals, 
>iit>KK I. Pahiasai'kjx. — 'Phis suborder includes the most 
members of the order, which make the nearest approach 
tbe Amphibia. The cranium i* at once characterised by the 
ifiAg orcr of its pottero-lateral or quadratic region, after the Laby- 
ibadotii manner, by the postoibital, !>quamosal, and upisihotic 
jMs; Typically the pfllate, which approximates to an Amphibian 
ttf and has been compared to that of Nyrania (fig. 825), appar* 
Hy has l»o flooring of the nasal passage to form secondary posterior 
|n. The skuU also has two temporal arcades, and the external 
ibct of ibe cranial bones is frequently sculptured, as in the typical 
tliytimhodont&. I'hc vertebral centra retain a notochordal canal ; 
C ntKober of sacral Tertetine wok limited to two, of which only 
ic flupports the ilium ; and inicrccntra may be present. The 
lira is of a l-abyrinthodont type, the ilium forming a triangular 
ue elongated in a direction olilique to the a\is of the sacrum^ 
kh which it articulates obliquely ; and there being no oblmntor 
■asMSi between the pubis and ischium. The humerus pmbably 
doogmg to this group differs from that of other Anomodonts in 
b dight expansion of the extremities, and in that the lower aperture 
ftbe entepicondylar foramen opens on the distal surface of the bone. 
Fawlv Pariasaurii>.£. — The type genus Pitriasaunts occurs in 
K Bemfon beds of the Karoo system of South Africa, of which the 
irtebtate fauna presenu a Triasaic facies. The best known species 
t inMdfKi) attains thedimen&ionti of a large cifM-odilc ; and, with 
e ■nfijrtunate exception of the limbs, the entire skeleton is knuwn, 
id has been described by Profc?ssor Seeley. In addition co the 
idpture on the booes of the skull, mucous canals, like those of the 
ll^hDthodonts, arc also present The teeth are of uniform siic, 
, altbotigh ai>chytoscd to the bone, arc set in distinct sockets, 


and were replaced after the Crocodilian manner; their crownil 
somewhat compressed and grooved. The premaxilbe 
have been small, as in the Amphibia. There are 39 
of which 18 are presacral, and two are anchylosed 
form a sacrum ; while wedge-shaped intercentra are also 
The neural spines are extremely short, and the centra of Uk 
vertebras are very small in proportion to their arches ; and 1 
lirst sacral vertebra supports the ilia of the pelvis. Small 
present in the caudal region ; and there was probably a 

Professor Seeley concludes that this very remarkable and 
like Reptile is a direct descendant from the Labyrinthodonts ; 
affinities to that group being displayed in the characters of the i 
in the notochordal canal, and the large arches of the vertebne ; ■ 
support of the pelvis by a single vertebra ; as well as in the 
of the pectoral and pehic girdles. The Utter features, together ' 
the general structure of the palate, being identical with those of tfl 
Anomodonts, there appears every reason for referring this frmily { 
suborder of that group. 

The genus Propappus is founded upon a humerus, fifOD 
Karoo system of the Cape, of the above-mentioned type, and dM 
is no direct evidence of its distinctness from Pariasatms. U 
innominate bone referred to Di^nodon honiups probably bdoap 
however, to Propappus, which may thus be entitled to AM 
The pelvis and sacrum described under the name of Du j iui^ 
Hgriups also seem to indicate a member of this suborder, since A 
ilium is of the same type as in Pariasaurus, and is connected «M 
the sacrum by only a single rib, while there is no obtunta 

From the general resemblance of its skull to that of Pariasaum 
we may refer to this family the genus Anihodon, of the Sertl 
African Karoo system, which, although originally regarded by Sir B 
Owen as a Dinosaur, must be included in the present order. 1 
agrees with Pariasaurus in the roofing over of the quadratic tega 
and the continuous replacement of the teeth, which are in a unifnn 
series ; but differs in the form of the teeth, which resemble those" 
the Dinosaurian A<anthopholis. 

Family Pariotichid*. — This family, although agreeing with tl 
Pariasaurida in the sculptured cranial bones and the roofing a\ 
of the quadratic region, differs in the dentition being of a can 
vorous type. All the known genera are from the reputed Permi 
of North America ; and the family is included by Professor Cope 
the next suborder. The three genera are Pariotkhus, £ctocynodi 
and Patttylus. In Ectoeynadon the first premaxillary tooth is tu 
like, and there is also an enlarged tooth in the middle of the nuu 


The no&trils are large and lateral ; and at the junction 

; of the bones of itic pelaic wiih the maxilla, the taoth-bcaring 

b vide, and supports four pamlle) rows of small obtuse teeth. 

p/ni, which was originally described as a Labyrinihodont, 

are more equal in size. 

>ROER J. Theriodontia. — ^This suborder, which is taken 

Or Baur) to include the Pelycxisauria of Professor Cope, is 

by the absence of a bony loof over the quadratic 

I of the skull, and the presence of oaly a single wide temporal 

(fig. 979)1 apparently consisting of a conjoint squumoso- 

and quadrato-nuxillary aniade. The mandible has no 

I incutty. 

KMDe esses, as in the American forms, the vertebne an: still 

kl ; iniertentra may be developed, to which the capitular 

of the ribs arc anirulaied, and there are not more than 

three sacral %-ertebrx. The dentition is fully developed. In 

of the African fonn& at least the maxillae develop palatal 

Id floor the nasal passage, and thus produce tall and nearly 

potrterioT nares, sirikingly like those of MiiEninLils. The 

bite mnain separate. 

ibase of the typical African forms in which the pelvis is 

the ilium is somewhat intermediate between that of the 

■uria and Dicynodontia, having a distinct but small obturator 

The buments i» usually more or leRii of a Dicynudont 

, having expanded extiemiiies, and the entepicondylar foramen 

[fb lower apenure opening on the palmar aspect of the hone 

iz) ; there b generally a marked thin flange on [he postaxia! 

opposite this foramen which docs not occur in the Dicy- 

evidently nearly related to the Pariasauria the present 
departs farther from the I>abyrinlhodom type, as is shown by 
tk>» of the roofing bones in the quadratic region, as wcH as of 
;Wpcnor tcmpora] arcade, and by the absence of sculplurc or 
canalf on the skull. This adi.-aiice U -iIbo indicated hy the 
icnt of secondar)- poatcrior narw, by the fuller aiiaclimcnl 
rf the lib to the sacrum, and the relatively larger centra of the 
Mxcfane of the higher types ; as well sm by the development of the 
(htoniior foramen in the pelvis. 

FiMiLV Tamnocepma L rn.^ — This family may be t.iken to in- 
tWe two gigantic Anomodonts from the Karoo system of the Cape, 
faoilxd under the names of Tapincaphalus and Tilanesuchus. 
Ifc former is known typically by the exireniiiy of the cranium ; the 
*nichrx probahly belonging to it ha^ng shon and notochordal 
Cddra. A pelvis, found in association with some limb bones, has 
txxa ikscrd>ed under the name of PKoeoiaurus, but there is no 



Fig sTg.-Lcfi laicnl upcci ofihe ikull or CoiEciaifrMr 
flattictfi : from tbe Kan» ly^tcm of Souita Africs. Re- 
duced. Or.Orbit. Onlyumeof thechcek-Ieclb Ullhown. 

evidence to show that it does not belong to Tt^tai^haha. 
associated bones show that the coracoid was distinct 
precoracoid, and that the short and massive humenu 
ectepicondylar foramen. In Titanosuchus the dentiticm 
probably the case with Tapinoaphaius) was of a camivwooi \ 
the humerus is characterised by the presence of an 
foramen {in addition to the entepicondylarX which pieicedi 
through the shaft of the bone ; while a bone incorrectly ■ 
as the pubis shows that the precoracoid was fused 

Family Galesaurid*. — Nearly the whole of the typic2l 
dontia of Sir R. Owen may be provisionally included in thii I 

since, althoa^ 
genera have 
others double i 
skulls of tbe 
so closely 
another as 
render it ini[ 
refer them to 
families. Tbe bndrt 
characterised by 4 
humerus (when knon 
being of a more elongated type than in the preceding ^unily, n 
by the smaller size of its members ; while the vert ebrae were probiU 
different from those of the latter, and had no intercentra. Ti 
dentition is of a carnivorous tj'pe, and differentiated into an antcrin 
or incisive, series separated by one large tusk or canine-hke toofl 
from a lateral series of cheek (or molar) teeth ; thus simulating di 
dentition of carnivorous Mammals, and more especially that ttT til 
pol>'protodont Mesozoic Marsupials. There are no teeth on ill 
palate. The majority of the genera are from the Stonnberg ■■ 
Beaufort beds of the Karoo system of South Africa ; and »c u 
mainly indebted for our knowledge of the group to the labaa 
of Sir R. Owen. 

In the type genus Gaksaurus, with which Nythosaurus is ideal 
cal, the skull (fig. 979} is much depressed, with the nares divide 
by a narrow septum ; there are 4 anterior and 12 cheek-teeth, (1 
latter having tricuspid crowns. In Lycosaurus (Rg. 980, a, c) 1 
have larger forms distinguished by the lateral compression of tl 
skull, the distinctly double nares, short mandibular symphysis, ai 
by the number of the cheek-teeth being reduced to 5 ; the derelo 
ment of the tusks being very great jSlurvsaurus, again, appes 
to be a neariy allied but still more specialised genus, in which, 
the reduction of the septum, the nares have united to form a sinj 



Qm*drMO (fig. 980, b) alao includes targe funns with 
eloped tusks, having compies&ed crowns with serratiMl 
th<»e of the Mammalian genus Macharodus ; the narcs 
Hdcd. Other geneta from the Karoo si'Siem shuwing Uie 
»re CynoiAamfita, Cynasuchus, and Stt>J»jK>siiurits ; the 
one of the nnallest known forms. Tigrisuthus, again, 
itvbcd by its single nares; while Gor^mps, of the same 
tuu a naiimr Battened skull, with the anangcmcnt of the 
at (rom that obtaining in all the preceding {jenen, and 


■^t,Jto—faf^*Mfl vlwirt af itwWkoUnf ^jmiMiwi; •, AnMftsr ^w of ifcc 
HM«; baa iIm Kara* ■TMwi>' Sa«b Africi. Hadam]. i indinia the link' 
»Mik (After 0-«>.> 

Ucate a diitina (ainily. Detttfrcsaurus anti othur forms 
t Permian of Russia, which arc included by Sir R. Owun in 
ad 'llieriodonlia, arc noticed hclow, 
LV CleMvdrowd.«. — This name is applied by Professor 

carnivorous Tbcriodoniia, distinguitthcd from ihc GaU' 

1 either by the developmeot of teeth on the palate, or by 
nocdinar)' character of their dorul vcrlebne, in which 
[ercenira are typically present All the genera are typically 
r reputed Permian deposit!) of North Ami^rica. In the 
nus CUp^Arvps the premaxillary and maxilLirj- teeth nre 
ted size, atKl the denuiry bone of the miindiblc has two 

tu&ks near its extremity. Teeth are alu^i borne on the 

ds; and ihe neural spines of (be dorsal vcrtchtic arc not 

dy elongated. Ja Vimetrod&n, the most rvntarkablc char- 

tbc extraordinary development of the neural spines of the 

otebraB, which resembled those of Ntiosaurut (fig. 981), 



with the exception of having no horizontal processes. Tht 
of the spine in one species is more than twenty times the 
of the centrum ; and Professor Cope concludes that then 
formed a kind of elevated fin on the back, of which it is ' 
to imagine the use. Naosaurus diflers from the precedii^ 
above-mentioned horizontal proce 
the spines of the vertebne (fig. 
The premaxilla had one tusk ; aoi 
were two similarly enlarged teeth i 
anterior extremity of the maxilla, 
which comes a series of some twdv 
of equal size, with compressed and 
recurved crowns. Numerous small 
teeth are also dotted over the palatii 
pterygoids. This genus has also t 
corded from the Permian of B< 
Other American genera included 
family by its founder are EmhoU^ 
Edaphosaurus, Archaobelus, Tien 
and perhaps Lysorophus — Tie? 
being characterised by the presi 
well-developed abdominal ribs. 

Here may be mentioned the geni 
eorhachis, from the Lower Pern 
France, of which the lateral chee! 
tion presents a considerable resen 
to that of NaosauruSy although it 
known whether teeth were present 
palate. The neural spines of the v 
are of normal type ; but it is not 
whether intercentra were present, or i 
the centra were notochordaL The h 
(fig. 98a) differs from that of the C 
rida in the contour of the distal ext 
Professor Seeley regards the above-mentioned humerus fh 
Karoo system of the Cape, described by him under the n 
Propappus, as indicating an allied form, but there is not! 
justify this association. Stereorhachis may constitute the ty 
distinct family, but there is at present no evidence to supp 
view that this genus (together with Propappus) represents a 1 
order, for which the name Gennetotheria has been proposed. 
Family BolosauriD£. — This family is also typically 
from the reputed Permian of North America, where it is 
sented by the genera Bolosaurus and Cfuiot^x. In the 
the teeth are fixed in shallow alveoli, and have their croi 

Fib- 981.— Anterior view of 
dorul veriebni o{ .VaeianrtH 
clavigir; from ihe Pennian of 
Texas. Onc-ftixlh nalurml tizE, 
Ct, Centrum. (Aftci Cops.) 



ly to the axis of the jaws. Those croims are 

their baie, and have^ a low apex vertically divided into 

ons, of whkh the inner one in the upper jaw is low and 

I, and the outer forms a curved claw-like cusp — the 

teeth consisting siinpl}- of an inner ledge and the outer 

i there being no cnlan;cd lusks. Afflarmfsaums, of the 

losits, nujr perhaps be referable to lliis family. 

Urge OenttrvinuriU, from the Russian I'crmiun, the prc- 

tceih (fig. 9Sj) approximate 10 the de^scription of tliu&e 

MmM, but there are lar^ 

'. teeth resembling those of 

wtrridr behind the five prc- 

■ teeth , the narcs beiiif; di- 

This genus may be regarded 

lenting a distinct carnivorous 

A tooth from the Karoo 

if South Africa, haiing the 

characters of the anterior 

DtJitfntiaurus, but with ihc 

Olden of the inner surface 

cjuvn rormitig ridges, has 

fcdc the type of the geni» 

in. hn associated series of 

B the British Museum may 

D the same form ; the vertc- 

notochordal, and the bumcrua 

D size with the one mentioned 

s Srilkiypus. Here may be 

the genus Rhopahdotty from 

nian of Russia, founded upon ria. <*.-Ant^,^. ^ ,h, u»«v 
■buuir minus which may he- f*" •^ hnin«nit of SurttthiKhit J»m- 
the same family. Perhaps, Oi»-t«if natttr.! ««. (Aft.r Cndir.) 
\ the mo&t remarkable specj- 

n these deposiu is the distal portion of a large humerus 
d mdcr the name Briihcpm {Rurasaurui) ; the proximal 
nothcT humous, which has received the name of Orlhcpits, 
' belor^ng to the same species. The former sipcciincn 
"table for ha\-ing both ectepicondylar and eniepicondylar 
condition elsewhere known only in 7'^ianosMchm and 
'K, Or Baur has suggested that <hi54 specimen m^y belong 
icbocephalian, but it is ceitain that it \% referable, as ijir K. 
pointed out, to the present suborder ; and it is quite 
X it may prove to belong to Dtuttrataums, in which case 

should be superseded. 
V OiAlttcnDX. — This family is also founded upon genera 



from the Permian of North America, and includes Diadttta,^ 
pedias {Empedocles), and Helodectes. The teeth (fig. 9! 
transversely elongated like those of Boiaamrui, and arei 
divided by a median vertical ridge ; but both the innet 
outer moieties are equally low. Their alveoli are not 
and the edges of the crowns are obtuse, with tuberosities od ; 
of them distinct from the apex of the main ridge. Profesurl 
regards this peculiar type of dentition as indicative of a 
diet 1'he brain-case differs from that of the CUp^refUm 
manner analogous to that in which the brain-case of the Va. 

Fig. gS^— LtlcriJ view of 
a prcmaxLllary tooth of DtU" 
ttrotaunti hmrmUttt I froin 
the Upper Permian of Rufwa. 
Half DBIural size. 

Fig. 984- — Laientl and palatal new 
of a poMmor toocb oT Bmftdim* mm- 
Utrii : rnuo tlu Penoian of Nonli 

is distinguished from the same part in other Lacertilia — that is, it 
continued between the orbits so as to enclose the olfactory lob 
in bone. Phanerosaurus, from the Permian of Germany, is refen 
by Professor Cope, from the structure of its vertebrae, to this or tl 
preceding family. 

SuBOHDER 3, DicvNODONTiA. — In this suborder the verteb 
have no notochordal canal ; intercentra are wanting ; and tl 
sacrum includes from four to five vertebrae. There is in no <3 
more than one pair of teeth in the alveolar borders <rf I) 
upper jaw, while there are none in those of the mandible. Tl 
palate is of the general type of that of the Theriodonts, but tl 
premaxillse unite to form a single beak-like bone, and the mandibol 
symphysis, which is very deep and laterally compressed, is likewi 
anchylosed. The nares are double ; and it is probable that in sod 
forms a part or the whole of the alveolar borders of the mand^ 
was sheathed in horn \ while the mandibular rami have laiei 
vacuities (fig. 985, b). There is a single temporal arcade, whii 
appears to be a squamoso-maxillary one. In the pelvis the Ilium 
much expanded in an antero-posterior direction, the expanded pla 



riy parallel to the ^crum ; and there is a small obturator 
The humerus (flg. 978) is expanded at the two ex- 
witJi a prominent dcltopcctoral crest.' ard with the lower 
of the entepicQtidylat foramen opening on to the palmar 
The memhcTS of this group -are found in the Stormbcrg 
iftjrt beds of ihe Karoo sj-stem of South .\frira, and the 
il GondwMtas of Central India. 

uv UtCTNODOXTlD*. — Thts family is characterised tiy the 

of teeth on the palace. The t^iK genus Diiynedon was 

known representative of the order, and was originally 

by Sir R. Owen from «pccimcn& brought from South 

:il}.-tUHBl T'm afih* ilnill t/f a1 DinwiJfH lattrtittfi. am! (b) Udmmhi BminJ: 
(rcB lb* Kafoa ipDia of SouiK AAici. Rnluctil. (Alttr Owan.) 

It is chaiacierised by the presence of a lusk-lilce looih 
Is. a) rto**"? fro*" s persistent pulp in each maxilla — the 
of the jaws liciog edentulous, with trenchant edges. The 
of the slcuU is rounded, the maxillx are not strongly ridged, 
BiTte arc approximated to the muzzle, and the supraoccipital 
a broad bar above the foramen magnum. This genus may 
ifco occur in ihc Panchel stage of the Indian (londwanns. In 
0. Hgriftfii the skull has a length of so inches. In Ftyehoriagtim* 

' The ikfaopceioral crew b itic ridiieon ihe ilgM ti'leof Ihe upper half of the 
' Thii Dcw nunc l» propcxteJ in Un of J^yficgiMfAus, which u preoccupied. 
VOL 11. M 


{Ptychognathus) there is also a pair of tusk-like upper tee^j 
the skull is angulated, with strong ridges on the maxilta, the 1 
far behind the muzzle, and only a very narrow supraocdpiB 
above the foramen magnum. The typical species are from 
Africa, but another representative of the genus occurs 
Gondwanas of Central India, which was originally dc 
Di^nodon orientali's. A very imperfect and flattened skeleton I 
the Karoo system has been made the type of the genus 
thus, which is said to be characterised by the small size 
canine-like tooth, and the presence of only two phalangeab ii 
the digits except the third. It appears, however, that these 1 
differences do not really exist, the difference in the humenii I 
due to a comparison of opposite aspects, and the numl 
phalangeals being apparently normal, so that this form 
belongs to DicyHodon. The same remark will apply to put 
skeleton from the same beds upon which the genus £iayattfm\ 
been founded. The genus Udeno^n {Oudeneden) is 
by the total absence of teeth (fig. 985, b), but is otbenricl 
closely allied to Dicynodon that it must certainly be included in I 
same family. The nares are somewhat approxiraated to the 1 ~ 
and the profile of the muzzle is rounded. It occurs in the 
system of the Cape Colony ; and some of its representatiia 
tained very large dimensions. CisUeephalus {JCist^xpiahu) 
prises smaller forms from the same beds, in which the skull is 1 
depressed, with the orbits directed frontally. There was a pair 1 
tusks in the maxillie. 

The name Platypodosaurus has been applied to a considenUl 
portion of the skeleton of a Dicynodont, from the Karoo systen^ il 
which the skull is unfortunately unknown, and which may prove K 
be identical with Udenodon, unless it belong to EndotkiodoH. Thl 
remarkably Mammalian structure of the pelvis, in which there i 
a small obturator foramen between the pubis and the ischium, i 
fully noticed in Sir R. Owen's description of the specimens. 

Family ENDOTHiODONTiD.t, — The remarkable genus EndMia 
odon, compri-sing large reptiles from the Karoo system of the Cape 
forms the type of a family distinguished from the preceding by thi 
presence of teeth on the jxilate. The skull presents a strong gen 
eral resembl.ince to that of Udenodon, but the muzzle is mow 
elongated, and the nares are terminal and overhung by the nnas.«in 
nasals. The alveolar borders of the jaws are trenchant, but the on 
surface of the palate and mandible carry one or more longitudina 
rows of columnar and cylindricil teeth. The remarkably Maio 
malian type of the palate of Endoihiodon is noteworthy. The skul 
from the same deposits described as Theriognathus seems to belong 
to Endothiodon. 



vtiJan has l>CT:n compared to the RhjmchoccphaJian JfAyncAa- 

iand also to Platodut; lioi although there is a marlced super- 

iblancc iKrtwccii the three forms in the palate and Iccth, 

is but apparent, since while in Endothiodon the 

borne 00 x MCOixlaT>' bony floor beneath the narial pa.-»dage, 

\VT geaesa they are wpponed on the pro[>er surface of the 

which the posterior lUrcs open directly, without the inter- 

of a scconclar)- passsgC: 

»itDEit 4. Piiocoi^raoKt&. — According 10 a recent ohserver 

los I*nxalefikam, represcnitxl by comparatively small forms 

Karoo system of the Cape, differs so decidedly from the 

satia that it is entitled to form a distinct .mhordcr, showing 

lignft of affinity with the Rhynchoccphalia. The yVoiw/a- 

have a full dentition, hut no tu<>k>likc teeth, and the narcs 

Although the pectoral girdle still has a distinct pre- 

|)ei its whole characters approximate to iho»e of the 

phalian genus SfikeaadMt. The humerui! also resembles 

3nding bone of the latter ; and in the skull the pterygoids 

forwmrds in the same manner to join the vomeni and exclude 

kpabuncs from the middle line ; while there are no secondary pos- 

: aarcs ; and teeth arc borne on both Uic pterygoids and vomers, 

be young of SfiJienodon. 

jr PljIcooontia. — Our sole knowledge of this group, repre- 
. by Plaadus {ftp 986) and Cyamcdus of the Middle Trias, or 
Ik of Germany, is derived from the skull, ko that we are 
IfRwot to a great extent in Uie dark 
wdieir true affinilieK. These formii, 
baling been rcf^rded as Ganoid 
icre referred by Sir R. Owen to 
' Sanroptcr^ia ; and the type j;cni» 
■bks the Noibosaurs in ihc back- 
. position of the nares and the form 
ibi ibe crania! rostrum. The skull has, 
hdccd, been said to present many points 
M mcmblancc to that of the Anomo- 
Awn, and more especially EmiotAiodaM, 
'to which genus tt i» considered by Sir 
. Owen to be closely allied. The re- 
in the form of the palate is, 
as ftlrcady mentioned, only a 
one ; the present form having 
r to the nana] passage, dnd the pos- 
iBMroarcs opening dircaly into the roof of the mouth by horizontal 
iVDiafei. u in the Sauropterygia. The skull is broad posteriorly, 
n^double naieSi a deep and apparently compound tempoial 

rio. gS6.— Till Impitrcct iialMc 
af Piiiotibu rigai : from lh< Mii>< 
chclkallt of TUyieuih Onc-fuinih 
iuliirsl*i>a. WticDtalircthe muule 
irouli] (otm a prvdiKBi) nniniin. 


arcade, and a postorbital bar. The palatal teeth {fig. 986) : 
paving-stones, and were probably adapted for crushing hndjj 
stances like the shells of Molluscs. In the upper jaw the 1 
are arranged in an outer or maxillary series of small ooe^i 
an inner or palatine series of larger ones ; all being imp 
shallow sockets and replaced by vertical successors. In the 1 
there is but one row of teeth. The number of palatal teeth ' 
the different forms ; and there are also modifications in the 1 
contour of these teeth, which aid in affording generic and 
characters. The premaxillary teeth may be of a more fJt\ 
prehensile type. Till the vertebrae and limb-bones are kaoR^ 
position of these forms must remain uncertain; but it 
remarked that all the known limb-bones from the Mu 
except those of Dinosauria, appear to be of a Sauropterygian \ 

In the typical genus Placadus the skull is comparatiw^y 1 
and has a long rostrum produced considerably in advance ti{ 
nares. The palatal teeth (fig. 986) have polygonal crowns, 
the palatine series being three in number on either side, and < 
approximated ; while the three premaxillary teeth are more <k I 
chisel-like, and are separated by an interval from thc^e on the ] 
The mandible has a long symphysis, and two pairs of cuttii 
The maxillary teeth may be either four or five on either 
Cyatnodus is readily distinguished by the great width and 
of the cranium, which has no distinct rostrum, with the nares ] 
at the muzzle and the premaxilUe fused together. The palatal I 
have rounded crowns, the crown of the last palatine being iq 
large ; there may be either two palatine and three maxillary, orIM 
maxillary and three palatine teeth. There were but two pain • 
premaxillary teeth, which are not chisel-like. In the Iowa jaw ih 
symphysis was triangular and comparatively short, and was proUbt 
devoid of teeth. 

Recently Dr Gurich has proposed the nanie Pleurodms for I 
allied form from the Muschelkalk of Silesia, but since this tera i 
preoccupied for a Crocodilian genus it will have to be changed 

Synaptosaurian Branch. — According to Dr Baur's sdiemei 
classification this branch comprises the orders Sauropterygia U 
Chelonia ; although Professor Cope and Mr Boulenger would iL 
include the Rhynchocephalia. The typical Froganosauria of I 
Baur may be merged in the Sauropterygia. Although the San 
pterygia and Chelonia present many characters in common, yet it 
not easy to give a definition of this branch. In all, however, t 
quadrate is firmly united to the skull ; and all, or nearly all, c^ t 
dorsal ribs articulate with the vertebrae by single heads. As a gei 
ral rule the palate is more or less completely closed, the pterygo: 
generally extending forwards to join the vomers. There may 



[one or two temporal arcades. A parietal foramen is present 

in the young. In all cases ossifications are developed upon 

aspect of the body, cither in the form of abtiominal ribs, 

|il«|||iiiii ; but there are none in the sclerotic of the eye. 

rib» are connected mlh the vertebra: by upper and lower 

i ; and when chevrons are present they are mainly or ex- 

' attached to the hinder borders of the caudal centra. A 

>id anchylo&ir^ to the scapula may be present in the pec- 

1e ; and in the pelvis the pubis and iechium have expanded 

led venlTsl surfaces, and the obturator foramen may be 

ted by the union of the ischium with the pubis of the sanic 

Tbtre is, moreover, a considerable structural resemblance 

the limb-bones of the more generaJiscd forms of the two 

these bones always having terminal epiphyses; and the 

in both may be of a very primitive type. The humerus may 

Ichber an eniepicondylar (ulnar) foramen and an ectepicondybir 

]) groove, or only the latter, or may be devoid of both. The 

[brrcT haAir uncinate processes. 

■ are still very much in the dark as lo the origin of these two 
altbodfb chc Sauiopterygia can be traced back in a form 
several Amphibian features, which appears lo have been 
' allied to the ptimicivc Rhynchocephalians. From the dis- 
ce of numerous segments in the verieliral column of the 
during development, I'rofessor Parker h^» 5ugj;esied that 
rorder has origin-ited from a type allied to the Sauropterygia ; 
tbdr plastron is almost certainly derived from, or developed 
, tbe abdominal ribs of a form allied eittiex to the Rhyncho- 
or to Mescrairuj. 
M II. Sauhoptekvuia. — In this extinct order the body was 
of any exoskclcton, while the neck was more or less clon- 
and the tail short. In the skull there is only the superior 
arcade ; the rurial apertures are lateral and inore or less 
Med to the orbits ; the prcmaxillx are very large ; and there 
; a weU-devdoped parietal foramen in the adulL llie prefrontal 
distinct; the posiorbiial may be separate from the past- 
bootil ; typically there is a transi-erse bone ; and the symphysis of 
Ik nundible is united by suture. The teeth, which are implanted 
B distinct socltcts and confined to the margins of the j.iw8,' have 
anod thaip crowns, with fluted enamel. Each rib articulates to a 
iogle vertebra, and in the cervical region the costal facets, which 
lay be either single or double, arc situated entirely on the centrum, 
■d generally are not prominent The vertebne are amphicmtous ; 
mI the ncunxentral suture may be either persistent throughout 

EjUcitiBing thai [be I'Ucodoolia ut dUtinct &am this order. 



life, or completely obliterated. All those vertebrse in whie 
costal articulation is on the centrum below the neuro-cestnl : 
may be reckoned as cervical ; their number varying from abt 
to nearly 40, The centrum of the atlas is well developed, and 
is a wedge-shaped intercentrum between the latter and the 
The true cervicals are succeeded by a few vertebrae in irtiic 
costal articulation is partly on the arch and partly on the ca 
for which the name of pectorals has been proposed. The ' 
vertebrje have the costal articulation placed entirely on dw 
and generally forming an elongated transverse process. The ( 
vertebrae are always furnished with true ribs, and also with dk 
bones, which may not be united below. The structure of dM 
toral girdle is very remarkable, and has given rise to consid 
diversity of opinion. In all forms the coracoids meet in a n 
symphysis, which may be short (fig. 987) or very long (6^ 

Fig. 9B7.— Ventral aspect or [he pectoral girdle of XalAmannu miraii/it: froa the I 
kalk of wilricmbcr^. KeduceJ. mt, Clavicle uid in(erclaTicl«i; te, SapuU: fl, 

cavity t ior, Coracoid- 

In the generalised Nothosaurus (fig. 987) the scapula has ■ 
small ventral portion, separated by a wide intert-al from that 
fellow, .interiorly to these ventral plates of the scapube 
is a slender arch consisting of a median and two latera 
tions, corresponding to a similarly situated bone in 
saurus. This arch is usually correlated with the interclaricl 
clavicles ; but from the deep-seated position of its represei 
in PUsiosauruSy Mr Hulke considers that in that genus it 
spends to the omosternum of the Amphibia, and if this ini 



It will hare the same homology in Notkosaurus} 

proceeds it appears ih-it the scapulse have tended 

ver^' larg« ventral plates, with a concomitant reduction 

:c dbappearance of the clavicular arch. The inter- 

itag« is shown in PUsiosaurus (fig. 997), where it will 

that the ^'cntral plates of the scapuhc are separated in 

lian lin« ; arxl the culmination in Cimoliosaunii, whc-rc ihcy 

X n>cdian sj-mphysis, and join the anterior extrvmilies of 

coidSf while the 

le ha» di!>ap- 

lilr Hulke regards 

filaie of the 

«s Tepret.eniiiig the 

of the Cbdo- 

t further evidence 

ircd to prove this ; 

that il rqinrscnts 

bcio^ obvioml)' 

In the pel vis 

lu usually form!) a 

te, while the isch- 

MDicwhat chopper- 

; in some cases the 

tod ischium of each 

jnite to enclose an 

(or foramen. Thc 

[ore strikingly like 

D( the Amphibia, and 

long symphysis, 

itu are subject to 

rrable variation : 

if the earlier Reneralised forms being adapted for progrcs- 

land, while in the specialised typen they are modified 

iddles. In all cases, however, the limhs are readily dislin- 

I liom those of the Ichihyoptcrygia hy the iclalivdy longer 

IS and femur, and the absence of inicrdiyital bones. 'Ilie 

o arc regarded by Professor Seelcy as showing signs of 

lan affinity. A peculiar feature in the liinb-l>one!i is that 

ihyses (Rg. 9S9) of the bumcru!. and femur arc enomiDu.sly 

and form btf;e cones at either extremity of the bones, 

or completely, nieelin^ in the middle of the shaft, which 

to 3 i>air of elongated cups. The bones of the palate 

Brllbh Mmnin Catalogue of Fo»i] Repiitia Mr HuIIec'n iniciprtta- 
hmnolovr <>f ^cx bonca wu _provi)iion«Ily iiOopted, bui ilic writer 
I that ibc MhcT inlerpr<Ulioa U p(obkt>t]> lli« Irue one. 

rriiri.ii i>(«L[ ot ihi |i»'.1onil ^irjlt rA 
• Iv' ) ''■-' ^•••^'riui ; fmm 'lit KtlneriJijo 
lai. Kniuunl. «, Stavulit . #t, Vtniril (prrcuni- 
ikI^IJ fixin <3r<!o. ' £i, C^lcnoiii <^k^iiy; /^ SinpulAr 
fMontca; (V, C9iS(«)d. lATtct Kulkb) 



never develop plates to form a floor to the nasal passage 
the posterior nares also open directly into the mouth by ban 
apertures (fig. 991). 

This order ranges in time probably from the Pennian ind 
tainly from the Trias to the Upper Chalk, and we are 
trace the gradual evolution of the specialised marine fonu 
those less widely separated from a normal t]rpe. All these n 
appear to have been carnivorous. 

Family Mesosaorid* — The genus Mesosaurus, origim^ 
scribed from the Karoo system of Griqualand in South A 
which is probably of lower Mesozoic age, but subsequently t 
in beds of uncertain age in Brazil, and described under the oa 
of Stertosternum, includes small reptiles regarded by Dr BuH 



FijZ- 989- — LongLludinnI Hctton 
of a Sauroplerygian humerus ; 
from ihe K-imendge Cl»y ; one- 
sixih natural liie, a. PnjxiiDal, 
b. Distal epiphysi-. ; c, Shah- 

fig. 950.— Ventisl upcct of tlw 
left pftCEora] limb of Mta^tmmrmi 
ttHHjdens: from thr Katod KVllcPi 
of GritiuaUnd- eni-/t EotepKoA- 
d^lar foramen of humcruii r, Ra- 
dtun ; V, Ulna. 

constituting a distinct order — the Prc^anosauria, — but «h 
appear to be so closely related to the Nothosaurida, that there 1 
be little, if any, hesitation in including them in the same on 
The Brazilian form was originally referred with some hesitation 
Professor Cope to the Amphibia. One of the most peculiar i 
ures of this genus, in which it differs from all other groups ax. 
the Amphibia, the extinct Palaohatieria^ and perhaps the Chelc 
is the separation of the fourth and fifth tarsalia, so that each m 
tarsal articulates with a distinct tarsale. The centra of the verte 

See Rhjmchocephalift, infra^ 


. miAll notochordal cannl, :ind ue small in comparison lo the 

, arches j while the ribs seem to have been anchyloscd la the 

and were of great thickness, like those of the next familj-. 

ffjfstcm of abdominal ribii was strongly developed. The skull 

elongated, and has slender recurved teeth, which were in 

jiltly implanted in distinct atvcoti. The pectoral girdle 

lo be very similar to that or the Niytltasaurjiiz, the pre- 

iDtciclavicle not being T-shaped ; and a similar cloae re- 

is presented by the pectoral limb (%. 990), in which 

Flnuncnis has an cntcpicondylar, or ulnar, foramen, like that of 

The pelvis is considered to have liad only a very 

obturator foramen ; the ischia and pubes forming broad 

plates like those of Amphibia and other Sauroptcrygia, 

tenninal digits vere devoid of claws ; and the feet were prob- 

fwdibed like those of frogs. 

Baur make) this genus the tf^ of the order Froganouiuria, 

he would also include the undermentioned genus Palao- 

; and regards this order as the connecting-link between .A.m. 

and Reptiles^ Although there is something to be siitd in 

of this view, yet the manifest affinity of Affsosnnn/t to tfic 

typical Sauroptcn-gia, and of PalxuhaiHria to the Rhyncho- 



I)*-— PatitaJ ■^MCt of ibe tnniNm ti SfSkemmrma mlrmtiUt ; from the MuHhclkalk. 
Itt MlanI tki. /■&«, ProDum*: inar. Pmenni nun: tv. Vomer : >■■>, Manilla ; 
Wiaj (*, FurnoU; «, Ala tt ia.: t. Owuliuic riditc uf lame , im, IJuoJiait ; it, 
I MHjn. Tta« powttiw uircnlly cX f»I U piabobly fiamtA by k dtitini:! Inninnc 

seem to render it more advisable to refer those genera 
tt die two orders in question, of which they will tcspccttvely form 
tbe most generalised stage. By this arrangement the intimate con- 
Mction of both orders with the Amphibia will be made manifest. 

Family NorHOSAURiD>E. — In this family, which comprises some 
foma of large size, the limbs were fumiihed with claws, and adapted 
to a certain extent for walking. In the &lcu[I tlie plerygoidi either 




diverged posteriorly, or gave off wings uniting in the 
upon the basi- and presphenoid, and thus completely 
posterior portion of the palate (fig. 991) — an approximatioB 1 
latter arrangement occurring in some Chelonia. The palili| 
also devoid of infraorbital vacuities. The coiacoids (fig. 987) 
a short median symphysis, not extending as far forwardi 
scapular articulation, and also had a groove ; while the 
plates of the scapulae were very small. Typically, the 
vertebrae have double costal facets, while the transverse ; 
of the dorsals are very short, and remarkable for the vertical 1 
tion of their articular faces. The humerus and femur are 1 
— the former (fig. 993) having an entepicondylar foramen, 
distinct distal expansion. The ischium and pubis did not 1 
enclose an obturator foramen. 

The known forms occur typically in the Muschdkalk, or 
Trias, of the Continent, but some of them range op into 
overlying Keuper, and one species is found in the Buiiter,< 
Lower Trias. In the typical genus Notkosaunts the sktiU 
991, 992) is long and much depressed, the length of the 
orbital exceeding that of the preorbital portion. The 

Fig. '/ji. — Ki^lit ]:iicr3i a>pn:i uf ihe ukull oT Xolkesamia 
mirnh'/h, ReLlui:iHl- (After .Ue^'cr.) 

rig- «}. — VeMnl •■ 
peel o( ihc rishi bnant 
of Cnt-'kiHrnmna. Ok- 
half latuial (iac. £■!/'■ 
Enlcpicandytu' lisnaa '• 
a, EcUtpicoadylar p««- 

of the palate are shown in fig. 991. Cenekimaurus is a closely 
allied but smaller form, in which the teeth are club-shaped. In 
Sinmsaurus the skull is characterised by the breadth of the facial 
portion and its stout teeth. The type sijecies is nearly equal in 
dimensions to the larger species of Noihosaurvs. Finally, Pisto- 



duOfiguished hy the posterior divergence of the ptciygoids, 

improbably by ihe luescnct; of long tran.wcrsc processes to 

vcrtehne; both of these features heing Plesiosaumn. 

ital ponion of the skull is very narrow. Although the 

of this family have lost all Iracen of a notochordal cannl 

centra of the rcttcbrx, yet the ossification of ihtse centra 

en place in the same manner by means of a sheath investing 

ii-T IjuuotAumD*. — ^TTiU family is closely allied to the 
ig. but the limbs approximate to those of the Pktiosaurida. 
dnill, a% least in one genus, has infraorbital vacuities on the 
v and the cowcoid has no notch at its gWnoidal extremity, 
femur alway* remains an elongated bone, longer than the 
>s and metapodials collectively; but the humerus maybe 
ly short, and is more or less expanded at its distal 
;y ; while in Lariogaurus, although not in the other forms, 
tost its fofotncn. The terminal pbalangcah of the pes still 
their clflws. In NeuslUomurui it is thought that cen*ical 
■mcK waniii^. The type genus Larinsaurus comprists one 
species from the Trias of Lombardy, and according to 
Bht, has both limbs adapted for walking. The pabtc is un- 
In yhulitnsimrus, from the Lettenkohlc at the base of the 
of Wiirtcmbcrg, Professor Scelcy considers that the [xrctoral 
bob had become modified into a paddle, although this conclusion 
is tKA aa:eptcd hy Dr Baur. It appears probable that a small 
Itptilc described from the Trias of Italy under the preoccupied 
■■K<]f pA(hypkMra, is not more than spcciliailiy sepMrable from 
^HB^aaaxirus. The type species of the latter was about one foot 
ia ki^h. and vsa probably of amphibious habiU. 

It ony be convenient lo notice here two ttmall Trlassic reptiles 
rtich are referred by Dr Baur to this family, although Dr Dccckc 
ramdcrs that at least the second has more affinity with the Lizards. 
TkeM genera arc Daetylosanrus, from the of Silesia, 
lad Matrvmerataurus, from the lulian Trias; the former being 
Jteon eettainl/ referable to this order, and perhaps not separable 
ficni Xfustuosaarus. In this connection it should be observed that 
Dr Bassani consider Afa/r^nurotaums to be identical with the type 
|Kdo of iMriotaurus, and that NtHsticosimms is not generir^illy 
qwible from the latter. PaekypUura is, however, regarded as 
Ertbict from Neuslieosaunis, in which case it will reciuire a new 
tme, as the present one is preoccupied. 

Finally, it should olso be mentioned that some authorities would 
gaid the two preceding families as constituting a distinct sub- 
der — the Nolhosauria — but the transition to the next family is 
sost oomplece. 



Family Plesiosaurid*. — With this group we come to the 
iideration of the typical members of the order, all of vhidi 
idapted for a purely aquatic life, and probably frequented c 
and estuaries. In the skull the pterygoidi 
verge posteriorly, and do not overlie the 
sphenoid ; while there were small infhud 
vacuities in the palate. The dorsal 
have long transverse processes. In the pectt 
girdle the scapulae have large ventral pM 
which may meet in the middle line ; and 
symphysis of the coracoids is much el 
and extends in advance of the scapular 
tion. In the limbs (fig. 994) the humenn 
femur were comparatively short and dista% d 
panded ; the former being devoid of a fona^ 
The bones of the second segment are likari^ 
very short, and strangely altered from the nom 
form. In certain cases, moreover, a third bM 
(fig. 998) articulates with the humerus a 
femur, of which the homology will be discoM 
under the head of the Ichtbyopterygia. Tb 
metacarpals and phalangeals are, however, SJ 
elongated, but the number of the latter is ii 
creased beyond the normal complement Further, the tennim 
claws have disappeared ; and the whole limbs were doubtless ta 
veloped in a common integument, to form paddles after the fashio 
of the turtles. The coracoid (fig. 988) is remarkable for itspei 

Fig. i»04. — DormI u- 
peci of tbt left pccioral 
limb of Pltthtannu 
Havikiiu! ; from ihe 
Lower Luu or Dortci- 
(birc. Rcduced.B, Hu- 
merus ; bj Radiui ; f » 

I^iS- 995- — SVtXtXoa of PUiitiamriH dvlicludinH : from the Lower Liai. CnUlyndKi'- 
(After Conybeare.) 

antero-posterior length, and has no fontanella This family includ 
some forms of huge dimensions ; its range extending from, t) 
Rhxtic, or Uppermost Trias, to the Chalk. 

The Pkiiosaurida, like the Crocodiles of the present day, dif 
greatly among themselves in the relative length of the mandibu 
symphysis, as is shown in the accompanying woodcut In 1 
present family it appears, however, that the result of evolution a 
specialisation has been towards the gradual lengthening of this sj 



wbcfeos imong the Crocodilia the tendcnc)* has l>ccn prc- 

I the ^>pOMte direction. 

fiiniiljr hu been divided inio a lai^e number of genera, but 
Evera) of lh<:f« sre not really distinct it mil suffice to adopt a 

number of such divi»ion». The genuti Pirsieiatms, as now 
cd, JB occlusiTely confined to the Upjier Trias (Rhaalic) and 
Owing to the beautiful prescrvatioD of many of the species 


I '"I 




L— TW NBUi4iUa in i(df«*cit (>nira nl PUtiaMmridr. Rtdveiij. >.. PiltniHtUi 
~ iM ikt Ctafotd Clari «iW'<i<tiEh naiunl tat : ■> T^<im»li>airta imiifta, (mm 
• jMfcBJc of India. oncMvvnih cuiunl lin ; (, PInUiutnu iteJitAKtirti, lunvlirifat 

3US has been long known lo science; and its remains were 
ibly dcwn'bcd in the first third of the present century by the 
[r Conybtare and Ilean Huckhnd, who with remarlcabte fore- 
nnied at the affinity of these strange and weird forms of Kep* 
the Chelonia. In this genus the skuil is either small 
mandibular symphysis (as in fig. g<)6, c), or moderately 
longer syniph)-siK and rostrum. The teeth are generally 
r, without carins, and the terminal ones are not much larger 



than the others in ihe more typical species. The nedt is' 
Xcss elongated, with the iintcrior vertebra: in mtMt cases vec 

The cen'ical ribs i 
ral aichcs arc (umlj 
lateil to llic centn 
Ycttcbrfc, but tracd 
sutuic usual)}' per&ii^ 
ccmcal region the 13 
the articulation of Ui 
nre u>ually double, M 
Icrminal liices of ihtr 
generally ellipsoidal, 
more or less cupped, 
meet characteristic U 
uf the genus are, ho 
tu be found in the pi 
ijirdlc (lis- 99 7). in 
the scapulx are nl 
small, and widely sep 
... ,,..j. , ,- , -,. , in the middle lir 

lOoneuhirc Kcducrd. ivMni<rvU>i(Ir; (T, Su>;r^ thcy FCSt Upon 

oChMVRu*. (AftnH«ik«.) Clavicle, With lu 

deep notch. The 
arc, moreover, long and rather narrow, with a median prod 
in advance of the glenoid cavity ; while the foraitKn bt 
Ihe coracoid and scapula is very lar;ge and open towan] 

fit- «g" — Veninl »Hp<v.T o( Ihi retto"*' l'/i,l'« i*! 


This genus may be divided into three groups. The Laagir 
(tronp. represented by P. rastialHs of the Lower and /*. iimpni 
the t.'pper I.i»», is cliariicieriscd by the comparatively cIongBted 
dibuhr ^yinphyiiis and ilie extremely slion neural spines and 
costal facets of the majoriiy of (he cervical vcricbra:. in the 
grotip we have /'. do/icAotlirus with its extremely long neck (fie. 90 
ihortcr- necked A ffmc/kinti, and ihc large P. Cimy^iri. all ocin 
the Lower Lias. These forms have a mnderaicly snon inandibula 
phy»» (fig. 996, r), while the cenita of the ceniral vcncbrje are 
Crc.itly eloiiKaicd, nnd there are double costal facets and moderai> 
neural spines in this part of the vcriebtal coliimn. The third g 
rcprcwnted only by /'. homuiospintlylus, of ihc Upper Lias. "1 
characterised by the great elongation of the tcnira nf the cer%ic 
tebne, which have flat terminal face^, and cnonnoutly tall nctiniM 
The resemblfince of these vertebne to those iif the cerxicah of the 
groupof the JiirauLC and Cretaceous gcnm Cimo/iouiMtuj sugge 
origin of the latter group from the present ; whereas on similar g 
it may be suggested thai the Ca-fm/H'mfr/iiu group of Cim.>//o!Jii/ 
originated from the tvpical group of /'(ru'osjurus. In all species 
present genus the radius and ulna (fig. 994} still retain evidence c 


rier as long bones, and are scparatfd by a well- ran rke<l 

exclusively Liassic and perhaps RhKtic genus is Ertt- 
wbicfa had a long neck and probablj' a small head lilce 

nwtu, bu( with a vtrv different n-pe of peciorfil girdlt. Thus 

icoids had no median production in advance of the glenoid 

vhSe ih« scaputx were brf^ and articulattd togcihcr in ihc 

fine, and posterwrty were united by their whole length wilh 

muU, leaving only very minute coracotdal fornmina. If an 

ride were present it had 

fased irith th« scapula.-. 

fce ipedcs is from the 

Lias, liut thiie is anothci 

ippcT Lias. The largest 

rf the family is. however, 

Mw^KJ, in which we may 

ently include those fonns 

td under the naiue» of 

smj, ElatoMsaarus, Man- 

I Ptfytotylm. Muricm'sau- 

d ColymhoiaHrui. This 

was originally described 

fce diidcncc of a very 

«cies (rom the CrcUceous 

I Jersey, with which Dis- 

v, and probably Elastn»- 
are flpeciAcatly identical. 

lew Zealand CTeiaoeouit 

i deMTibed as Mamsaums 

fly allied ; and it has yet 

ro>'cd that the lyfic species 
■pecifirally distinct from 

C. censlricftts. Many of the other species differ con- 

Ij (roQi these fotnw, but if generic divisions are once 

: 5ccm5 impossible to know when to stop. 

genus in the abo\'e extendtxt series may be characterised as 
I1ic tcvlh and akull arc relatively small, the niatidibular 
i* K short, and the neck usually very long, with the anierior 
Iv >malL The vertebrae arc more or leas elongated, 
' >i' ihc neural arches and ihc cervical rilts completely 
to the centra in the adult ; the costal articulations always 
iin(;kr facets in the cervical region- In the pectoral girdle 
) the scapulte ha%-e very larjte and wide ventral platL-s, meeting 
iddle tine, without any trace of an tntcrclavicle, and usually 
do«rn a median procc&s to join the ooracoids, and thus com- 

KlH- M^^ ~ VcDtnl aviHtc of (be rljht 
if^sA Cimtiitimnrui IntAaiiftriiLt ; Iriini Ui« 
(Afler llulkc.) 

pletely closing the acapulo-concoidal foramen.' The 
998) is usually longer than the femur ; and both these el 
arlicubte distally with cither two or three (figs. 998, 9- 
Wnoi, which in the latct instKiiccs Icnc all resemblance to 
mal elongated form. The i&chia arc relativel)' short. 

Thit eenut may be divided into two ^'roups, accordinit ta »h< 
venebnl centra have nearly flat or deeply cupped cenira. In tU 
group we have the inrgt C. trtiMentUf ni the Kimerid^e, and tbs 
C. fin'Mfui {fifi. iooo^/j)nnd C. fiuhardii)Mi{fig. iooo)of the OxS^ 
in both of which the humerus aniculaic* only with the radios a^ 
the latter ^prri^'t being di«tinpui«hed hy it» shorter cervical 

Fi([.TO-— VciHtnl ii>mi ui (.1111 uf iht jitni jitUii; limli 
of Ctmfiliatmmrtii ptirfisneinMi ; from thv Finland Oolite. 
Kaducsd. T. Pnuist ; f. VvtmtM banl«r; i^Tfliiai 
it, ribulB: ^'. >li>inuli>K.ii*'ir ihtpnifonn: /jlTtbiiiie: 
tit Jntflrmcdiuin ; t j, Filiulbv i / I'OftiajiifeL tkrv«t. 
(After Kiilkc> 

pen «l Hit (_ 
Vtaon^ liMb of 
MMnu KtakariSt 
itK Ontcri CU; 
vichlh naiuni i 

tjlAix: r, KadiMil 

r. UwiLii.: i. 

diuin; It, Ulttitf<i 

In C. p^rl/ofuiuHt, o( the Fortknd Oolite ana Purfacck, the 
three tliorl bones articulating with the humerus and femur (6^. f 
» in the Crctnccous, however, that «c meet with the largest rep 
tivu of this K'^'^^P' vrhich comprise C. iimstrittuj, of the Europei 
and Chalk ; C. vetusiiit and C, pltUyurus, of the Crclaccoiu 
America ; and C. ffiuuUt in the corresponding strata of New 1 
These were caormous reptiles, with an estimated lengib of beq 

' Ocewionalty, a» in C. iturvMpeaa't, of the Oxfoid CUf, ihb bi 
ftUent, l>ni it is not known wh«th«r [hit it on'y an indivklual pecMliariif 



leet. aod havinf[ ni^arly 40 nn'icil verlebrx. In the sMond 

wfaicD apparently corresponds to the genus Polyc&tylus of Pro- 

" Spc, *« e have, in lh« Oxford CUy, C. oxoHirmis and C euryments^ 

Ui« ituntenif ^iniculatcs with only lh« radius and ulna, and in 

Igt the larger C. troekanttrius, in which the pisiform also joins 
I (fin- W8)« Ttc rcry imatl C valdiHiis, uf the Wcaldcn, niav 
rireibwaicr habita [ wliilc in the Chalk wc find the latter C. 
This group apfwan to Ua^-e been aho represented in the 
of North Anicrii:a and New 'IksXsmA. A specie» from the 
of K^n^A^, described under the name 
fpKy has three bonca anicubtinK 

imperfectly known gcnut Pohptythodom 
■), or the Middle and Upper Cre^ 
of Europe-, appeaiK to be allied (o 
\rits, but with a relatively liujjtr 
nd teeth, and probably with a much 
Deck. The teeth, which are very 
in the Camtiridge (ircrnsand, luve 
crowns, with strongly - marked 
(Ch a considerable numlicr gene- 
llhort of the humniit. These Kcp- 
Biut liav-e attained huge diincnuonti. 
ncra wc have now to consider indicate 
ent branch from that to which the two 
ling gcnvra belong. The lirst of these 
\MmaiasaMrui (in which JihitmalenSiturus 
e iocluded), typically occnrring in the 
Oolite of Wuncrabcrg, but extending 
iard» to the Lower Lias and upwards 
Kitncridgc Clay of England. In these 
tbc skull and teeth were relatively large, the latter being 
carirutcd. and the mandibular symphysis (Rg. 996, ti) com- 
ly short, with the finX. five ur six teeth enlarged. The neck 
■1, IL M 

wtl vtpmi -jfib^ left ^^IvK 
^rcJkmai {torn the Oxford 

CUy. OdE'iiith iiniiinil 
■inc. />. L>i<ml h>1f i>r 
feimsr ; r.Titiia./ Fihufi; 
r,TiInnlE: i, Intrrmeiliiim : 
/". fibuUrc. The libia It 
drtiRi raihar ion unialJ in 
pniiarlion I9 th« Abulo. 



was short, and the cervical vertebrse have comparatively short 
distinctly cupped subcylindrical centra, carrying dooUe ( 
facets ; while the arches and cervical ribs were firmly arttcuhl 
the centra. In the pectoral girdle the scapulae and coraaadi 
of the general type of those of PUsiosaurus ; but the cl» 


Fig. looi. — Anterior part of the pectoral Rirdle q{ PtlmtfUtii /Ailardkiu: fron dt 
Clay. RedDced. «»>, iDlcrclavicle ; le. Scapula ; /.ur, Vcnlnl plate of do. Thi 
aspect a ibatm. 

arch was greatly elongated transversely, and was probably 
lapped by the scapulae. The humerus was longer or shortei 
the femur, and articulated only with the radius and ulna, 
were considerably elongated, and separated by a distinct ini 
This genus is represented in the Lower Lias by T. megacef 
and T. arcuatus, and in the Upper Lias by the gigantic 7! C 
toni, which attained a length of some twent) 
The type species, only known by detached 
and vertebrae, occurs in the Great Oolite o 
Continent ; while T. indicus (fig, 996, b) is I 
in the Upper Jurassic of India. 

In the genus Peloneustes, of the Oxford 
Kimeridge Clays, the coracoids (fig. 1601) di 
appear to have been produced anteriorly u 
middle line ; while the scapulae have their v« 
surface broad and flat, and the dorsal surfat 
duced (fig. 1002). The same figure also shon 
extremely small size of the interclavicle. The : 
were very long. The mandibular symphyaf 
996, c) is greatly elongated ; but the vertebrae 
with those of Tkaumatosaurvs in the firm a 
ment of the arches and cervical ribs to the o 
although the centra themselves have the terminal faces fi 
and with a transversely elliptical and somewhat angulated coi 
The radius and ulna are nearly as broad as long, and have 
a very small interval between them. The most specialised ] 
of this branch is PHosaurus {Ischyrodcn, Spondyhsaurui or 

Fig. 1003. — Crown 
of a looih of /'//tf- 
taurus hroihytiirus ; 
from the Kimeridge 
Clay of Elv. One. 
hat) natural stie. 



t\ wluch thus occupies a Mtnen-hai Mtnilar po&ition to the 
'd by Fofyptftkfi^n in the preceding branch. This genus 
-cnttfd Itjr scvctsl species of gigantic Reptiles ranging from 
' -rd lo the Kimrndgc Oayi but of which derived remains 
> found in the Lower (Ireeiisand of I'ollon, in licdford- 
Thc akull and tcclh (fig. 1003) are relatively very large ; 
fbntwT having a somewhat shorter mandibular symphysis than 
\^tiotiemsta. The teeth are caiinatcd, and in the Kiineridgian 
ibe space between the two carina is nearly flat, and 






•M^ — DUsnmef ibftaniralwpKtoTlW p*lTic ipnll* ofa jouns nioaur. Th* oppw 
M 4fv tha prtio. uid ih* lonw the iichia. Tti ib* iidiill Ihr tichia bcoomt moic tlanioiRl, 
\mm faMtf \kMmh ilioaU \un% Imm plnoHJ tnon oody |>i.>*lkl- (Afiu Phill>|H-} 

of ridees. The neck is very short, and all the cervical 

(fig. 1005) are relatively large, with nearly flat terminal 

to the Teiy short centra, which in the anterior region have 

distiixt and often very prominent costal facets. A peculiar 

Jorc of all the vertebra is, that the arches were only articu- 

bud to the centra by cartilage, so (hat they aru aln'a\'s found 

detached. 'Ilic pectoral girdle was of the general type of that 

of Pti»tuusUt, but it is not improbahie that the inieTLla\'ic!e was 

abtcnt. 'llic humerus was ahottcr than the femur, and tlie 

and ulna in the Kimeridgian forms have become much 



shorter than in that genus, and have scarcely any intent 
space, but in one of the species from the Oxfoni Oay 
have the same form as in the latter. The general airangeme 
the ventral bones of the pelvis is shown in the woodcut 1 
huge P. maeromerus, of the Kimeridge Clay, the length o 
lower jaw was nearly six feet, and that of the femur one yai 
which some estimate can be formed of the gigantic dimei 
attained by the entire animal Pliosaurs were widely distri 
over Europe, and have been described from England, Ge 
(as Ischyrodon), France (as Liopleurodon), and Russia (as ^ 
saurus). No remains of this genus have, however, been hi 

Fig. 1005.-^ Anterior (i) Mid latenl (i) Mp^u of an anterior ccrviol vnubra of PSm 
vacromena ; from the Kimeridge Clay. One-fifth nalml liie. (Alter FUG|» 

recorded from America. The less specialised characters 
Oxfordian forms, as shown by the structure of the teeth, an 
longer radius and ulna, indicate affinity with Peloneustes. 

Finally, it may be mentioned that in addition to the i 
already recorded the terms Piptomerus, Orophosaurus, and 
naules have been applied by Professor Cope to Sauropterj'gia 
mains from the Cretaceous of North America ; while a tooth 
the Kimeridgian of France, described under the name of Ha 
saurus, has likewise been shown to belong to this order, alth 
originally regarded as Crocodilian: 

Order III. Cheloma. — With the Tortoises, Turtles, and 
allies, we enter upon the consideration of the first of the ex 
orders of Reptiles. In this order the cervical and dorsal ver 
are not numerous ; the body is short and wide, and has a m( 
less complete bony shell, of which the ventral part, or plastron. 
sists of few elements of dermal origin, while the dorsal, or cars 
may be in great part of endoskeletal origin. There is gener 
horny epidermal exoskeleton. The skull may occasionally hav 
temporal arcades, but more generally only the lower one is pi 
(fig. 1007), and in some cases even that may be absent. The 
(fig. 1024) are single and terminal ; the premaxillae very small 
there is no parietal foramen in the adult ; but there is a di 



bone (f^. 1034). The deninry hones of the mandible 

iy fused logelher ; the postorbilal is welded with ihc posl- 

, wd nsuall]' the jirefronlal with the nasal ; while the supmoc- 

I h pcolongcd backwards. Tlicre is no transviTsc bone. The 

k completely dosed by the junction of the pteij-goids with 

ihcnoid, and often with one another {fig. 10 1 7 ^is). In all 

forms teeth are absent, and the trenchant jaws ensheathed in 

Each rib articulates at the junction of two vertebrae ; there are 

■>!■ Vm*ni mpKt of dw Aiknm of a rovnt Lasginlitad Tunle (Timlmtitrit^i 

j»4Alh« > b n roB m i n ii ,J . Mnth I«di>«*il< Th* tnrscwduKi prwooncoMl e/appoaM 

^Mvclo* wfi tMuu J M «fck)y lepafaud (ran OM anotho'. (AftwOiMn.) 

■aribsi in the cerrical region, and no transvcriie processes to the dor- 
BnitthneL The verlehne may have proccelous, opisthoctelous, 01 
P^ifaiazlotis centra in difTercnt parts of the column of the same in- 
itHMaiL In the pectoral girdle (fig. 1008), which i^ situated within 
Ibe ribs, the coracoid is the widest of the three earlilagc bones ; and 
he comcoid and ptecoracoid of opposite sides arc respectively con- 
iccted in the middle line by ligamentous tissue only. The con- 
betireen the scapula and piecoracoid is short, and there is 



no trace of a sternum. In the plastron (fig. 1009), derdo 
the ventral aspect, the epiplastrals {es) and enioplastral (r) 
spond to the three plates of the Labyrintbodont thoracic 

Fig. iix>7.^Lcfl lalcral wpecl of (he akuU oT Tttuda. ^, Pmnuilla ; ■>, ll>nIli,:iC 
Prerronial and naul \ /, Fronu] ; *, Pui«uU ; 0, SupraoccipiEal ; 1, }v%m\, behind '■4iich alw 
quadratojugHl, >nd above ihe pcnlfninlal : (, Quidnte, ibowitiE the incmnpletenciB of tbnf* 
pinic ring poiierioily ; the bone above thiiring in the muMnowT 

and apparently represent the clavicles and interclavicle. Th« 
arc in addition paired hyo-, ^po-, and xiphiplastrals ; and in seme 
forms (fig. 10 12) there are meso^oitnii 
intercalated between the hyo- and hypo- 
plastrals. The pelvic, like the pectonl 
girdle, becomes in the adult placed on 
the inner side of the ribs, and has the 
pubis much larger than the ischium (Gg. 
1006). The two latter may be anchjdosed 
to the xiphiplastral, and the obtuiator 
notch may be converted into a forameo. 
The humerus has an ectepicondylar groovy 
which is occasionally converted into a foci- 
men. There is a centrale in the carpus 
(fig. 829), and in the Oulydrida also in 
the tarsus. The tarsus may probably be 
regarded as having five distinct tarsalia, 
as in the Amphibia and Mesosamrta. In 
both the manus and pes there are almys 
five digits, with a variable number of 
phalangeals. Not unfrequently the bones of the palate develop 
inferior plates to floor the nasal passage, and thus produce second- 
ary posterior nares with a vertical aperture. Since the paUeonto> 

Fig. itoa.— Right ikie of the 
pectoral gitrlle of Tuttida. i. 
Scapula ; a, PrcCDracoid ; e, 



; generally has to d«al witb what ii rrequently and convenientljr 

the tMelit — that i5» the carapace and plascron together,^ 

must b« more p»niculArly directed to its structure in the 

tTi»iral forms. Commencing with the epidcmial skulctun of 

ronn« in which this '\s fully developed, wc find that il con- 

^ homy shields, which uisualljr have their cdjjcs in apposi- 

bui occa^onally overlap (fig. tost). On the upper surface 

astt five vetitimj shields in the middle line, on cither »idc of 

there are four eostait ; while die horder hasi 24 or 25 smaller 

of which the median anterior one (when present) is termed 

Id/, and the double or single posterior shields ore known 




d4; •. fintMlutnl (inUfUanUc); At. KyoplocnU i fi, HyiMirUiUal i ^i, Xl|i)>cpUi- 


x^rmidats; the remaining eleven on either side being reckoned 
marximais. The position of all these shields is exhibited in 
1017. On the ventral aspect of the phstron (fig. 1016)1 
are Bsually six pain of shields, of which the most iintcrior 
tenned gtlars, the nesti kumtrah {pattguiari)\ the next pedo- 
<: then the ahdominals ; \Smi Jemorait : and finally the anah. 
iome cases, howe\'er, there is an additional infer^^ar (very 
ly doul>le), which is usually (as in fig. loii) placed between 
gttlan, but in some eases may be situated below the gulars, 
(KtBg then surrounded by the gulars, humerals, and pectorals, 
as in Ckehdina. In the extinct Anhaoihtiys, of the Wealdcn, 
■here the intergular occupies the latter position, there is aLto a 
fcries of apparently single inkrftHerai, inleraMominal, and fnttT' 



/emaral shieids dividing the normal pairs of plastral shidds; 
is probable that this maybe regarded as the archaic type; 
in a Chelonian, mentioned below under the name of 
there is a median series of intervertebral shields dividing the i 
vertebrals into two lateral rows. In some cases tmj 
shields separate the marginal from the plastntl shields. 

The bones of the carapace, although following the same j 
arrangement, do not, as will be seen from the figures, by any i 

Fig, loog ill. — Upper surface of the ampuct of Thmlaaetktljn cmrwHti, with ibc 

ftbklH^ removed- Ine dark Lines in the middle region indicaie theboundvici of the*] 
the Ainuou^ one* the sitturet between the bone*, hm, Kucbal bone ; Pf^ t*yv^ <to. ; tbc 
above thitk markcil n and the next are the HuprapyEa]% \ above ihese^ tbe eight nrtnli, <£\ 
the Krft it tnarked n; c,c, Cottal bonea; m, m, Marginal do. i v,v, Vcnsbni ihiilili Hi 
reduced. (After Owen.) 

correspond with the overlying shields. In those forms with 3 wdt 
developed carapace there are typically (fig. 1017) eight median 
neural bones formed by the expansion of the spines of the dorsal 
vertebrae (fig. loio) ; these being preceded by a miclial, and fol- 
lowed by two or more suprapygah and a pygal, all of which have 
no connection with the vertebne. While, however, the nuchal is a 
cartilage bone, the pygal and suprapygals are of purely dermal origin. 
On either side of this median row is a series c^ from seven to nine 



k (fi^. 1 009 /Us) fomted by the developnien t of a plate on the 
of ettch rib (fig. loto) ; whil« the sidc'$ of the carapace 
by the eleven margfHa/s,' which arc dermal ossifica- 
. eigfat of which receive the estreinities of the ribs from the 
the ninth. In all young individuals, and in many of the 
funics (fig. toagiis), the cosul hones do not extend to the 
of the nhs, and consequently leave vacuities on the inner 
marginals, but in the Lind Tonoiseft and their allies (fig. 
carapace ia entirely bony. In some instances, agAin, the 
Uof neural bones may be reduced (as in lig. 101.4, where 
jie but M.-ven), and very rarely they aic entirely wanting, so 
fhe cosiaix meet in the middle line. 1'he two suptupygals 
be reduced to oitc, as in fig. 1014. Similar variations 
k respect to the degree of ossification of the plastron ; bince 

— TManMWMdiMi Hiroagh Un thdl cfC-WkwnurfMi. Koluceil. c.Vttitbtti 
. bjaodad Manl iplne; r. Rib; d, ComJ 1«mi m. Uarsuul do.; A flutroo. 

|d idl ytnitig individuals, and in the existing marine forms (fig. 
i^iroagbout a great pan or the whole of life there are vacuities 
^ the bones, in the land Tortoiaea and their allies the bones 
teotuKCted by suture. 

DC marine Tunics the plastron is totally uneonneaed witli 
p|Hoe; but in most other forms the hyo- and hypoplastrak 
P lotigcr or shorter peduncles underlyinj^ the marginal, and 
le cues also the cosials of the carapace, and thus form well- 
il axiiiary and inguinal luitrtsses. These peduncles are 
; in some of the exiaing Pleurodira ; while the inward protn* 
' the buttresses t& most marked in the Indian BaUigurs. 

3\t variation occurs in the &lcuUs of the dilfi^rent groups, 
I only be mentioned here that in &oiiie genera like Ckdydra 
^leiome the supntemporal fossa is more or les« rnniplctcly 
vra by tbc development of plates extending ijutnaida fiom 
lleial and backwards from the postfiontal; this roof being 

BOiMlly the Dunber \A iiuT|[inaU nuy be IncrcaKO It) twelve 01 rcUuccd 


most complete among existing forms in Cfulone and DenmA 
where the parietal joins the squamosal In such cases, in adl 
to the inferior temporal arcade formed by the quadrate, qtari 
jugal, and jugal, as in fig. 1007, there is also a superior anade 
stituted by the squamosal and postfrootaL Whereas, howero^ 
arcades in the Crocodilia (fig. 10S9) are separated by the infii 
poral fossa, in the Chelonia they are in immediate contact. 

The feet may either have all the digits free, or enclosed in a 
mon integument to form paddles. 

The humerus of existing Chelonians is a very peculiar bone 
actcrised by its extremely prominent globular head ; btu in a 
Jurassic forms {e.g., Acichelyida) this head was much smaller, m 
the bone departs less from a normal type. On either side of thii 
there is a projecting ridge, of which the radial, or preaxJal, one (em 
termed the lateral process) corresponds with the deltoid crest oTtbe 
codilian humerus ; while the ulnar or postaxial (mesial) procctf 1 
sents the inner tuberosity of the same. In all Pleurodiran Tettod 
the radial process is comparatively small, and the ulnar process |il 
in the same transverse line as the distal surface of the bone. Ini 
Cryptodira, however, and more especially in the land Tortmtti 
radial process forms a thin plate extending towards the ventral « 
and the ulnar process becomes twisted round to the same aspect; 
causing the pit between the two processes to form a narrow hinnd-ib 
channel instead of being very broad and open. At the same tin 
shaft becomes extremely curved. In the marine Turtles, where the 
is nearly straight, the radial process tends to become aborted, ai 
attain a position more or less below the head. In the AthccaU, 1 
likewise have a nearly straight humerus, the radial process, whD 
scending on the shaft, tends to an excessive development 

In time this order dates from the Upper Trias ; and it 
attained great development in the Upper Jurassic, from whicl 
it appears to have gone on increasing till the later Tertiary. 

Considerable diversity of views obtains as to the classifii 
of the Chelonia, but according to the system now followed i 
British Museum it may be divided into the two suborders Atl 
and Testudinata. 

Suborder 1. Athecata. — This group contains those fonns 
have been usually regarded as showing the nearest approximat 
other Reptiles, and therefore representing the most generalisei 
of the order. Dr Baur, however, takes the opposite view, a 
gards them as the most specialised group, which has tended to 
or less completely lose the carapace. Before, however, a de 
opinion can be given on this question it must be determined wl 
the absence of a bony connection in this group between the pa 
and pterygoids is to be regarded as an acquired or as an 01 
feature. It may be observed that Dr Baur regards the gro 
closely allied to the Chelonida, but if the undermentioned T: 


be rightly relerred lo it, we have at once a great 

lo the acceptance or his view. 

'■nboffder may be briefly elufactcriscd by tKc clrcomstince 

caiapace ts entirely of dc-rmal origin, end quite separate 

vettebrse and ribs, and may consist merely of a series of 

bones, or of marginals tni)i a •■in^lc median liors:)! row of 

ftcutes, or of a number of snull incgular scutes, with longi- 

row5 of larger ones; while ihc pb&tron (fig. ion) has no 

(interdavicular) element. The cranium 1$ charaderiKed 

absence of renicaJ platen ccinnecting the paHetals with ilie 

AH the forms are of marine habits, and consequently 

of the limbs are moditiei] into paddles Hkc those of 

noticing tbe two established families, it may be obscr^-cd 

imperfectly known Pstphodtrma of the Upper Trias of 

•nd Engbnd ie founded on a specimen which appears to 

■pace of a member of this suborder, although it has been 

I that it is not Chelonian at all. This presumed carapace is 

of a number of polygonal scutes, traversed by longitudinal 

of kedcd scutes. Here also may l>c mentioned tlie ^enus 

TlWnaikui, founded upon the anterior portion of a toutlied 

from ihe Upper Jurassic of North Amcnco, which Pro- 

^'Ur^h regards vi showing affinity with the Chclonia, and 

nuy possibly indicate a getwraltscd family of the present 

IILV Protostegiu^ — This family is mually regarded as the 
|:9|iccialised of the two that arc yet established, althouj;h an 
view is taken by Dr Baur. The carapace, according to 
ition of that authoHiy, is represented merely by a row 
nal scutes ; but the plastron is strongly developed, and 
of very thick ossifications. The t)-pe genus ProiosUga 
Ijfptcally in ihe Cretaceous of North America. It was con- 
by its dcscriber Professor Cnpe lo have possessed a solid 
, but ibc liones which he regarded as probably dorsal appear 
lo tbe plastron. It was also suggested that the dorsal 
were proccelous, with traces of transverse processes ; but 
: TCrtetns are probably referable to the cervical region. The 
(species inaioed very large dimensitins. An allied form from 
Upper Cretaceous of Italy lias been described under the name 
Ti ^vtnfiargis (fig. loti), but further e^i)!ence is required to 
pravc its right to generic disltnciion from the American form. It 
W^ indeed, been asserted that there were nu marginal bones, 
wt according to Ur Baur this is incorrect In the Cambridge 
ircetoand and ibc English Chalk there occur humeri of Athecate 
;bcloniati& which have been provisionally referred to Protoitega. 



It is uncertain whether the remains from the AmericaD Oct 
described as Atlantoclulys belong to the last-named g^ius. 

Family Dermochelyid^ — This family is characterised by! 
a carapace composed either of a median row of large and 
scutes and lateral marginal ro^m, or of a mosaic of small id 
scutes, or tessarse, traversed by longitudinal rows of laijei 
The plastron varies considerably ; and the humerus, which ii 
flattened type of that of the Che/onida, is distinguished fia 
of the preceding family by the great development of its radii 

Fig. ID] T.— Ventral a-<ipect of the pUiiron and ihoracic re^on fif Pi^fiii^mji'i mm 
from the Upper Cnuccous of luly. One-fifuenih utuiml liie. {Afur CB|icBii 

cess, which is situated near the middle of the shaft. The sk 
the temporal fossae completely roofed, as in the CkehmiJt 
an open tympanic ring, but has no bony floor beneath the 
passage. Whereas all the members of the preceding family 
moderate size, several representatives of the present one a 
huge dimensions. The earliest representative is the genus Eosp 
of the London Clay, of which the one known species was ori 
described as Cfulone gigas. The skull is of the general type 
of the next genus, but the carapace apparently consists on 
median row of very broad and large carinated scutes, and 
a row of marginals ; the structure of the plastron is not de 



ii was doubUess devx>id of lessane. The allied Psepluh 

iging in Europe from the Middle Eocene to the Upper 

tod also found tn the Upper Eocene of the United States, 

Tucd by the presence of a complete tessclatcd carapace 

Dci. In ihc carapac* the longitudinal rows of larger scutes 

rinaicd, ami arc more appmximalcd than in the existing 

e carapace is aho thicker than in the btter ; and there 

matpnab. It is also su^gt^tcd that the carapace may 

homy epidennal .shields. The Kkiill is short and much 

, THc existing genua Dtrmochelys (SfiAargis) is represented 

M) weU-known I^athery-lurtlc, and is characicrtsed by the 

t a tesselatcd pla&tron ; by ihc caiinalion of the scutes of 

rows of the carapace ; by the comparative thinness of the 

which is devoid of epidermal shields ; and by the longer 

: vaulted skull. It is, moreoTer, worthy of note that in 

IB the preceding genus, there U a distinct nuchal bone at 

lor extremity of the carapace, corresponding to the nuchal 

e&tutlinata; but there are no margin;i1 ossifications. The 

ing species of DrrmorAeJyi attains a length of ne:irly five 

a species oi /'se/AtffAcntj is estimated to have been as 

ten feet in length. The skutl of Et^i^harps, altbou^ih 

ger than that al Pstphopharus, docs not apparently indicate 

iiger carapace. 

iMUt a. TESTVDrSATA. — This suborder, for which the name 

ora ' b also employed, includes by far the great majority 

irdcr, or all those forms commonly known ait Tortoises, 

a, and Turtles. The group is characterised by the tniddle 

4 the carapace being formed of bony plates, developed 

' from the ribs and the neural spines of the dorsal vertebra^ 

it is (irmly welded. The outer surface of the carapiice is 

acrally smooth and overlain by horny epidermal shields, but 

me sculptured and devoid of such shields. The parietal 

■ tl>c »kull in all ca.sc3 send dovm vertical descending pblc^ 

By either unite direcdy with the pterygoids, or be separated 

D by the intervention of the columella or epiplerygoid. It 

Bbeerved that in nearly all the Mesozoic fomiK the vertebral 

ire very wide, and that this condition obtains in the young 

ater forms. This suborder may be divided into four sec* 

nc. and not improbably tw<^ families of the third section 

marine habits. 

ON 1. Amphichrlvdia. — This section is formed for the 
n of certain extinct Chelonians, mostly of Mesozuic age, 
Knbine in a remarkable manner the characters of the two fol- 

nane » objcctituuMr. ai bcine cmployd foi an order of lljdroid 
I \vUt mfira, vot. i p. 203). 

1 092 


lowing sections, and may probably be regarded as the soni 
the earlier ancestral types from which those two sections took 
They are all characterised by the presence of a mesoplasti^ 
and of an intergular shield in the plastron ; and the pelra 
may not be connected with the xiphiplastrals. The entopU 
rhomboidal. The skull and cervical vertebrae are unknown. 
Family Pleurosternid^. — All the members of this secti 
be, at least provisionally, included in this family. In addi 
the characters given above, it may be observed that the 1 
fully ossified, and that the carapace has a complete se 
neural bones, of which the hindmost articulates with the 1 

supr^ygal bone. On ti 
position that the Cbt 
plastron is derived fi 
system of abdominal n1 
those of the Rhynchoo 
and Sauropterygia, it t 
evident that the meso] 
of the present group 
archaic feature. 

In the typical genus . 
s/gmum (Megastemam 1 
gerrAum), which ocean 
monly in the English Pi 
and is also found in tb> 
land Oolite, the shell is 
and depressed, with co 
mesoplastrals (fig. 101 
large and wide entopla: 
single intergular shield, t 
nuchal shield. In the 
the pubis articulates ' 
smooth facet on the 
plastral, thus foreshadow 
complete sutural union 
occurs between these b< 
the Pleurodira ; but in the young it appears that there was n 
articulation. The neural bones of the carapace are hexagoi 
comparatively long ; while the vertebral shields {as in so n 
the earlier Chclonians) were relatively wide. Further, infram 
shields (shown in fig. 1012) were developed between the shi 
the plastron and the marginal shields of the carapace ; wh 
extremities of the xiphiplastrals were notched. The bones 
pectoral girdle and the humerus approximate to those of the t 
Pleurodiran genus Chelys. 

Fie- loii. — The plastron of Pliurattiruum 
Baltocki ; from the Purbecli of I>orMi»hirK. One- 
ihird natural siie. ig, Inter^lar <cute ; g, Gular 
do. ; ptc. Pectoral do. : at. Abdominal do. ; /cnt. 
Femora] do^ ; an. Anal do- ; e-py EpipUstral 
bone; m'/, Entoplasiral do.; ky.p, Hyoplailnl 
do.; Ml-/, McsDt>laKiral do.; Ay./. Hypt^lmral 
do- \ xp, Xipbiplastral do. 



confusion bae arisen in regard to this genus owing to a 

having been described under the name of Plattmyi Bullocks^ 

I' the erroneous impression that it had been obtained from the 

Gay- Remains of Plmrosternum are extraordinarily abun- 

ibe Purb«cfc of [>or»et»hire; and inelude speriinens of All 

>ni the newly Hatched >'uung, with a carapace of a ccupte af 

in length, to adult specimens which arc close upon twenty 

Curiously enough, however, no specimen of the skull 

Ito have been obtained. The young appears to have differed 

ly in the details of the shell from the adult ; thus, not 

s the puhis, as already mentioned, entirely unconnected 

pUMTon, but the marj^nal bones cncrwirhcd in a rcniark- 

I manner upon the front of the niirha]. Moreover, it seems 

very )-oung individuals the vertebral shields were divided in 

^JSUmk/r/, of the Lower Greensand of Bavaria, we have an- 

geniis also furnished with complete mesoplastrals, but ap* 

iy without any articulation K-twccn the pubis and the 

The shell is ornamented with a pustular sculpture rc- 

[hat found in the genus Tretcs/ernum, mentioned below 

the Cheiydrida. The plastron differs from that of Plturo- 

in that the xiphiplo&trals were not notched ; and there 

to have been a nuchal i>hield. 

next two genera, constituting the family Bainlda of Professor 

may be at least provisionally placed here. Both are devoid 

Ibony attachment between the pelvis and plastron. The genus 

cArAi (Heltmys\ typically from the Lower Kiitieridgian Htlio- 

ptuc limestone of the Continent, is readily distinguished by the 

of irregular ridges and prominences on the carapace (fig. 

j)^ and bj* the width and irrcguUr contour of the neural bones. 

BtetopbLStRlIs are small, and widely separated in the middle 

; and the inlergular shield is single. 

genus Bacna, from ihc Enccnc of the United Slates, has the 
xU meeting only by a point in the middle ; and is further 
}lc for the presence of double inlergular shields, and the 
ofa una]] additional costal shield in advance of the norma] 
costaL The caudal vertebne are opisthoca±lou& Professor 
who places this genus in the Cryptodira, regards it as indi- 
generalised type, showing marked sij^ns of affinity with the 
, and exhibiting uaces of an imperfect connection between 
pdvis and the pla&tron. The extinct North American genus 
hifytkffrax cbould perhaps be also placed in thi-s family, although it 
tk not certain that it is not alhed to Uie Ciyptodiran Adoms. 

Here sHo may be mentioned the very imperfectly known genus, 
JlnkMchtfyt. of the Engh&h Wcaldcn, in which, as observed abovOi , 



the paired sbietds of the phstron were sepdinted hy a medi 
of azygous sliictds, which probably extended backwarib fro 
Icrgular to the cxtrcmit)' of the xiphiplaslral. If the separei 
arc rixhily interpreted., it would appear that the plastron 
mesoplastral clement, which, instead of being situated in tin 
of the bony bridge as in Pieurosiernuta (fig. loizX lutd D 
part in the formntion of the axillary notch, fonned the aoia 
of ibis bridge and the whole of the notch. The hyopbstr 
therefore be entirely above the hiter notch, as in the exisli 


Fig. wty—Cv^UMOt PlatjKMyi Oitra4a^tri : from Iht l.0w*t KinuUgiui 
On*-tbird naiural wiv- (Afifr W^^tt.) 

nxliran genus Sternoihtfrus. I'ossibly also a Chelonian, 
Lower |uraii»ic of Stnncslicid, dciicnbed, upon the evidence 
impressions of the epidermal shields of the carapace, u 
name of Tcsttido UtrUkhndi, should aUo find a place in this 
The name ProfotJuJyi may be proposed for this form, wh 
certainly nothing to do with Ttstkdo. 

Skction a. Plecrodira. — The members of this section 
a rule, characterised by the complete ossification and union 
carapace and jilastron. and by the full development of the in 
bones, which are connected with the ribs, as well as by (he 
union or anchylosis (synostosis) of the pelvis with both the 
and the xiphiplastral part of the plastron. Very frequently 



bone, ind some (fig. 1014) or occa»ion%lly all of [he 
aic absent ; and when epidennaJ shields are developed 
liar (as in (ig. loi^] is present on the pUstroii. This 
bowc\cr, somctirats found Jn the next section, in which 
1» may also be reduced in number. The cntoijlastral 
to the preccdit^ section) is either oval or rhoraboidal ; 

ilastral bone (compare fig. 101 a) may be present. 
&kult of existing and Tertiary types the t)-mpanic cavity is 
iy mrrounded by the (luadxate, which forms an unbroken 
e articular »UTface of ihc quad- 
a concavity for the reception of 
ie 00 the mandible ; the pteryft- 
c inery broad and wing-like ; the 
may be absent, so that the pala- 
may meet in the middle line ; arnl 
may be distinct nasals, and a suture 
Tundibular symphyt^iv The ccr- 
renehrw hare well-developed trans- 
proccMcs, and single terminal ar- 
while there are never more 
thive phalangeals to the digits. 

Picurodira are fimher character- 
by their irvibility to rcinct the head 
within the carapace ; hut the 
ia bent on one tide and die head 
IviM^ht within the margin of the 
In all cases the lab)Tinth of the 
is completely open from behind. 
Ai the present day this section is al- 
eiclusivcly confined to llic southern 
jibcrc ; but in the Boccne forms 
or le&s neariy allied to exisiinj^ 
wtsc widely spread over the north- 
beraisphere, and in tlie Nfesuzoic 
were many European repiescnla- 
6m cf the groupt ^V■ith ihe exception 

4P«dam*in\ the existing fornu are carnivorous, and the whole of 
^^Km are thoroughly aqtudc. 

V\si- ici4.--Tb« tlchl haif irf ih« 

UbcB iftdicaa* iba bcwadMrw of iha 

At least one of the two earliest known genera of Tesiurtinat-i has been 
afarcd to this section ; and from the occurrence in maoy exi*tiny forms 
if wpuair nasals, and of a mesopLastral bone, we may regard this sec- 
bn a* retaining et'idcncc r4 a close alliance with a primitive i^cncntliscd 
tfpt ahich hat been tost atrwintc existing Cr>-ptadiT3. tince it is nciirly 
"ETtain ttiat features like ittcsc if once lost would not reappear, except 
H abonniialittcs. The presence of transverse processes to the cervical 



t'crtebra: would also seem to be a generalised character ; and, tn 
evidence atforded by the above-mentioned Pleuroi/ftintm and A\ 
^helys, the presence of an intergular shield should probably be B 
regarded in the same light. The anchylosis of the pelvis to the pk 
as pointed out by M. Dollo, would, however, seem to prevent m 
regarding the Pleurodira as the ancestors of the Cryptodira ; 
more probable that both sections should be regaraed as 
branches of a common stock, probably represented by Amphich^ 
earlier age than those yet known. 

It would seem proliable that the palate of the earlier Mesoioic 
dirans approximated to the Cryptoairan type, and that the peculiv 
dibular articulation and closed tympanic ring of the existing fan 
acquired features. The skull of the Mcsozoic types appears gened 
have been roofed over after the fashion of the modem Turtles ; tluii 
ture having apparently been common to many of the earlier fbims (I 
this and the following sections of the suborder. 

Family PROGANOCHELYiDiE. — This name is proposed 
Baur for the reception of the genus Proganoc^fys, fVom the Kfl 
of Wurtembcrg, which he regards as a Fleurodiran, althou^ 
ing from all other members of the section in that the piastron 
only with the edges of the marginals, without giving off axilliij 
inguinal buttresses. It is suggested that mesoplasttal bona 
present, while the carapace is compared to that of PlaIjA 
7'he latter resemblance suggests that this form might belong to 
Amphichelydia, but a study of the figure of the type specimen 
hy Professor Quenstedl, under the synonym of Psammoehtfys, 
that its extremely imperfect nature scarcely permits any 
opinion to be formed as to its true affinities. Whether the 
Tectly known and apparently aberrant genus CheJyfherium of 
Keuper is an allied form cannot yet be determined. ! 

Family Pi.esiochelyid,*:. — This Mesozoic family is characteAl 
l>y the total absence of the mesoplasttal element in the plastron, ■ 
also by the circumstance that the pubis alone is united with 4 
itiphi])lastral ; while there may or may not be a complete serin 
neural twncs articulating posteriorly with the suprapygaL The pi 
tral shields are separated from the marginals by the intervoitioa 
1 series of inframarginals ; and the plastron, as in the succeed 
families, is connected by long buttresses with the carapace. 1 
liumcrus is of the type found in existing Pleurodira ; and the sect 
3igit of the manus, when known, has but two phalangeals. 1 
ikull is known only in a few cases with certainty. 

The typical genus PUsiocfulys and the closely allied Craspedock 
vere originally described from the Ix)wer Kimeridgian Lithograj 
imestones of IJavaria and France. These genera ha\-e a Uiick d 
*ith the full complement of neural bones, which posteriorly join 
Irst suprapygal, and with the vertel)ral shields of the carapac 
uoderate width. The entoplostral bone is relatively wide 



; the nuchal is tnit slightly emarginate : and the intergular 

doable. The sur£i<:« of the carapace is uhuaIIv marked bf 

Sorae at kast of the Chclonians Frum the KimviidkHan of 

which havt been dtscribed under the pTCoccui>ii'd natnc 

1 tnar be indudcd in PUsiixhtiys, and this gcnuii is abo 

r.!i_-'! in the English Wcaidvn. 

Chclonians, fiom the Kimcrid^an of Hanover, described 
ibc tiatne of Cluiomides (likewise preoccupied), not iniprol>at}ly 
fofins connecting Plaiochetn with the undemicnlioncd 
•iyi. In one specimen refened lo the type species of Chtl- 
tfacnc are but se^-en neural bones ; but tliv vertebral shields 
et than in typical species of Hylafxhtlyi. If the>e forms 
y eotitlcd to rank as a distinct genus they rcqwirt: a new 
A skull uid to have been associated with the shell of the 
: species of Ckthmdes has the temporal fosste roofed over by 
(^ while the palatines arc described as meeting in the middle 
>lAer the raanncr of many existing Fleurodira. From the occur- 

&of Pttdochelyi and the undermentioned IJyiaochtlyi in the 
en, and the apparent rarity of the Adchelyidit in the same 
it it probable that ihc nwrnbcrs of this family were of frcsh- 
B probable that the genus PartuMys of the Lithographic Lime- 
(wbidi was considered by Professor von Zittel to be indis- 
^ fsaoi Atkhelys) y^ really identical yf\)S\ Piesiochelyx : in 
case the former name should sujieisede the latter, and the 
name Paraehtlyida replace Plcsmhelyidte. The humerus of 
gCDus resembles that of recent fleurodira, and is. widely diffcTL-nt 
that type of humerus which appears to be rcfcrahle to the 
Hda. The number of the phalangeals in the digits of the 
of thi» form (and al&o in IJiotAefys) is 2, 2, i, 3, 5 ; and thia 
u a decided approximation to the existing Pleurodiran genus 
iM, vrhich dilfen from all other freshwater forms now living 
there are only two phabtr^eals in each digit. 
name ilylaithtiys has been proposed for an allied genus of 
typically represented by the so-called Plcurvstrrnum 
\btm of xht English Pcrbcck, which as shown by a shell 
the Wealdt-n presi-rved in the British Museum, belongs to the 
t faiudy. This genus is distinguished from Piesioehelys by the 
wider vcrtchfal shields of the carapace, in which the width 
taf eseecd three times the tength ; and also by the ruirruwor and 
laDond-tthaped entopiastral \yanc ; as well as by the circumstance 
|at the neural bones ^1 l<^^i usually, do not join the suprapygals 
Id arc generally interrupted in the middle of the wrics ; while the 
ipopl aMfa! bone is relatively shoner. Both in this and the pre- 
there may be a vacuity in the middle of the plastron. 


It would appear that the Purbeck form described as Piairw 
emarginafum, as well as the Wealden specimens to which the 
Chelone Belli {eostata), and Platemys Dixom and ManM 
been applied, are likewise referable to Hylaeehefys. It is, mo 
very probable that a Chelonian, from the Upper Greensandd 
to which the name Flastremys has been given without deso^ 
also referable to this genus. 

The genus Idiochelys (Chehnemyi) and the allied Hyir^ 
the Lower Kimeridgian of the Continent, would appear to be 
allied to the preceding. Idiochelys resembles Jlylteochelys in 
tremely wide vertebral shields, and also in its small and dii 
shaped entoplastral bone, but differs in that the number of 
bones is generally much reduced, so that many of the costal 
in the middle line ; while there are also differences in the c 
of the lower plastral shields. This genus, which is known 
immature specimens, was indeed regarded by Professor Riit 
as essentially Pleurodiran, although there was no absolutely d 
evidence in support of this view. The shell is thinner than ia 
ehelys ; and the skull has the temporal fossae roofed over b; 
and apparently had a long sutural union between the postfrwd 

Mesozoic Chelonians or Uncertain Position. — It will t 
venient to notice in this place several Mesotoic Chelonians 
known to us only by the skull, of which the serial position 
at present be determined. A large skull from the Portland 
originally described as Chehm plaiyceps, but subsequently mi 
type of the provisional genus Stegochefys, is characterised 
incomplete roofing of the temporal fossae, and the meet 
the prefrontals in the middle line, while it is stated to have < 
nasals. The palate is unknown. The size of this specimc 
gests that it may perhaps be referable to the Cryptodiran 
Tkalassemys of the Kimeridgian. 

In the Wealden of Belgium there occur remains of young ( 
ians to which the name Chitraeephalus has been applied. T 
no mesoplastrat bone; and the skull is characterised by itselo 
form, open temporal fossre, and the approximation of the 01 
the nares. 

In the Upper Cretaceous genus Rhinochelys the shell ; 
known by fragments. The skull (fig. 1014 his) has an i 
temporal arcade, the temporal fossas are completely roofet 
the palatines meet in the middle line, the nasals are distim 
the prefrontals, which are separated from one another ■ 
frontals, the pterygoids are comparatively narrow, and k 
emarginate, while the symphysial suture of the mandible i 
terated. The humerus is of the Pleurodiran type found in 



Tt has been suggested that this genus is Plcurodiran ; and 
prove to be the case it will be interesting as shovrtng an ap- 
jn in several cnnial features to the Cr>ptodira. In tts 
rc3of the »kull of this genu.«: 3p|>roximnie$ to the Cktionida. 
\ fit Shinoihtfyt are ver^- abundant 
^Osifarii^ Greencnnd, and they 
I met with in lb« Gault and the 
Chalk. Fiagmentaiy Chelonian 
front the Cambridge Grcensand, 
the name Tnuhy^rmodutys 
applied, are not improbably 
to this gcntu. They are char- 
by Aeir pustulate external sur- 
t}ke pustules beirtg niueh larger 
the Amphichelydtan genus //f/<*- 
of the Loirer Greensand. A 
similar, although l(r&« marked, 
OCCU13 in the existing Ckelo- ^^ „,^ ^ _^, ,„,^ ^ 

ILT PEt^KEDt-siD^_- Reverting J'^X:S™V™?^:^a 
ia»sideration oT undoulrted Pleu- r^y^fr -l^I'T'^ii.^^fc 
the present cxbiing family agrees /— '. h^-o-""*- 
Ampbichelydian PSmrestemida 

presence of inesapla:slrat bones, but is distinguished in 
the pubis anil ischium have a sutural union with the 
TTic shields of the plastron, as in the next family, 
contact vith the margirmls, owing to tlie absence of infra- 
Thc skull.1 of existing types ha%-c an infratvmponil 
and in Podocnemii, alone among existing representatives 
KCtion, the temporal fo**ae are roofed o\-ci ; moreover, the 
Is are in contact in the middle line and are fused with 
the palatines, owing to the absence or abortion of 
vomer, meet ; and the suture between the dentary bones of 
mandible is obliterated. The second cervical vertebra is 
:lous ; and the neck is completely retractile within the 
Further, the series of neural bones, as in the next family, i« 
lete and b rtot connected with the suprapygals. Finally, !n 
'thb and the ncitt family the characters of the t)-mpnnic and 
regions of the skull are those mentioned at the commence- 
EBt of the description of the Pleurodira. The earliest known 
ircseniatives of this family occur in the Up[Kr Creiaccous of the 
ated Sutes, where we meet with forms apparently cl«sely allied 
the existing Podocnemis. The genus Bothrcmys has been described 
the evidence of a skull from those deposits ; while the name 
^krmfkys has been applied to portions of the carapace and 


plastron which not improbably belong to the same fonn. Bt6 
differs from Podocnemis by the presence of a distinct vomer, 
not improbable that allied forms occur in the Cretaceous d 
Zealand. The only existing genus which appears to be represa 
a fossil state is the above-mentioned Podocnemis, of South At 
and Madagascar, which occurs in the London Clay and the E 
of Northern India. This genus includes the largest existing 
sentative of the section. The carapace has seven neural I 
and the mesoplastrals are small, and do not meet in the i 
line. The shell of a large Chelontan from the London Clay, 
has been referred to this genus under the name of P. Beb 
may not improbably belong to the genus Daeockefys. The b 
founded on a mandibular symphysis from the same dqmdl| 
is characterised by a large spine-like process on iti onnj 
and the serrated margins of the alveolar borders. Its sertd p 
depends on whether it is specifically identical with die i 
mentioned shell. 

Family Chelyid*.* — In this family the shell is much ti 
than in Plesiochelys, and is characterised by the absence i 
mesoplastrals, and the reduced number of neural and supr 
bones ; while both the piibis and ischium unite with the pit 
The skull has an incomplete inferior temporal arcade, owing 
absence of the quadratojugal ; and is further distinguished 1 
separation of the prefrontals by the frontals ; the distinct 
(except in Chtlys); by the vomer dividing the palatines; »■ 
the persistence of the suture in the mandibular symphysis. 
temporal fossae are open, but there is an arch connecting the p 
with the squamosa], not found in the Pelopmdusida, and wb 
probably a remnant of the earlier type of roof The fifth and 
cervical vertebrae are biconvex. The neck cannot be fully ret 
within the carapace. As typical genera we may notice the 
American Ckelys, Hydraspis, and Plaiemys ; and the Aus 
Chelodina, Emydura, and Elseya, in which neural bones are a 
Remains of Chelodina and Emydura occur in the Pleistoa 
Australia; while Hydraspis is found in a fossil state in the 
Eocene of India, and Platemys has been recorded from the 
ceous of North America. 

Family Miolaniid^e. — This is an extinct family represente 
by the remarkable Miolania, from the Pleistocene of Austn 
which the remains were originally referred to the large Vara 
the same deposits. This family, according to Mr Boulenger, oc 
a position in the present section somewhat analogous to that 
by the Chelydrida among the Cryptodira ; the caudal ve 

* Frequently incorrectly given as CMyJida. 



and the temporal fossae of the cranium roofed 
The cmuum (fig. loij) is rcmvkablc for canying 
pain of hom-likc proccues (on wliich account tlic name 
tijr* ha» been proposed); and the tail was protected by a 
bony sheath (fig. 1014), somewhat resembling that of the 
>nt Edcntaiei, The species of which the skull is shown in 
:ul muM have attained huge dimension;!. 'I'hc carapace 
>n are tuiown only by fragments. From the structure of 

y ♦* 

\ an),— JTM^aaft 0»ml. A. ADiertor Tbar nf cnninm ; >, Ri[hi kuanl iiptrt af Ih* 
Mf aTlW —Jul Jialli; (ram llbc Plattoain et Aunnlia. .Much nduod. <iUlv 

wn it h inferred that Mio/am'a was herhivoroux ; while the 

of the terminal phnLingealx and the solid caudal sheaih are 

live of terre^tirial habits. Tlic hones of the pelvis are fuaed 

Ian innotninste bone, and some of the cervical vertebrae had two 


Labould be observed that Dr Baur disseni* from the view ihai ihiH 

I b a PleurtMlinin. and would refer ii to the Cryjumliran family Tes- 

ySr; bui there appear 10 be several mi sconce pi iojis in ihe argu- 

laddaced in uijiport of this view. Sir K. Owen do«s not, indeed, 

_n>M the Chclonian nature uf/l/(i>/it;v//t, and regards ic as CDiistitut- 

' a distinct gruup. under the njiine of Ccralosauria ; but thi? view i* 

fetber at variance with the facts, and there can be no questiuii but 

', ibe genus is a iiue Chelonian. 

FaHiLv C\RErnK:iiLLYiD.t. — ^The last family of the Plcurodira 
tlftaiacterised by the absence of horny shields, and is typically 
it«l hy Carettixhelyi of New (iuinca, in which the neural 
are very small and do not touch one another, and the limlw 
IK paddlc-shapcd. In the Ixiwer Koccnc of Northern India we 
find the extitKt genus JhmUfulys, which difiers from Carettothtiys 
in baling »even large neural bones in contact with one another ; the 
BUnne of the limbs being unknown. 

SlcnoK 3. CitrrTODiitA. — In this section the ossification of the 
cmpBKX and plastron nuy be either imperfect or complete, and the 
iwo nay or may not be connected at their edges ; there iit no bony 
Bttachment of the pelvis to the plastron ; and as a general rule, the 

1 103 


full complement of suprapygal and neural bones is [ffesent,! 
there are numerous exceptions. Excluding the Ampbid 
forms, no known Cryptodiran has a mesoplastral bon^ i 
the majority of cases, especially among existing types, tl 

troD has DO intaf^ 
(fig. 1016). Theaiui 
when present, is dtha 
rhomboidal (fig. loi 
f-shaped (fig. loog),! 
epiplastral joins the 1 
tral. In the skull (fig 
the outer border of d 
panic cavity is always 
notched posteriorly ; tti 
rate articulates with 
cavity in the mandibl 
the pterygoids are o 
rively narrow and 1 
emarginate. There ai 
more than three pha 
in the digits. The sat 
caudal ribs (as in the 
dira) ardculate partly 1 
centra and partly w 
arches of the vertebra 
the cervical vertebn 
very imperfectly d« 
transverse processes, 
isting Cryptodiran has distinct nasals, and at present tha 
decisive evidence of the presence of these bones in any fossi 
The Cryptodira are further characterised by their power of 
ing the head by a sigmoid curvature of the neck in a vertic 
directly within the carapace. This large section comprises t 
majority of the existing Testudinata, and at the present 
mainly characteristic of the Northern Hemisphere, being 
absent from Australia. The habits of existing Cryptodii 
be carnivorous or herbivorous, and either terrestrial, fluvi 

Family Acichelvida:. — Under this name may be grc 
number of generalised Cryptodirans often spoken of as the 
semytfes, and in some cases as the Eutystemida. They o« 
cally in the Lower Kimeridgian Lithographic Limestones of tl 
tinent, and are abundant in the Kimeridge Clay, while they s 
till the period of the Lower Greensand. They were p 
mainly of marine habits, since they appear to be veiy rare 

Fig. 1016.— PUilton of Kathiga Iteftam; from 
India. Onc'half natuiat size ^, Gular shield ; 
Aitm. HutneraJ do. ; fiec, Pcctoial do. ; oj, Abdo* 
minal do. \/im, Femoral do. : attt Aikal do. ; #./, 
EpipLutral bon« ; fivf./, EntoplastraL do. ; Aj/^fi, 
H^ioplaiirB] da ; *-/y, Hypopksuat do. ; x./, 
Xiphiplnslnil do. 


Purljeck and W'L'alden beds. They are chflracterised by 
or \c3i hcattsbapcd carapace, of which the costals arc 
perfectly ossified, and do not unite fompletely with the 
'Ilic plastron .has, moreover, a vacuity in the centre, 
for a long period or throughout life, but it had large 
and inguinal buttresses for connection with the rarapuce. 
has a rer}' imperfectly developed head and a slightly 
fthafl ; aod the limbs were not modi^ed into paddlc;s. Tlie 
bone of the carapace has no costiform processes ; and the 
■1 foBSK of the skull vcrc more or less completely roofed 
f bone. 

S>eir cofdiform, and frequently imperfectly os:>ified carapace, 
^kktiyida agree with the modem CheUnidie ; and since the 
r form of the neurals of Trvpiticmyj is fuiother feature only 
tb eUewbcre in the latter family, there are strong; grounds for 
Rg the one family as the direct ancestor of the other. This 
aUo exhibits certain signs of alSnity witli the Pleurotliian 
%efyida ; white the simplicity of the hutnerus is a very general- 

genus TkaJastemyt, which includes some very large forms, is 

Icfised liy its long and flat neural hones, in which the ntiterior 

surfaces arc mu<^ shorter than the poaterior ones ; while the 

al shields arc narrow. The carapace is well ossified, and of 

nbk thickness. The type species occur* hoih in the Litho- 

tone and the Kimeridgc Clay of England ; the unde- 

onian from the latter deposit at Ely, to which the name 

has been applieti, being apparently identical. As al- 

ed, the Hortlandian Utt^ehelys may IJlctrwise be llie 

aiMl the genus b repiescnied in the I^orsetshire Ptirbcclc. In 

genus Aeiditfyt {Euryslemum^ Achetonia, Palaomeduxa, 

) the neural bones of the carapace arc flat, with short itnd 

listinct aniero-Litem! suifaces, and the cosWis are well ossi- 

Kttebral ahiclds of the carapace being comparatively wide. 

seems to lie confined to tlie Lithographic Limestone. 

'ktfyt, of the Kimeridgc Clay, we have a lai^c form 

very imperfectly o&sified carapace, in which the costils are 

!y thin, and the neural bones arc long and six-sided. The 

surfaces of the neurals are much shorter than the 

tcnJ, and in the 6fU) neural the latter surface.<i arc deeply 

The Dcurals especially in the hinder pan of the rara- 

strongly ridged, having the form of the ridge-tiles of a 

the hinder pan of the whole carapace is itself roof-like. 

bral epidermal shields were very wide, and the borders of 

' Thii Dame is (be euliett, but it pieoccupjed. 


the areas which they overlie on the carapace are conqna 
fluted. The last genus, Tn^demys, while ' having the roo 
posterior neurals of Pehbatockelys, is distinguished by tJie i 
regularly hexagonal form of these bones, owing to the lengA 
of the anterior and the shortening of the posterior lateral tm 
One species from Hanover referred to this genus shows the n 
feature of a median row of numerous small intervertebral i 
dividing the normal vertebrals into two lateral series. This 
occurs in the Lithographic Limestone, in the English Kim 
Clay, and the Lower Greensand of Switzerland {T. va/ai^im 
The detached neurals are readily distinguished not only by 
shape, but by their exc^sive thickness. 

Family Chelvdrid^ — This family is confined at the p 
day to North, and part of South America, where it is repm 
by the Alligator-terrapins ( Chtlydra), and the larger Snapper (J 
elemmys). The skull is more or less triangular, and very 
behind ; the temporal fossa is partially roofed over, but the 
mosal is widely separated from the parietal ; the bones of the 
do not develop plates to floor the narial passage ; and the tyn 
ring of existing forms, like that of the Testudinida, is in gm 
closed behind. The nuchal bone of the carapace develops i 
rib-like (costiform) process on either side, which underlies the 
ginals; and the complete union of the marginals with the i 
does not take place for a long period ; while in the pi 
vacuities frequently persist till late in life, and the plastron 
may be relatively small, and unites with the carapace by gomj 
the hyo- and hypoplastrals not giving off buttresses to undeil 
carapace. In existing forms the caudal vertebne are opisthoa 
and there is no bony union between the ischium and pubis 
same side to enclose an obturator foramen. The head can 
completely retracted within the shell. The humerus is not flat 
and ha.'i its radial and ulnar processes large, and directed ti 
the ventral aspect. 

If all the genera provisionally included in this family are 
referred, it will be the oldest representative of the order whii 
exists. The genera may be divided into three subfamilies, t 
ing to the presence or absence of sculpture or epidermal shie 
the shell. It is, however, by no means certain that all these 
should be included in this family ; but until we know the ski 
caudal vertebra; of all the genera it is impossible to decit 
point, and it may eventually prove that there is a more or les 
plete passage to the Dermaitmydida from this family, sinc< 
already are some indications of a connection between them. 
the American Cretaceous genus Toxoehelys is stated by Dr I 
be a true Chelydroid, but to have the procoelous caudal vt 



and also to have an open t>-mpanic ring. The 

axe typically rcptt-svnti'd by Aiostira, of the Ui>per 

of KattH America and England, and wc may prnlhibly 

in the same (^oup Puudotrionyx {A/ihalidemvi), of the 

and Lower Eocene of Euiopc. In iheae forms ihc :thell 

Tcnnirulated sculpture, but epidermzit shieldK appear to be 

although Ur Baur auiea thore are traces of them in the 

gena»; the plastron is well de\'eIopul ; nnd ihe neural bones 

laced to seven, as in Dtrmatcmys. Amttira, which, on ac- 

of the presence of only ten marginal bones, I>r B;iur places 

Dtrm-itfmydidte {^aurotypida), difTcrs from Pseudatrionyx 

kcutpturinj; of the nt-ural bones. The skull of Pseudotrionyx, 

referred by Sir R, Owen to Plafemys. agrees csfcntialty 

of Afaerocltmrnys, although thcorbits arc less lateral. The 

sah&nnily, Trt/osttrnina, is represented by I'retosieruum, 

English M'caldcn and Furhccic, with which Peftechetys, of the 

of Be^um, is identical. I'ossihty the imperfectly known 

Atnencan Cretaceous genus Compitmys may prove to be a 

allied, if nM identical, type. The shell is studded with 

tubercles, and lius epidennal shields; while the pListron ts 

than in the C/ulydrimr, and has an interguLir hliield. This 

I bowever, accompanied by only five paired plaslral shields, as in 

ys, instead of the six found in PUuresternum (fig. loia). 

1 "TT***' bone is deeply cmarginatc. 

Fnallf, in the typical subfamily Cktlydrimr the shell is not 
and has epidermal shields ; the plastron is relatively 
and generally ha& a median vacuity; while there is the full 
of eight neural bone^ of which the last articulates with a 
VTP^ and thus prevents any of the costaU from meeting in 
^■uddle lirve. The type genus Chelydra i.s represented in the 
or Lowest Eocene of the United States, and aNo uccurt in 
iVpper Miocene of Switzerland, ihe MiddK- Miocene of Sl>Tia 
it has been described as Chcfydivpsis), and the Ixiwcr 
of Ro(t, near Bonn. No foBsil fonns liave hiiht-rto Iwen 
to Mturoclemmys, which u distinguished from Chtlydra by 
mare deeply emaiginate nuchal, and the lateral orbits of the 
■U, which is very Urge in proportion to the shell and linib-hones. 
Fajiilv CiNo-fTEHNtn*. — With the American genus CiMOsStrmim 
•e ODcne to a family readily dinlingiii>hcd l)y (he toUit ali.sence of 
Ae entoplastT3l element of the plastron. In the exi^^ting genus the 
aknll has open temporal fosse, the nuchni lias a costiform process 
like that of the Chtiydridtt, the pelvis is of a Testudinaie type, and 
the caudal vcrtdwa; are proccclous. The alisencc of the entoplastral 
occurs in the Swiss Iwcene genus DilkyroxltrnHm, which lia,s 
iogly been placed in the same family. It is distingui:>hed from 


Cinosternum by the presence of eight neural bones, and die ! 
bridge connecting the carapace with the plastron. 

Family Dermatemydidve. — The genus Dermaitmys, of 
America, is taken as the type of a family which iocludes 
typus of the same region, and in some respects cmnectii 
CinosfernidtB with the Chefydrida. The nuchal bone has a ' 
form process, as in both those families ; the temporal fouc ft\ 
skull are not roofed over ; the caudal vertebre are procceloui; i 
in the pelvis the pubis and ischium of the same side do not i 
to enclose an obturator foramen. The shell is well ossified, 
the type genus the union between the costals and marginab i 
not take place till a late period. Baptemys, tiS the Upper 
of North America, which has been included by Professor < 
DermalemySy is distinguished from the latter by having the 
number of neural and suprapygal bones, so that none of Ae ' 
meet in the middle line. Here may be mentioned sevenl 
North American forms, most of which are referred by 
Cope to a distinct family — the Adocida — mainly on account of I 
circumstance that the ribs do not develop heads to articulate i 
the vertebrEe ; but since the same feature occurs in 
among the Testudimda, it cannot be regarded as of &mily ' 
In Homorophm, of the American Cretaceous, there is no 
gular shield, and the vertebral shields are very narrow. In Ai 
of the Upper Cretaceous, and Agomphus (with which Dr Baor i 
Amphiemys is identical), of the Eocene, intei^ular shields wOl 
present. The former has traces of sculpture on the shell ; while ii 
the latter the epidermal shields are very thin, and the sur^ure oflhl 
shell is marked with a faint vermiculate sculpture. We are thui IbI 
on to the European Trachyaspis, which is probably an allied ta^ 
and with which one or other of the American types may peila|i 
prove to be identical. This genus, which has elongated veitdial 
shields recalling those of Dermatemys and Baptemys, occurs typidfl) 
in the European Tertiary, and has also been recorded from ^ 
I^wer (ireensand of Switzerland, although the latter fonn has bcfl 
referred by Professor Riitimeyer to Plesmhelys. A nearly entir 
carapace has also been obtained from the Tertiary of Egypt Ti 
surface of the shell is covered with a distinct vermiculated sculptin 
like that of Trionyx, but thin epidermal shields were present. 

Family Platysternid*. — The existing Burmese genus Plei, 
slermim is represented by a very small Chelonian forming the on' 
member of a family which connects the Chelydrida with the Tat 
dinidis. Thus the temporal fossae of the skull are roofed over, tl 
pelvis is of a Chelydroid type, and the caudal vertebra are most 
opisthoccelous ; but the nuchal agrees uith that of the Testudinu 
in having no costiform process. 



T TlSTODiNiD*. — Following Mr Boulenger's *TTaiigement, 
isive bmily is taken to include the Cisttidimda and most 
'mydidm o( other writers, and may be briefly characterised 
n ; The Utnbs terminate in free digits ; epidermal horny 
Ire aluays present, Ihii there is no intecguUr shield : the 
I is ovoid and fully ossified ; the plastion in the adult is 
ed with the carapace either by suture or a straight articula- 

I b always fully ossified. There is no costiform process to 
bal bone; the temporal fossae of the skull arc not roofed 

bone ; the caudal vertcbnc arc procccloua ; and the pubis 
lium of either side unite to enclose an obturator foramen, 
merus has a wcll-de%'cloped head, and its &haft is more or 
red. The skull is of nearly equal width throughout its posl- 
ortion. 'I'bc Terrapins and Tortoises, as the existing mem- 

this family arc commonly termed, exhibit great variety of 
oroe being aquatic and others terrestrijU ; while some, again, 
riTorous and others herbivorous. Some of the terrestrial 
we the normal two epidermal caudal shields of the carapace 
geiher into a single large shield. 
My commence our brief survey of this large family with the 

group of Batogurs, represented by Jiata^r, Kac/mx'i, liar- 
id allied types. These include freshwater TenapinB, frc- 
oT large size, characterised by the great development of the 

and inguinal buttresses of the plastron, which dix-ide the 

■ the carapace into chambers, and also by the presence of 

more Ktrong ridges on the oral surface of the palaie, running 

to its alvetJar borders. On the plastron (fig. 1016) ihe 
tetween the humeral and pectoral i^hields is below the etuo- 
Kaehuga (including Pan^hura) has the fourth vertebral 
loogated, and o\'erlying parts of four or five neural bones; 
nor Dcurals being elongated and hexagonal, with the short 
erioT, The typical A', lineata, m which the fourth rcrtebral 
\ broad anteriorly, is represented in the Pliocene Siwaliks of 
vhicb aiso yield the existing A'. MoM^oia. In a second 
Paffgihura), in which the fourth vertebral shield is narrowed 

E' 1 at its junction with the third, we have the existing A". 
. tet6) in the Pleistocene and Pliocene of India. JJar' 
1017) is clutracterised by the shoniie>s of ihc fourth 

II shield, which usually eirtends over three neural bones 
I the (igurt), and by the third vertebral shield uveiiymg 
t three (in place of two) neumls. It is represented in the 

by the existing //. TAttrxi {fig. 1017). With the exist- 
ican genus Ckrysemys we come to forms in which the 
and inguinal buttresses arc much lcs.s developed than in 
and the neural bones are .shorter than in HardtUa. 



In Chrysemys itself the sulcus between the humeral and 
shields of the plastron is situated entirely on the hyopIastnJtl 
fig. 1016; and from the presence of this feature, it is 
that the so-called Emys testudim/ormis and £. M^rtMota, 

London Clay, should 
ferred to this genus. '. 
existing genus OcaA, 
China, and also in the I 
arctic and North 
C/emmys, the hut 
sulcus is placed 
more anteriorly, and 
quently cuts the ent 
bone. A similar iatHn] 
frequently found in die 1 
called £Mys erasta {A 
lietuis), of the Upper : 
of Hampshire; and 
pears that this form 
be referred to the 
Palaochefys, of the 
Miocene, from which 
is probably not 
Emys wyomingensis. 
Upper Exx:ene of NoA 
America, appears to be ■* 
ferable to the same gtoK 
Ocadia, it may be obsenc^ 
agrees with the herbivotOH 
Batagurs in the presence of ridges on the palate ; but these riclpi 
are absent in Cltmmys and the undermentioned forms, whicli M 
purely carnivorous. 

These forms may be divided into two groups, according is to 
whether the plastron is united to the carapace by suture, or sin^ 
by ligament. In the former series Clemmys is represented in thl 
Pliocene of Algeria by a species closely allied to the existing C 
Uprosa of that region. The characteristic Oriental genera Damaot 
and Bellia are represented in the Pliocene Siwaliks of India ; ih 
fossil Damonia being apparently inseparable from the living h 
Hamilloni. In the second series, where there is a more or la 
complete transverse hinge in the plastron at the junction of th 
hyo- and hypoplastrals, and the buttresses of the carapace may l 
wanting, we find remains of the existing Emys orbicularis {£. I 
Iraria or Lutremys), commonly known as the European Pond-to 
toise, in the Pleistocene of England and the Continent. Cistiui 

Fig. 1017.— Cdrapaceof //arrfr//a Thnrgi; India. 
XH, Nuchal boiic ; ni-nS, Ncunl do. ; ify. Supra- 
py^al du. ; fiyt Pygal do. ; ci-<8, CoiibI do- ; 
lf^l-Hlll^ Marginal do. Reduced. 

I ainioat exclusircly terrcstiialf has been recorded from the 
ntol Miocene. The Indian genus Nmria, which is repre- 
rt the cxistii^ N. (Chaibauia) tricarinata in the Siwaliks, is 
Uied from all ihe preceding forms in thai the neural bones 
Hr >bon side placed postcrioily. The exiinut Plyeho/^astcr, o( 
VI Miocene (Upper Oligncene) of France, is rhamcterised hy 
Mcnttitu junction of the hypoploalriil with the carapace, and 
retcnoe of a hinge between the hypo- and hjroplxstrals, and 
the coniDur of the neural and cosul bones, which approxi- 
I ibotc of the true Tortoises; from which we may assume 
I form was mainly of terrestrial habits. In both genera the 
•pectoral sulcus cuts the cntoplastral bone, 
e bmd Tortoises forming the laat group of the family, the 
loncs arc generally very short and wide, and may he either 
Mil, or altcmateljr ictragoruil and octagonal ; while the cotttal 
Ire ^nerally ahernately short and long at chcir inner and 
Urcmitics ; the suture Iwtween the marginal and cosia! Ijones 
coincides with the sulcus dividing the corresponding shields, 
caudal shield is generally undivided. In most cases the 
kctwccn the humeral iind pectoral shields is behind the enlo- 
bone. The digits dilTer from those of most of the preceding 
by Ireing adapted solely for walking, and devoid of wcIm ; 
te metaoirpats are shorter, and the humerus h verj- much 
with ibt radial and ulnar processes approximated. The least 
existing forms arc Cynixi's and lyxts, which have neiir.Tl 
those ol Ilomepui, but costaU of the ordinary Emydinc 
appear to be unkno»Tt as fassils. I/adrianus, of the 
') Eocene of the United States, includes generalised 
of large size, in which the neural bunes are elongated and 
with the shorter lateral surface* posterior ; the costals do 
,c in length ; the vertebnd shields are narrow ; and the 
it divided. The genus I/omnpus, which now includes 
of small size confinwl to Africa, has no ridge on the 
and the neurnl bones are short and hexagonal, with short 
lateral surfaces, ami the caudal shield is single. It is rcprc- 
in the Upper Miocene of Switzerland by the so-called £niyt 
, and in the t.ondon CUy by E. Camptom. In ^yltmys, from 
lie River Miocene of the United Slates, the short neural hones 
cir shoncr lateral surface placed anteriorly, the posterior cnsuils 
, alternate in length at their ejttrcmities, and there is a single 
ahield. All the remaining forms may be included in Testu4o, 
h the pabte has or>e or two ridges ; the neural bonei; iiiuaily 
of an allemaiion of small tetragonal and larger uct:igonal 
rhile the anterior exlrtniity uf the ejiiplastrals is more or 
Ckcnod. la the akull the pterygoids arc wide, and dcpteiwed 


in the middle (fig. loiyi/j). Usually the caudal shield is 
but it may be divided. The ribs have no heads for aiticulatiaa| 
the vertebre. The forms described as Manuria and 
may be included in this genus, which comprises a laige re 
species. Exclusive of the Jurassic so-called T. Strickian£, 
has been already mentioned under the head of TVv/arA^s, ' 
earliest occurrence of the genus appears to be in the Upper '. 
Phosphorites of France ; it is, however, not certain that the 
from these deposits may not be referable to one of the 
genera. A marginal in the British Museum indicates an ind 
about 30 inches in length. T. gigas, of the Miocene of 
Loire, is a still larger form ; and species of la^e dimoisions 
occur in the Lower Miocene of Allier and the Middle Mioceoei 
Gers, as well as in the Pliocene of France and Greece 

The huge T. {Colossochelys) atlas, of the Pliocene Siwaliks of 
was one of the earliest of the lar^ extinct forms brought to the 1 
of science, although its size has been greatly exaggerated. It ap, 
indeed, that the length of the carapace was about six feet, or one-tl 
^eater than in T. elephantina, of the Galapagos Islands. This sp 
IS remarkable for the great development of the eptplastral bones, 1 
formed a pair of hom-like processes ; and is also noteworthy for 
anchylosis of the three bones of the pelvis into an innominate bone. iM 
skull, which probably belongs to this species, resembles in structure <m^ 
skulls of the recent giant tortoises of Aldabra. The carapace had a» 
nuchal shield, the caudal shield was divided, and the limbs nR 
covered with bony ossicles, as in the existim; T. emys of India, to wbiiA 
this form was probably allied. The large T. perpiniana, of the Plioceai 
of France, in which the depressed carapace measures nearly four feet ii 
length, and the limbs were likewise covered with dermal ossicles, m 
probably also nearly allied. An unnamed species, from the Pliocene rf 
the Siwalik Hills, has a skull resembling that of the Galapagos tortooa 
(fi^. 1017 f>is) ; which are characterised by the backward extension of At 
opisthotics. We also find in the Siwaliks T. Caulleyi chaiacteriscd by 
its small epiplastrals ; and T. punjaiiensis, which appears to hai-e bed 
a form allied to the smaller existing Indian T, emyi. Remains of giiM 
tortoises also occur in the Pleistocene cave-deposits of Malta. 

Finally, we must not omit to briefly mention the giant tortoises of ibe 
present epoch, which, it appears, have been driven from the contiuaitil 
areas by the competition of the higher types of life to seek refuge in 
islands, where they attained an extraordinary numerical development, tiD 
their haunts were invaded by man. These tortoises formerly existed '* 
great numbers in the islands of the Aldabra group, lying to the north-wtd 
of Mada^'ascar; in the Mascarenes, which comprise Mauritius and Rodri- 
Ifuei ; and also in the Galapagos, or " Tortoise-Islands," which lie off the 
coast of South America. The Aldabra Tortoises are characterised bf 
their deeply excavated palate, short opisthotics, and the presence of a 
nuchal and of double gular shields. Some of them were living in the 
year 1877. In the Pleistocene of Madagascar remains of two very laigt 
species have been found, both of which present the characters of the 
Aldabra forms. The Mascarenc species have no nuchal shield, and onh 
a single gular ; and the whole of them have been extenninated by humai 



Tbrir skalb rrwinblc ihmc of the next group (fig, tof?^it) in 

bt exca>4tion i)f the p:iiaie, and ilie prrutuccd opi^ihotics. Finally, 

ipayra tonoiies. i. cniuidciable nuinlier nf vhich were found in 

K and Abingdon liUnds during the voyxgt of Ihr Challenger, 

ttnpiiftbcd bv- ibr double and the ubsence of the nuchnl 

FnxD tht Pleistocene of South America, Dr Mar«node$CTibe« 

XB tortoises allied to the CalapHgos forms : one of ihem being 







tit. — KitMBl lupicl of ibc cnniun of Tnltnif pwnn/*!"" ■' '""" *'■• Calnj^oi 

- »» i h lfil* IMIUT'I tUc. HH. AuilitucT ipatiur* ; it. Buiutiiiiii >1 ; tt, Ruinih* "Old i 

l]4c «/ q uaJMtt : Ki. SJuilU : <•-, OuHpiul condyle ; «f, Ogiiiihiiiic ; /«/, Palaiin* I 

LUlUi #1*. Futxtoid . /■• (juKilnii : t^n. N|uiinoul : iv/. Siipratxci[iUiil tpine '. 

u ihe probable anceswr of T. nigra {tiffhaniofius) of ihe 
^roDp. The carapace of 7'. tlepkanliKa measures four feet in a 

ILT CheijdsidvE. — The lost family of the Cryptodira differs 
H the precedir^ in hanng the limbs modified into more or 
Dnplete poxldles or fliiipers (fig. loiS), whkh tn the existing 
ha%-e only one or two daw<i. Thu t-arapacc is broad and 
or less depressed, and is very generally heart-shaped (fig. 
although in some of the early forms it is rounded at both 
Mlics. 'llic nuchal bone has no cosliform process ; there arc 
larger or smaller unas*ified spaces between the coslals and 
lala of the carapace (hg. 1006), but in very old individuals 
,y oliliteraie in some forms; and the pta&tron is not articu- 
it. P 



Intcd to the carapace, and has digitated lateral terminoiio 
generally a larger or smaller median vacuity (fig. 1009).] 
humerus is moic or less flattenoi, with the axis of the 
erally placed nearly ini mediately above that of tlie shaft, 
radial process small, and placed more or less below the 
developed head. The caudal vertcbnc are prococlous, aoAj 
ccrvicals extremely short. In the !>kull the temporal foaM<j 
completely roofed over by bone, so that the liquamosal joiiB\ 
parietal ; and the bones of the palate unite for a longer or : 
distance beneath the narial pasuge, .so as to throw the 
,.nare& more or leas backward (% loso). The prefrontalt, 
Imosi Ttftudinidie) alwa^'s form a re-entering angle posicrio 
the tjinpanic cavity Is quite open posteriorly, so as to 
stapes. In old individuals the vacuities in the sliell tend to 1 

ni. loil.— Yaaii|<rflb( Kawtubill Twtia (rMnw ladvintajL Uaib f«luNA. 
CAft« B*)l.] 

literate, and in the more generalised extinct types this teixlenqr i 
much more marked, and ic is quite probable that in some 
they may have completely disappeared. I'his more complete 1 
cation of the shell in these generalised types indicates aAinity 
the preceding families, and probably more especially with 
Adfhefyidit ; and the same is indicated by other feature*;, sudii 
the more marked constriction of the shaft of the humerus, andtll 
more oblique position of its head, together with the ptolulrie 
encc of claws to all the digits, Some wTilcrs, indeed, consider th* 
more generalised types as entitled to constitute a distinct family 
\i\c Propltunda ; but their close relationship to the existing 




forhida this view. Wc hnvc already suggested reasons for 

thi& family as twing descended from the Mesozoic j4a- 

and froai this point of view it is interesting to note the 

> of an intvtgular shield in the plastron of the exi.'^inj^, and 

ft therefore of the fossil foini.s »ince, as we luve already 

fHaa appears to be an archaic feature. Ilie same observa- 

■pply to the open tynipanic ring. At the present day 

' four living species of Turtles — viz., the Loggerhead 

ijoiMefyt atrtrta, fig. 1006), the Mexican LogR<:thcad (7*. 

t), the Hawksbill [Chtione imbritata, fig, 1018), and the 

I Turtle (C. mydas), all of which arc of purely macine habits. 

■wksbill alone is carnivorous ; and is further peculiar for the 

thai in the young state the epidermal shieldn imhricate, 

of uniting by their edges. Ii has been suggested ihat the 

genen wer« of estturine rather than purely marine habits ; 

it IxMTie out by their occurrence in the estuarine deposits 

[ihe I<ondon Clay, to the exclusion of the murine genus 

remarkable of the extinct genera is Lyloloma {Eur!as/ts, 
Glassocheljs, or ErquelinMsia), which octuts typically 
a ibe EocoK of North America and the London Clay, and is also 
KpTscnird in the Middle Eocene of Brack Icsham, as well as in the 
and Cambridge (ireensand. In the t)'pical Eocene forms, 
Kxalled Citiatte fflanimenium and C crassiwstatum of the 
day, die sicull it as large in pToporlion to the shell, as in 
vnj among the C^efyJruitr. Th« skull of tlie adult is 
ieaukablc fot the extremely backward position of the |>osierior 
wbicfa art approximated to the basioccipitol ; and for the 
lattd width of the mandibular sj-mphysis (fig. 1019, a). The 
has k>w alveolar walls, and no oral ridge ; the nares and 
are directed somewhat upwardly ; and ihc bar between the 
a narrow, '["he shell is characterised by the great extent of 
jhoMiificaiion, as well as hy the rounded posterior cxircmily of the 
an|»ce. and the suiural connection of Che broad xiphip1a.<itrals in 
)At median line. The head of the humerus is » oViliquc, 
nd iix ihaft much constricted. The limbs were clawed. Curiously 
noogh, in the young, and perhaps also in the adult of some of the 
imOo' forms, the po^erior n.-tres were plac«d much le^ back- 
mdly, and the mandibular 5>-mphj-sis was shorter and less 
tutened. It is probable that at least some of the Nonh American 
Eocene forms described as Otieopygfs, PrepUura, and CtUapieura, 
ate not separable from this genus ; while a turtle from ihe Creta- 
cnes of .^uMraiia, origmally described by the preoccupied name 
if XittfiMrfyt, but which has been provisionally designated A'tfAv 
probably ohio nearly related. 

1 1 14 


Closely allied to the preceding is the genus TJia/atsxka 
represented by the loggerhead (fig. 1006). The skull is, hoi 
relatively smaller in proportion to the shell, which is disting^ 
by the presence of five costal shields ; the carapace is faeait-^ 
the xiphiplastrals are narrow, and join only at their extren 
while the terminal phalangeals are flattened and have but o 
two claws. The shell is distinguished from that of Ckelom \ 
more complete ossification, and its T-shaped entoplastral; 

Fig. 1019.— Oral oipKt of ihe left half of ihe nuuiilibic of I^Mrma crmuimfmtmm 
L'htlaiu Hiyiias (u). Reduced, llie former from the Lower £oc«ic of Belsium, ibe 1 
cent, r, PosLiriiL-ular proccu; p, Splerual ; jv, Geniohyoid xrcove; ^, k, loMftion of i 
muscle; k, (Juranuid process. 

the humerus is more constricted, and has its head placed 
obliquely. The position of the posterior nares and the form < 
palatal walls and mandibular symphysis closely resemble the 
parts in the young of Lytohma. In very old specimens the ■ 
ties between the costals and marginals completely close up. 
genus probably occurs in the London Clay,^ and also in the M 
Eocene of Brack lesham. 

> As indicated by a. mandibular syrophysii iu tlie British Museum. 



'^iofhtfys, of the l^ndon Clay, the skull (fig. loio], ^^tuch 

: and bu deeply emorginate pterygoids, diflTcn from thai 

genera by the presence of an oral ridgi! on either 

^tbc poJatc and in the middle of the tnandihubr s>-inphysi», 

these arc present in the Mexican Ixjggerhcad. The 

in the young of Lytoloma, is very long, and extends 

tlo join the ptemaxillx ; and the pterygoids are charac- 

tbc forwaitl position of their ectopltrryguiii processes. 

•eerTK to ha%'e been much like thai of Thalassoefufys, 

only four coiiia] shields, and its ptutron was perhaps 




' fnix 

.— FraattI lad palMalMpecu of ■ jraaiiE cnmimtif Mrtillt^tfyt iiitrkf/i: (nm 

Oaf. Ovi*g M (■■Hiifttj iha iwlMal TtHgu v* Imprrfwily il»ilr>ra><!. ii, 0<- 

J, raMccifml, c, Pukul. mm) 4. rronnl [(oJitmal ihirldi; tr. Ila.>loaipiial;_f > 

■id: 1^. filial iflfMJ poteMOl^o- ; f, Vomei; wa. Manila ; /nr, PnrnuuHa. Th* 

iadkcfroMBlufca tan>E«aud «titi tbesiof As. loaa, in ohidikiMnftnitrMi. 

_ ossified, although the wphiplastrals united extensively in the 
■iddle line. The mandibular symphysis is convex, and com- 
irdy shorL 'I'he skull of the type species was originaily 
as C^^*K cundctpi ; while the shells figured as C. suh' 
and C. amttxa are referable to this genus. 
In the typical geniu Ckehnt the »ku1l is much smaller in propor- 
to the shell than in TAaJassix/uJys. It is characlerised by the 
■ore or less nearly vertical position of the nares and 0[l>itN and the 
width of the bar between the latter ; by the tall alveolar walls and 
llfac oral nd|{c» of the palate and mandibular symphysis ; the long 
poygpids, with the cctopter)'goid processes placed soine^^hat back- 
nnlly; the forward position of the posterior narcs ; and the more 
rle» marked shortneKs of the convex mandibular symphysis (fig. 
:oi9. b). The skull, moreover, has no occipital epidermal shield. 
Is gcDcnlly present in the other Torms (fig. loio). In the 

shell the carapace is either heart-shaped, or pointed ai 
(fig. 1021); the vacuities are la^ and persistent ; the aM 

is tong and dagger -1 
and the xiphiplutn 
blender and KpanB 
1009). The humeral 
little constricted, with i 
nearly on the axis of 
and the coracoid is 
more slender Ihaa in 
swluiys ; the Green 
tttydai) being more f ptf 
tlvese respects ihi 
Hawksbill (Sg. 1018). 
easiest occurrence C 
genus appears to be 
Cambridge GreensanC 
Cault, where C. /atm 
very massive mandibltr 
what Tcsembling that 
H.itvk.sbill. 'Iliis fom 
[joissibly be identicaJ % 
Beiiiltdi of the Oal 
toai), which is onljrde 
known by vcrj" young 
and has been made the type of the genus Cimoliockelys. 
topmast Cretaceous of Maastricht the gigantic C ftoffmdi 
pear« to be allied to 6*. imbricala, but has a shorter and 
palate and mandibular syniphpis, a more deeply emarginate 1 
and the co^tal bones extremely short. From the latter <i*ery 1 
iscd character Dr Baur regards this species as entitled to . 
distinction, and has proposed the name of AlhpUurvn. An 1 
large, and perhaps closely allied turtle occurs in the English 
In the higher Miocene of Bordeaux C ^rondka appears t 
funn closely allied to existing types. 

Sbctio!* 4. TmowvcHoiDFJi. — The last section of the « 
includes the mud-turtles or soft-tortoises, of the freshwater* c 
Africa, and North America ; all of which are of aquatic ai 
nivorous habits. These forms, which may probably be r< 
as extremely specialised types, present the following dist 
features. The shell is sculptured and devoid of epidermal 1 
its enioplastra! being in the form of a chevron, which divi 
epiplastral from the h)'oplastTal ; and the marginals, if | 
forming only an incomplete scries at the pasterior extrcmtt) 
carapace, and having no cotmection with the ribs. The lo 

I tin|ri>hCh>lli. IUi)uoc<l. (Aflcr Omii.) 



zhnc have no distinct iranini-crsc processes, and tlie eig)ith 
with the first doisal Milely by ihe xy^ppophyses, thus 
log die most remarkably complete hinge-joint among the whole 
ipVenebracn. The Toiirth digit in each limb is chanicicnscd by 
||f not less than four phaLingeals. The form of the tympanic 
|i|D(the qtudratf, mid the tebtions of the latter to the mandible, 
lllfaesunc type as in Ihe Cr)-ptodira ; and wc find a similar 
pee of anion bct»'een the pelvis and the plastron. In the 
pOB (%. loax) the chief distinctive features arc to be found in 

Eie ; ihu* the broad pterygoids are -ieparated from one another 
ixubpbenoid, which extends forxi'ards to join the jKilntinest ; 
B. airing to the small *iie of the vomer, uniting exten^ively 
Ik middle line ; and the whole structure of the palate being of a 
■Boditan type. Ajs minor character of the skull, attention may 
ttRctcd to the open temporal fossae, and tlie forward po&ition 
direction of the small orbits, as well as to the extreme 
it of the backward processes of the supraoccipiCal and 
The sacral and caudal ribs articulate only with the 
i of the vertebrae, and there arc no chevrons. The plastron is 
I etuirely »iCpnratc from die carapace, and has a large median 
' and digitate extremities, as in the marine Chiionida. Kur- 
[the head and ttcclc are retracted within the carapace after the 
manner, but, owing; to the peculiar structure of Ihe 
I cervical vertebra, in a still more complete way. hi all casett 
tetior lateral surfaces of the neural bones of the carapace are 
I iihoner than the anterior. 
I marked resemblance of the palate and the general aspect of 
[Trionychoid skull to that of existing Pleurodira ii> a circum- 
thii does not admit of a ready explanation from a phylo- 
poinl of view, unless we adopt the somewhat improbable 
an that the Pleurodira gave origin to the Trionychoidea at a 
: when their pterygoids bad attained their full width, but while 
Etympanic ring was still partially open. 

Fawly TwoHvaiiae. — The whole of the members of this sec- 
I may be included in a single family, which is however, suscep- 
of division into two subfamilies. In the first, or Emydina, 
(opiithotic of the skull unites with the pterygoid 10 divide the 
nor aperture of the auditory labyrinth into two foramina ; the 
-and hj-popta-stral of cither side arc fused togtjther; the sciilp- 
tse of the carapace is tubercubted ; and marginals may be present. 
Tbe existing Orienut genus EmyJa, in which marginals and a com- 
plete tenes of neurals are present, i.t represented by numerous species 
in the Pliocene Siwaliks of India, some of which attained very large 
dimensions. The African genera CyclanorMf and Cyclodtrma, in 
which nurgituU were not developed, are at present unknown in a 



fossil state. In llie second subfamily, or TrioHychina, 
hypoplastmis remain distinct throughout life ; the 
not join the opisthotic lichind the lab>Tinlh ; the sculpcu 
consists of sinuous ridges or pits ; and Di.irginal bones _ 
developed in the carapace. 'I'hc type genus Trionyx is wid 

tributed at the prei^ent di 
extends downwards into tbe 
Kocene of both Europe t 
United States, and is aba 
settled in the Upper Cietao 
the latter area. Manj'spM 
remarkable fur showing dta 
types of skull, which ia otM 
Acatton has a broad pabte ■ 
for crushing, while in the i 
palate is vcty narrow, 
species, in which the iki 
subject to ihi.s variAtic 
short neural hones 
cosials : while in the 
vAw forms there are but 
of coslals. All the Eurc 
forms agree with the map 
species in having only otu 
neural bone between tbe & 
tals. In Europe this genus a 
to be unknown above tfc 
Miocene of (Eningen, audi 



of Tnmic ramttlifMi ; ftoni Che P Icj.iio- 
can* ti! IndiL KiduMd. m/, Snpnocci' 
Diial; fur, rmrittat: fit^, PwtrnHilal ;^, 
Vriviiial ; /•■/, Fnlimttal + nnal : mr. 
MuilU i /■, Jueal: fi/i Qiuiltai cjunl ; 
.on ullier «ioe of 

or. Orliil, Till 

nr.Orli.i, me orocwiM on ulher «i<)e of i'Jtut-wiv «i <.^iiiii| 

:l:::^SSS;±Tr,ir;fy'^'IK:''c'pt^ ^remely abundant .n 

lh«eic, oT which ih* innri boMir •riicululei EoCCnC (LoWCr OllVOCenc) 
lln lafitMixipiial 1 tJieinlilly to ihe ,, ~. . . , .. 

' it th( proaii:. well. The cxistinjj Indian i 

occur in the Pleistocene aoi 
cene of that country. Of the American Teriiar)- species A 
some have eight costaU, and in some cases only six neural^ 
Lower Eocene and Upjwr Cretaceous forms described by Pn 
Cope as Plastcmenus may be included in the type genus, sir 
thickening of the plastron can scarcely be regarded as a | 
character. 'I'be same remark applies to the Eocene Axtslut, 
United Statc.<>, in which the plaiitron is smooth, as in some c 
forms. An apparendy di.stinct type, in which the outer mar 
the costals is deeply grooved, occurs, however, in the E 
Upper Bocene, which has been named Autaa>tMefyt. Final 
exclusively Oriental genus Chitra is represented in the PI 
and Pleistocene of India by roinains of the one existing spec 
member oi 



CL.4SS REF2'IUA — (onHHtud. 


MC Branch. — According to the orrangetnent pro- 
by I>r Baur, ihis branch is t.ik<;i) la include the four orders 
in the heading of this chapter. These order:^ pr<^cnt a 
instance* or the divergent vicwrs obtaining among difltTcnt 
as to the classiiication of R«ptil«. Thus Professor Cope 
to place the Ichthyopterj-gia in a group l>y themselves, 
be reg-irda as of equal value to another containing all the 
white Profeswr Scclcy at one time regarded this order 
ion of the Dinosauria. although its relationship to the 
been long before pointed out by Sir K. Owen. Pro- 
has also proposed to separate the Squamnu from all 
Reptiles under the name of Cccnosauria, and lo include the 
dtoeephalia in a second division as Palseo^tiria. Prorcssor 
hile not going so far as this, places the RhynchiKcplialia in 
nqMosaurian branch ; while Professor Huxley would include 
bat group and the Protcro^una in the Squamata ( I ^iccrtilia), 
hich the Ophidia are eicludcd, Ur Baur, again, merges the 
sauna in the Khynchoccphalia, and regards the latter as a 
I order allied to the Squamata. 'I'his middle course is fol- 
io the present work, altliough the Proleio&aurio, in accordance 
views of Professor Seeley, .ire provisional ij allowed lo ranlt 

present branch may be collectively characterised -a* followa : 
eniition is very gently either acrodont or pleurodonl, with 
eth of the adult aiKhylosed to the jaws ; but the leeih may 
planted in a gTOO^'c, and remain free. In addition to their 
in the jaws, (ccth may also he developed on the palatine 
;oid» and very rarely on the vomer. There is very fre- 


quently a parietal foramen ; the superior tempoial arcade is | 
^ly present, but in the more specialised foims the inferior ii 
wanting. The quadrate may be either movably or 
attached by its proximal extremity to the cranium ; a < 
very generally present ; and the ramus of the mandible 
a lateral vacuity. The precoracoid appears to be fused wiflit 
coracoid, and may be represented by a precoracoidal procea f 
1040), or its position indicated merely by a fontanelle, while ia I 
IchthyopCerygia even the latter is wanting. A T-shaped int 
and clavicles are present in all the earlier and a large proportioaj 
the later forms. Usually all, or nearly all, of the dorsal ribs i 
late by single heads ; the dorsal vertebrae either have short or 1 
mental transverse processes, which are never placed entirely 00 1 
arch, or (Ichthyopterygia) a pair of facets on the centrum;!^ 
with one exception, there are not more than two vertebne in 4 
sacrum. Abdominal ribs are present in the earlier, but are l(nt| 
most of the later forms. The humerus frequently has an ectepie^ 
dylar foramen. The number of phalangeals in the digits of peril 
dactylate land forms is generally 2, 3, 4, 5, 3, in the manns^ ■ 

2. 3. 4. 5. 4 in the pes. 

In no case are secondary posterior nares formed by the devda 
ment of plates from the bones of the palate to floor the nn( 
passage; the posterior nares consequently always forming moK I 
less slit-like horizontal apertures in the roof of the mouth. Ndthi 
is the posterior extremity of the palate ever completely closed h, 
the junction of the pterygoids with the basisphenoid, after tti 
fashion obtaining in the Chelonia (fig. 1017 6is) and some Sum 
pterygia ; there being always a vacuity between the hinder atim 
ittes of the pterygoids which displays the presphenoidal rostrum. 

Order IV. Ichthvoptervgia, — The Mesozoic Ichthyosauis od 
their allies were large marine Reptiles, with the body long, *i>d 
shaped somewhat like that of the Cetacea (fig. 1023), witbcil 

Kig. loaj. — Gieaily reduced rctoraiion of ih« ^keleion of /cilAjraanrmt ermmtmiil frg« il 
Lower Liu of DorseLshirc. lite pelvic limb is nlaiively too luge. (Afm- Ovco.) 

either dermal or epidermal skeleton ; the limbs being modified ini 
paddles, in which the component bones were in apposition on i 
sides, and the phalangeals were oval or polygonal, and increasi 
beyond the normal number. The skull (fig. 1034) has the Eac 
region produced into a long rostrum, mainly formed by the p 



in the upper jaw ; and the naies are consequently approxi- 
to ihe orbit, which is of veT>' large size, and has a ring of 
in [he sclerotic ; thert: in a targe parietal foramen ; and 
two tnoponU arcades connected lORcihcr by ihc supra- 
il < i>rosquainuul or supniquadrate), which roofs over the 
rai fossa, after the Labynnthodont fashion. Other char- 

^~lt*4atad bit lu«ml view ct tb* Uiud of //Afiwwawiu infoMnAiii; from Ihg 
I ml tXniabin. tmt, Pnnuuilto: J/^. MaxiUa-, .V, K*ra>i £«. Lachiroul; 
M>ll Fr, tt^itl: Pa, Partvul; Pi. Ptxlfmul : SI. Supnitnivrvl ; Xf. S<tw)- 
(er, Quit>a((i«K>l i C'-^wdnxi ?■. Jusal ; Pl.O, |-oiioibiMl. <A/t«r llwhr-} 

features of the skull arc ihc small fronlals ; the presence 
I oputhutic in the occipital region ; the di.'«tinctnes« of the post- 
frotn the |x»trrotilal, and of the lachrj-mal from the pre- 
; ihe firm ^sture of the quadrate, which dots not, howe^w, 
By unite with the plcrygoid ; and the presence of a foramen 
; the middle puniori of the {|uadratojugal from that bone, as 

Pm. ■•«.— Tb* otncrnm tl an tattnai .I'li— I kcnel.ra at liklijr*H-rvi, vipwol in t«ii«n. 
Ml«BilBaMwigriuHllcAUici>ln*»e(t>; from i)w Kiaicrldi* CUy orwilitbin. R*diHt>l. 

Iw casting Rhj-nchocephalian genus ^hmod^H, and the presence 
rr cotiuneOa, or epipterygoid. 'Hie palate has essentially the same 
Enctiue as in the last-named genu«. \i in the Squamata, there U 
D lateral vacuity in the mandibular rami. The teeth are confined to 
le jaws, and are implanted in a continuous groove, nithout nnchy- 
ta the bone. Their crowns arc sharply pointed, and ore usually 

1 122 


cylindrical and deeply fluted (fig. 1032), but are occasioniOj^ 
pressed, carinated, and smooth (fig, 1028). The Tertebial 
is primarily divisible only into a precaudal and a caudal 
there being no sacrum. The centra (fig. 1025) are ami 
more or less disk-like, and either deeply cupped or nearly flat 
either side those of the precaudal series (fig. 1025) carry a ; 
tubercles for the articulation of the double-headed ribs; while ■ 
caudal region these tubercles coalesce. Superiorly each centnmii 
pair of flattened surfaces for the attachment of the neural 
which are united merely by synchondrosis. In the cervical 
the tubercles for the ribs are placed near the summit of the 
surfaces of the centra, and they gradually descend on the ( 
till they reach the base of the lateral surface in the posterior | 
caudal and caudal regions. In some forms the upper costal tnb 
of the anterior vertebrae is placed either partly or entirely on I 
neural arch, and it is convenient to term such vertebne, or 
homologues, cervicals, and the remainder of the precaudal 
dorsals. The centrum of the atlas vertebra is well developed, 4 

Fig. io>6.— A, Ventral atpcct ot (he pcctonl Eirdle and riaht limb of /Mkfnmiinii M 
Btcdiw. n, Pelvic glrdte of do. Redund. s, InierclnvicTc ; i, Cbvicle ; c. Scapnk; 
Coracoid : t, Humcni*; /, Ulna; £, Hadiui 1 H, Iliuni i u, Iichium ; /, Pubii. (Jli 

there is a large intercentum between the atlas and the skull, ai 
another between the former and the axis. There was no stemur 
but a complex system of abdominal ribs was developed. T 
dorsal ribs were devoid of uncinate processes ; and ribs occur 
the caudal region. In the pectoral girdle (fig. 1026, a) there 
no precoracoid ; but clavicles and a T-shaped interclavicle, like i 
corresponding bones of the Lacertilia, were developed. The co 
coids were large and expanded, devoid of any fontanelle, and i 




: anothur at their junction. The tlircc bones of the 

Ifi.' 1026, b) arc weak and rod-tike; the ilia have no coii- 

' h ihe sacral r^ion of the I'enebral column ; and although 

and ischium of opposite sides meet m the middle line, 

ihe same side do not unite to enclose the obtmaior notch, 

cms and femur arc relatively Khort ; but the radius and 

:itill vliuTter, and niay be reduced to ohiong bones in 

the transvcrM: diamctct 15 the longer. 

has no foramen. The slruc- 

tbc paddles will be noticed under 

of the different genera ; but it may 

that when there is a difTcrcncc 

sue of the limbs it is the peciorai 

the larger, from Ihe less special- 

ure of the limlw of ihe earlier 

Di Baur regards the Ichthyopterygia 

iants of l:{nd animals ; a conclu- 

F«hich is suT>|>oned by the argument 

[had ihcic Rcpdles originated directly 

Fishes, as was formerly supposed to 

: cue, ^cy would have retained ihcir 

In regarding this order ns elofiely 

to Sf^atodoK, ])r Baur lays strcs.s 

the general similarity in the cranial 

ar>d especially the presence of 

foramen between the qiindrnte tmd 

ajagal ; the identical structure of 

idxluminal ribs; and the remarkable 

of the pectoral girdle, which, this 

remarks, is only comparable to that 

[tbecKi&tiitggcnuA, especially when young. 

Ilbe dcuU and vertebral column the Ich- 

))gia retain evidencva of their Laby- 

it descent, which are lost in the 

' orders of this branch. 

hi& order is known to range with cer- 

from th« L'Kwr Trias to the Upper 

It was also widely distributed tu space ; its remains having 
discovered in the Arcric regions, in Europe, India, Coram, 
jTonh America, the cast coast of Africa, .Australia, and New 
^buid. Part of a jaw referred to Ichthyosaurus has been de- 
Pnbed from Malta, which, it has t)ccn suggested, may be of 
UJoocfK! age, but this requires confinnation. 

It nay be teinarked that the humerus and femur of this order 
Ite quite unique in that, instead of having convex condyles for the 

dmw, and iluiil iipicD of 

tamu : fr^in iho KimtridKO 
day. One-ihinf mtinral *itt. 

oti r<rt ndiui ; ■. Do. for 

1 124 



articulation of the epipodial bones, they present distinct i 
(fig. 1026 *k) for their reception. 

Family Ichthyosauri d*. — Although Dr Baur makes the | 
Mixosaurus, Ichthyosaurus^ and Optkalmosaurus the types 
many families, yet it seems on the whole more com 
include all the known forms in a single family. The most 
ised group with which we are at present acquainted is the 
Mixosaurus, from the Upper Trias of Italy, which is founded I 
a small species presenting the following characters. The 1 
and femur articulate distally with two bones; and the ndimi 
ulna are elongated, and separated from one amither by an ii 
throughout their entire length. The teeth are mull, and xAK 
cessively numerous. It may eventually prove that all the 

forms are referable to Aii genn. The 
genus Ichthyosaurus is charactoiaed by the 1 
presence of teeth throughout ttw jam; andl 
the normal articulation ^ tht hmiemi widi 1 
the short radius and ulna, iriudi ace in cIcmc i 
position (fig. 1027). In die pebic limb 
femur similarly articulatei onlj with the 
and fibula at its distal extremis ; and thus 
extremity in both the humenu and famir bu < 
two articuUr facets (fig. ios6 i£r). The 
which articulates inferiorly ividi tibe radius 
ulna at their junction is the intennedium (fi^'i 
1027,/); while the one 6Uing tq> the angle bfr 
tween the latter and the nulius is the radiile; 
the opposite angle being occupied by the ulnm 
Below the intermedium we have the cenud^ 
which in the more specialised forms may be 
double (fig. 1031), and below this the renuis- 
ing carpals, metacarpals, and phalangeals, wfaiti 
are combined to form a pavement-like structiR. 
The bones forming the continuation of the line 
of the intermedium in fig. 1027 correspond Id 
the middle or third digit of the tj-pical manu; 
and the structure of this limb indicates that it 
was derived from a four-fingered ancestor, since the poUex, or fint 
digit, is not represented. 


Fi([. 1037. — DorMll 
aspcti of Ihe left pec- 
toral limb of IrhlkfB- 
saunts latifrottx; from 
The Ij>wer Lia'i- ke» 
dui:ed. A, Humerus; 
r» K.idiu«; u,. Ulna: r, 
In termed mm. (After 

Specimens have been obtained from the Lias of Wiirtemberg and 
Barrow-on-Soar, in Leiccsiershirc, which show the contour of the in- 
teK'uncnts of the padrflcs. It appears from these thai on the anterior 
border of the paddle (fig. \oT] bh) there was a comparali vely narrow 
band of integument, which was covered by minute homy scales ; while 
on the posterior border there was a much broader flap, which appears 


incd parallel bands of muscles set obliftucly to the axii of 
In the Latipinnalc group (fig. 1037 ft«) the posterior flap 
mcnt was narrow in proportion 10 llie bony fratncHork of ihc 
ad it was piodoced infrriorly into a loti); poini below the distal 
n the Lon^pinriAie group, however, the imegiiinciit* were much 
pniportkin to the bime^ and Chey tenninniccl inrcrioily in a 
tremit)', «luch only reached a short diftt;ince beloiv tlie distal 

inttr lomu attained a length of from thirty 10 forty feet : and it 

'it pnrfable that the extremiiy of the lAil was provided with ;i 

WW fin. AH the species wrre riirniiomus; iind, a» wc learn 

Ui—ttn td tlu kft pcCMas) limb uf tiAlkyruanu iifirmt^in, viimd fram 
f^sa, ml •txiwinc I'm contour sf Iht inr*f(irnfni. Orx'hall naninil *ln. Am, Kn- 
^ Trii — ! f^ Ubu ; P, RfediaU ; «, lni«tiTi«diuin , ■', Vliara ; «, e', C*nii^(>a' From 
Ltai it Bain* «n-Saai. 

: K»-<»tlr(l coproliies, their food cnrtsisted to a considerable ex- 
Aancnd Tiahe^. These coproliics funhcr tell us that the inicstiiic 
Tided with a spiial valve, as in the Selachian lishes, E'crliaps, 
,the most remarkable circumsiancc connected with their internal 
By is the not unfrcqucnC occurrence of eniitc skelciont of small 
kiab within the ihoraeic and alMdoininal Ciivily of Iiir^er ones: 
null specimens being invariably uninjured, and bclnnt;ii)g to the 
peon a& the one in which they arc contained. This lends Pro- 
Seeley to conclude that some stkccics or individuals were vivi- 
, and that ttic young were probably produced of different relative 
different species. There n also evidence to show iliai in some 
any yoon){ were produced at a birth ; the number being perhaps 
^.c character. It cannot, however, be taken as proved that all 
iir» wcic viviparous, since in such oihcr Sauropsida nntl Ich- 
in which the same mode of development occurs, it is not dls- 
lany entire group; and it is n otc wort ny thai the three specimens 

1 126 


with young ligurcd by Praressor Seelcy all belong to a ^ng 
This gems may be divided imo iwo groups from the struoi 
pectoral limb. In the !j>neipintii»tt, or Ica&t spccLili&cd rtoui 
onknott-n ^bovc iho Lias, the orbit is generally nf vcrj' large di 
The pectoral limb (fig. 1027) ii ctinniLienhed by having the I 
or thnt nnKinu from the in termed rum, coiiiprisinv nwj a M 
tu(lin»l row of bones, and consequently having only one cenin 
there are usunlly only four longiiudinal rows of phalnn)[cah in 
and the radius is nearly square, with a distinct notch in il 
border (%. los/ii This group may be further subdivided ii 
groups. One of the moftt aberrnni fiirms it ihc hu^ f.ffatyoi 

Lower Lias, uhich attaittci 
(ti nearly ff»rty feeL This 
characterised fey the pelvi( 
bcinj,' nearly as larjie aa ibtt 
but still tncirc readily by 
(fig. 102S), which have cai 
carinatcd, and smooth cm 
the roots covered Hiih a 4 
cement. /. loHcMtfliioH is 
Low^r Liassic form with ( 
and fluted teeth. Anothf 
indicated by /. I^nuirritr 
lifi/rs>ns {l^tngimtrrii), in ■ 
cninial rostrum (fig. 1029)1 
oloiiijatcd and very sknd 
paddles (fig. 1027) are n 
ibr Ihc large size of the Oi 
bones ; and in /. ttnttirpitiik 
a small circular vacuity b«i 
radiu* and ulna, indicatti^ 
remiuint of the large ^aa 
in Mixwanrut. Lastly, A 
„ . , I J ti . _ f /tV, J, tttlatnti£uj, and , 
iDaOidr JckihwiaumfiaiifiiM: rnini iiic afc tnrce Upper Uassic rai 
U-<r LIm or Iw-tAlf.. this group indicating a ira 

ihc next group ; the radi 
latter species having no anterior notch. The akull of /. MttA 
shown in %. lojo. 
In the more specialist^ nr LalipiamiU group, which imnsei 


Fi(. nog.— Left bUnJ lirr of ib« (kull of /<*ri^rn*>mu /«y(fW«u ; bvaihi 1 

Lia» 10 the Chalk, the orbil is usually relatively smaller than (| 

iiroup. The peanral limb (fig. 1031 J has the third digit, or tli 
rom the iniemiedJum, with a double longitudinal row of txines, 
arc consequently two ccniralia. There arc, moreover, never 


1 127 

tdin^t ro»-*or p)uilanK«nls in thi» limb (ihc itiarginAl roiv-sarc 
tig. 1031}; lUid the radius is Iranstcracly cloitK-iitrtl (.ind thus 
ly rcmo«'cd bvai the nonnal t}pc) with an entire •inCcrior 
t t» imponani to obscnc that m this group the spliitinu of 
di^it And the two ccntnlui ntc evidently acquired, and not 
characters. Like the Utit this xroup may be divided inio 
b In the i^-pical »ubsroup, which n ciinhned in ihc Lias, 
■'.fig. 1033} nave their roots sdongly Huivd, and ihe dorsal 
the hamcnis has no sirongly-marited trochanteric ndcc. /. 



'f is the species least widdjr removMl from the preceding' 

hiie the typical /. communis is distinguished by ihc cxirmio 

(he paddmk In a tccond tu>>grtiu|i, which ranges fmm the 

lay to the Chalk, the roots of the teeth arc invented with a 

*? of cement : the dorsal aspect of the humenis has a pro- 

TDcbanicric ridge : a.nd the coracoid dUTcrs from ihal of the 

^ tobgrtnip (fig. 1016; by the alisence of a posterior notch. This 

I cuoipnses / trfgottus, :ind probably other allied fnrms from 

td and Kimeridge Clay* : and also the wctl-known /. mmpylo- 

I the Upper Cretaceous of a large portion of Europe, to which 

■^fnnn me corresponding strata of India, and perhaps also /. 

from the Cretaceous of Australia, are probably closely allied. 

rr is in some respects even more specialised than (he follnw- 

A. The species referred to this genus fmnn the I'olar regions 

dervd 10 be of Triassic age : and a species from Ceram may 

Cretaceous beds. The genus is unknown in America. 

nost specialised representatives of the family are the genera 
mosanrus, from the Oxford and Kitncndge Clay and piol>>il>ly 
i Cretaceous of England, and Bapianodmt {Sauranadpn), 
K Upper Jurawc of North America. In the latter teeth, 
|p n dental groove, were totally wanting, but in the fonner 
were prc&ertt. In the fortner (and probably also the 

Fin- IL^JI' — Vfioi rAl itf pttt ol the rifhi ^KVivriiJ ll'Vtb (waiiliTi^ i - j . :i A 

tauiia iKlrrmtHiut: (rum ihc Lower l.iu of doucUMnliit*. Kolu.ol f, Kalii 
Vina: r, RaitUlc; i, ImeinieJiunti /, ftnart: <i, n, CmiralU. (Aftii Ha«'>i< 


by Dr fiaur nvith th« radiuji, ulna, and pUifoim ; the 
ing bones in the pelvic limb (fig. 1034) being the tibia, fibuU 
ihc Kotnolo^ue of the pisiform. The clavicles differ from ttv 
Ichihyoiaurui in bciiig Btparate. Both limbs appear 10 have 


width, and the third digit, cw the one arising from the 
iiim, at Idst in ihc American genus, 

of two longitudinal rams at honex. 
c spcaes of O/^tha/nwsaurut was of 
! sixe, and occurs in the Oxford nnd 
re Qayx ; it is churactoHscd by the 
} m the size Df tlic ihrte distal facets 
wncnu. In O. iantiibrigiensis, of ihc 
|K Cfeenund, whtd) inay belong to 
i'-n, dteac three facets have, however, 

neviy equal in luie ; and this form 
iffcfcn to radicate the highest evolu- 

die ovdcr. It should l>e nitrniioned 

fessor MsTvh and Mr Kulkc difTvr from 

r in their inieqireiaiion of the three 

r tbe >£Corxi segment of ih« limb of 

US ; correlatinij the middle bone with 

raicdiutn, and the posta.'nal one with 

i in the pectoral, and the fibula in the 

iob. Finally, it should be observed 

bnar, frum the Cambridge (irccnsand, 

U under the name of Cefankr^taurut, 

uUed as belonging to a member of the present order, but 

N b«cn referred hy Mr Hulke to the suborder Pythono- 


•I : rroiii ihc l.oym Liu 

1 aMHsJ fin. «. Huafnit ; «, 1'rocta*BMrio rids* of <!«. ; r. Kiiliin ; 



morpha of the Squamata, appears to approximate in Mr 
ihc femur nf IihiAyosaurus mmpyled^n, and may therdi 
hflong to this onler. 

Order V. Pbotkrosauria. — The genua /^rcttretaunts^ 
Middle Permiaii of 'I'huringia, is regarded \iy Professor 
presenting such peculiar features that it is entitled to or 
Itneiion, aliliough ))r Hnur woiikl include it in the Khynchi 

The skull is very i 
known, and aJthough 
Scelcy has attempted il 
tion. Profcivsor Credi* 
oui that the specimen! 
ju-^ttfy the figure. Thci 
doubt as to the mode 
mcnt of the teeili to 
but they appear to h 
anchyloscd to the bo 
(-avitics beneath thcra, 
not, as has been 
implanted in distinct 
Teeth also occur on 
time, pterygoid, and vca 
Professor Sceley conadi 
the palate was closed 
dorsal vertebrae arc 
ous, and have no ii 
hut the ccrvicals appeari 
lieen optslhocoelous, 
markahle for their 
were imcrcenira in ifc 
cervical region. In all J 
tebrx the arches were M 
to the centra ; while tit^ 
xAi the costal aitic 
placed unusually hig 
posterior caudal rerlel 

'-> O O .; 
^^0 >J 


Kic- ]^u-~i^>r^'ki u^T^^' ^ ^^' 1^'^ y^^ 

limb of l<it*lxm*iim hMhiu; frea Ihc Upper 

Ju/fcuK c.r N'unh Amcrka. Koluml. T, 

Tibia: I. l-'ihula : f. Iton* ntinuntini ih* 

tiUjfvnn L»f Tilt manu« ; t^ TibUk ; r Ocf* *i^^')t 

InivrMviltKiii : i (ritfhr \iil<), Kilnilftr* ; / Vn^ j- -j j i * 

ilctcirmincil brini. Tli> l>rn Hnnu Iniviath <h« dl^lOed tieUTal SptnCS. 

Mid Huikc > inal nils were lully 

(fig- »ojs), and were 
of the Rliynchoccphfllian type. It has been suggcaled I 
itium may have had a prcacetabular production; but the 
girdle 5cems to have been of the Rhynchoceplialian 
(.tavick's nnd inicrclavitle, according to Dt Credner, 
bling those of Sp/i^aodon, 

According to Dr Baur there are two centralia in 



*rofeK«or Seetey describes a centrale (navicular) in the Ktrsuit. 
ctoral Umb is conadcrably shorter ihai ihc peh-ic (left uide 
1035) ; and while boih ha%'e five digits neither presents any 
nmcly marked divergence from the Rhynchoccphahan plan 

ore. In the type species there were the normal two sacral 
; but according to Professor Seck-y, P. Linki diflcis from 

• known members of this branch in having three sacrali. 
Itghly prob^ile that Pny/ftvMurus should be regarded as a 






Fif. ■■u.~P»rt of (he tninfc awl peclartt lioUt of Prvlfimitrut 
Sftatff Ifraaa ite l^miiii'i uf Tliuriniiij. Reduced. (Alter vni 

Specialised Rhynchocephalian. The imperfectly- 
known genus Haptodtii^ from the Peniiian of 
France, may not improbably indicate a Reptile 
more or less nearly allied to ProUrosatinir, 
although on the other band it*; affinities nwy be 
with the Rhynchocephalian PalaohaUeria. 

Order VL Rhvnchocbphalia. — Thjs order 

is represented at the present day solely by thu 

remarkable genus Sphenotion {Hatteria), and may 

lie provisionally characterised as follows: The 

mal appearance is usually more or lcs.<i lizard-Uke. In the skuU 

quadtate is immovably fixed by its proximal extremity, and 

a by suture with the pterygoid ; an inferior temporal arcade 

resent (fig. 1039); the po»torbiial, at least in Sphenodon^ \% 

ie di»tincl from the postfronlaJ ; the (ulalc is closed anteriorly 

the iDedian union of the pterj'goids, which, in the living form 

east, extend forwards to meet the vomere, and thus separate the 


palatines ; ^ and the premaxills: never united. The denti 
usually acrodont. The ribs may have uncinate prtxresses, i 
dominal ribs are always develof)ed. The vertebrae may be 
opisthoccelous or amphiccelous, and intercentra may be letiit 

Dr Baur observes that " the Rhynchocephalia, together *: 
Proterosauria, to which they are closely allied, are certab 
most generalised group of all Reptiles, and come nearest, ii 
respects, to that order of Reptiles from which all others tod 

This order, if we include Palaohatteria, dates from the F( 
but only comparatively few forms are at present known. 1 
be provisionally divided into three suborders. 

Family Pai.*:ohatteriidj«, — The genus Paleeohatteria, 
Permian of Germany, is regarded by Professor Credner, ite do 
as so closely allied to the existing ^henodon that it is refa 
him to the same family. There can, however, be no questi 
that it is entitled to be the representative of a distinct (ami! 
probably of a distinct suborder, Dr Baur, with whom Pr 
Cope is in accord, would even go farther, and remove this 
altogether from the Rhynchocephalia, to place it with Mta 
in his order Froganosauria. There is, indeed, much to be 
favour of placing both these primitive Reptiles in a single] 
but this appears to be outweighed by the resemblance of the 
the true Sauropterygians and of the other to the Rhynchocepl 

The skull much resembles that of Sphenodon, the jugal d 
posteriorly to join the two temporal arcades ; but the supi 
infratemporal fossae are much smaller, while there is a s< 
lachrymal, and the premaxillse do not form a beak. Teetl 
not only on the palatines, but also on the vomer and pterj-gi 
in the young of Sphenodon ; and Dr Baur suggests that tht 
a parasphenoid. There are intercentra between all the ve 
in which the neural arches remain distinct from the centra 
are also two sacral vertebrae ; and the ribs have no uncina 
cesses. The teeth of the jaws were acrodont, and anchyk 
the bone. The pectoral girdle presents an approximation, 
form of the clavicles and interclavicle, to that of Sphenodo 
the expanded proximal extremities of the clavicles recall the 
thoracic plates of the Lab>Tinthodonts, and the coracoid i 
like that of the Sauropterygians. This bone, indeed, like thi 
of the pelvic girdle, ossifies by radiations from the centre al 
manner obtaining in Sauropterygians and Amphibians. The 
girdle is widely different from that of Sphenodon ; the pub 
ischia forming wide flattened plates like those of the Sauropt 

' The same arrangement obtains in the skull of Nothaiourus, repm 
fig- 991- 

and the ischia being nlmont identical with those 
jfUophibU. The tanus also resembles that of M/s^saums in 
Rtc distinct bones in Ihc distal row; while the humcius (as 
kltcr) ha*, an entepicondylar, or ulnar, foramen. 
'■I*aitrokatt€ria is a primitive type conntciing the taicr Rhj-n- 
hftlik with the Amphibia there can be no reasonable doubt ; 
bough it presents many points of affinity with M<sosaurtt$, 
escmblances arc those which we should expect to find in all 
Hul tj-pcs, and do not necessarily imply that all the Toims in 
hey occur should be placed in the same group. 
HtDEtt 1. Si]iLfiiK>SACxiA. — This group is represented by the 
CiMmfiosaumf, typically from the Lower Eocene of North 
iBi, ■whith may \k regarded as a somewhat specialised Rhji)- 
halian, showtnj^ a remarkable aAinity, in the general structure 
Mcull and vertebral column, to Ilyptroda^d^n. Dr Hauc ob- 
thai this genus agrees with ihc other Rhynrhoccphalia in the 
Oodition of the otic bones, and in the nature of the coatat 
Ittotu ; and since ii h,xs a fixed quadrate, two temporal ar- 
■nd altdominal ribs, there appears no good reason for itx 
jaeparatioR. The adaptation to an aquatic life has, hown'cr. 
jffly prodt]ced considerable structural modifications of secon- 
^pon. The correctness of these observations has been re- 
)ly shown by the researches of M. UoUo, who finds that the 
re of the palate, and the position of the posterior narcs, and 
hcial arrangement of the teeth, closely accords with that 
^ in Hypavda^don. The suborder may be characterised 
hrs: The facial portion of the skull is produced into a long 
b; the splcnial bone enters into the mandibular symphysis ; 
% and pelvic limb^ are elongated : the vertebne are amphi- 
I; and the ribs have no uncinate procesites. 
ULT Champsosaurid^.. — In the type and only known family 
res are single and subterminal ; the maxillary and anterior 
bular teeth are large and not fixed to the bone : there is a 
>r tnudler teeth on the palatines and vomers, seimrated by a 

from those of the maxilla ; while tlieie is also an irregular 
t small teeth on the ptcrj'goids, which arc completely united 
middle line ; the posterior narcs form very narrow slits on 
les of the palate ; and there is no parietal foramen. There 
nc twenty-five presacral vertebra;, and the ncuro-ccntral suture 
istent. Renuinii referred to Champsoiaums have been fotind 
liy In the Lower Eocene of North America, but also in the 
fondtng horizon of Relgtum and Rheims ; the latter s^iecimcns 

been described under the name of Simftcioidarus. One of 
ecimens from Rheims indicates an animal of about nine feet 
but other examples appear to have been of considerably 



palatines ; ^ and the premaxillas never united. The dentitiai 
usually acrodont. The ribs may have uncinate processes, ud 
dominal ribs are always developed. The vertebrae may be 
opisthocoelous or amphicoelous, and intercentra may be letaioed 

Dr Eaur observes that " the Rhynchocephalia, together with \ 
Proterosauria, to which they are closely allied, are certaiii^i 
most generalised group of all Keptiles, and come nearest, in ■ 
respects, to that order of Reptiles from which all others took i 

This order, if we include PalaehatUria, dates from the P( 
but only comparatively few forms are at present knowiL It i 
be provisionally divided into three suborders. 

Family Pai-eohatteriida:. — The genus PalaohatUria, di 
Permian of Germany, is regarded by Professor Credner, its desoS 
as so closely allied to the existing Sphenodon that it is refeird 
him to the same family. There can, however, be no question 
that it is entitled to be the representative of a distinct fomilT, 
probably of a distinct suborder. Dr Eaur, with whom Profa 
Cope is in accord, would even go farther, and remove this ge 
altogether from the Rhynchocephalia, to place it with Mesoaa 
in his order Proganosauria. There is, indeed, much to be said 
favour of placing both these primitive Reptiles in a single gw 
but this appears to be outweighed by the resemblance of the one 
the true Sauropterygians and of the other to the Rhjmchocepl 

The skull much resembles that of Sphenodon, the jugal di' 
posteriorly to join the two temporal arcades ; but the supia- 
infratemporal fossEe are much smaller, while there is a 
lachrymal, and the premaxillse do not form a beak. Teeth 
not only on the palatines, but also on the vomer and pterygwds* 
in the young of Sphenodon ; and Dr Baur suggests that ^ere «■ 
a parasphenoid. There are intercentra between all the vertebn 
in which the neural arches remain distinct from the centra \ thtJ 
are also two sacra! vertebrse ; and the ribs have no uncinate pn 
cesses. The teeth of the jaws were acrodont, and anchylosed 1 
the bone. 7'he pectoral girdle presents an approximation, in d 
form of the clavicles and interclavicle, to that of Sphenodon; bi 
the expanded proximal extremities of the clavicles recall the btd 
thoracic plates of the I^byrinthodonts, and the coracoid is ma 
like that of the Sauropterj'gians, This bone, indeed, like the boo 
of the ])elvic girdle, ossifies by radiations from the centre after ti 
manner obtaining in Sauropterygians and Amphibians. The peh 
girdle is widely different from that of Sphenodon ; the pubes ai 
ischia forming wide flattened plates like those of the Sauropterygi 

' The same arrangement obtains in (he skull of Neiieiatmts, icpresenled 
fig. 991. 



ixiwmnts, and the ischin being Almost id^ntic.^ vith those 
Unphibia. Tbc tarsus also resembles ihat of Afesosaums in 
fe dutinct bones in tKc disinl row; while the humerus (as 
tier) has an entepicondylar, or ulnar, foramen. 
^'aiachatleria it< a primitive typeeonntciing ihc later Rhyn- 
dia with the Amphibia there can be no reasonnhlc douht ; 
ough it presents many points of aifinity with Mesosaurus, 
(cmUiUKCs are those which we lihould expect :o find in all 
ul types, and do not necessarily imply chat all the forms in 
ey occur should be placed in the snme group. 
IDEB I. SlM,r.r>OSAUBlA, — Tliis group is repffsenied by the 
hainpsoiaurmi, typically from the Lower Eocene of North 
I which may be regarded as a somewhat specialised Rh)ii- 
llian, showing a remarkable affinity, in the general structure 
uQ and x-ertcbral column, to Jlyptrodapedon. I)r Raur ob- 
At ibia genu-) agrees with the other Rhynchoccphalia in the 
Kiition of the otic bonex, and in the nature of the costal 
oru; and ^mce it ha& a lixcd quadrate, two temi^oral ar- 
nd alidominal ribs, there appears no good reason for its 
•eparation. The adaptation to an aquatic life has, hovi-e\'er, 
ily produced contkidentble structural modificationx of &econ- 
►ort. The correcincss of these observations has been re- 
f »hown by the researches of M. Dollo, who finds that the 
I of the palate, and the position of the posterior nares, and 
sal urangement of the leclh, closely accords with that 
g in JJy/tradapedon, The suborder may be characterised 
■s: The facial ponion of the skull is produced into a long 
; the splcnial lx)ne enters into the mandibular symph>'5is ; 
and pelvic limbs are elongated ; the vcricbne are amphi* 

and the ribs ha%-e no uncinate processes. 
A CHAMPSOSAVRiii.t. — -In the type and only known family 
IS are ungte and subterniiunl ; the uiaxtll.iry and ^interior 
llu teeth are large and not fixed to the bone ; there is a 

stnaller terth on the palatines and vomere, separated by a 
Tom those of the maxilla ; while there i.s also an irregular 
small teeth on the pterygoids, which are completely united 
liddle line ; the posterior narcs form very narrow slits on 
% of the palate ; and there i-i no parietal foramen. There 
t twenty-five preMcral vertebrw, and the neurocentra! suture 
lent. Remains referred to Chitnifinnaurus have been found 
r in the Lower Eocene of North America, but also in the 
nding horizon of Belgium and Rhciins : the latter specimens 
>ccn described under the name of Simtniflsaurus. One of 
rimcns from Rheims indicates an animal of about nine feet 
I, but other examples appear to have been of considerably 



larger dimensions. The Simsedosauria may perhaps be 
an ofTshoot from a stock related to Hyperodapedon. 

Suborder 2. Sphenoiwntina. — This group is chat 
the short and more or less triangular skull, in which the 
are produced into a distinct beak ; by the longitudinal 
palatine teeth, separated by a groove (into which the hinder 1 
bular teeth are received) from those of the maxilla (fig. 1037); i 
by the presence of uncinate processes to the ribs. 

Family Rhvnchosaurid*. — This is the most specialised : 
and as being most nearly related to Champsosaurus, may be ; 
first. The nares are single ; there are no teeth either in die \ 
or in the opposing part of the mandible, which were probably < 
in horn ; there may be more than a single row of palatine 
and the presacral vertebrae may be more or less opisthc 

Fig. lOjS.— Lefi lateral view of ihc okull of Hyttrtdaptdtn Gtrdtmi, as tntorHl by Prd 
Huilcy ; from the TrUs of E)|{in. Reduced. Or, Orbit ; /, Infmein[iofal Icwt 

The most specialised genus ffyperodapedon was originally aait\ 
known to us by some very imperfect specimens from the Keupci^ 
or Upper Trias of Elgin and Warwickshire ; but the subseqneot 
discovery of a nearly entire skeleton in the former locali^ his 
enabled Professor Huxley to illustrate its full affinities. Remiilll j 
referred to the same genus also occur in the Maleri stage of At 1 
Gondwana system of Central India. The European species attained 1 
a length of from six to seven feet, but the Indian form must hate ! 
been nearly or quite double these dimensions. The skull (fig. 1036) 1 
is remarkable for its depressed and triangular form, in which it ns 
sembles that of the Chelonian family Chelydridx ; and also for dK 
upward direction of the small orbits ; the reduced size of the infra- 
temporal fossas ; the strongly-cur\-ed and thick premaxillary beak ; 
the diverging clawlike processes of the mandibular symphj-sis ; and 
the absence of a parietal foramen. Professor Huxley considered that 
the forked extremities of the mandibular symph}'sis embraced the 
premaxillary beak, as is shown in the figure ; but later observations 
indicate that they were received in a pit beneath the beak, which Pro- 
fessor Huxley regarded as containing the apertures of the posterior 



he U«cr probably really fonned very small slits on the sides 
Btc, as in Ckampsosavnis, and opened l>eiwwn the rows of 
and palatine teeth. By far the most remarkable feature 
tr, to be found in the upper den- 
a the palatal surface of the pecu- 
lar compound bone (lig. 1037), 
y be conv-enienily termed the 
Ila, allhougli anteriorly it prolj- 
odes the vomer, and posteriorly 
lie pterygoid, there are, on the 
maxillary side, several rows of 
Mnujal teeth ; then comes the 
br the reception of the edge of 
lible. Of) the inner side of which 
I two or more romi of similar teeth 
ilhc palatine and vomer, and prol>- 
by the pter^igoid. And it is in- 
to notice that while in the type 
le lar;ger number of rows of teeth 
p the palatine and vomer, the re- 
ndition obtains in the Indim form 
\l\. Ici the type species there 
K> be no foramen to the humerus, 
j presacral vcncbrn arc slightly 

rious ; it is however, not improh- 
the vertebrae oi the Indian form 
tccclous. The extremely solid structure of the pnlato- 
this part to be the moitt frequently preserved ; and 
are rery common in the Indian Gondwana^. 

Huxlc7obsen-esthiit ttis very intcreslinc to note that this 
~ atuincd itA urciiicst degree of specialisation as early as the 
fyptrfiAiptdon DeinK in all 
B more modified form than 
h appears therefore to be 
that in inc Permian, or pcr- 
earlicT, there must have cx- 
s diflertn^ less from the 
U9 than either Hyptn'^ui- 

Pi(- toiB. — Kif hi JciTBml At|>«1 of lh< 
typical uenUS ftkynch^iaU- *kMcr kkrmtlt>,»mrM4 ^nurfi: trgm 

the English Keuper, of (Ar« Omn.) 
r type skull is represented 

ijS, there is but a single row of palatine teeth ; the orbit 
and lateral; the infr-iicmpora] fossa of considerable size ; 
Vj beak loiiji and slender, iu fonn being not 

richt paUitt-muiillik of Hsftrv- 
Mtdliculi and fllanronL) 


correctly shown in the figure of the type skulL The wt 
scribed species indicates an animal about three feet in I 
This genus serves in some respects to connect J/j^erodaftdn 

Family Sphenodontid*. — The least specialised family o 
suborder is solely known by the existing New Zealand , 
Sphtnodon {Hatteria), of which the cranium is shown in % 
According to Professor Huxley's definition, this genus is cbn 
ised by the divided nares ; the presence of a single tooth on ■ 

side of the prema 
beak, whicit wh 
sheathed in hon 
the single row o 
atine teeth ; vm 
amphiccelous ver 
In the palate the 
goids unite anti 
with the voDU 
exclude the pal 

Fig. loig. — Riahl lateral view of the cnuiium of i">**iMrf«i r .». __ j' 

/■•t/a/Bj, ofNewZMland. (After Gdnlhet.) "Om the meOiaf 

Additional disti 
features are found in the presence of intercentra between J 
vertebrae ; in the large size and the lateral position of the < 
the well-developed parietal foramen ; and the large siae of the 
and infratemporal fossae (fig. 1039). The humerus is rema 
as having both an entepicondylar and an ectepicondylar foran 

The Tuataras, as these lizards are called by the Maories, are m 
sized reptiles of extreme rarity, and with nocturnal habits. The a 
border of the dentary bone of the mandible is received in the 
between the palatine and maxillary teeth, and in old individuals b( 
as hard and polished as the teeth themselves, of which it eventna 
charges the functions. 

Suborder 3. Hom^osauria. — This group includes ! 
genera of Mesozoic Reptiles, in which ^e premaxillse di 
apparently form a beak, and the ribs were devoid of ur 
processes. The dentition is acrodont ; but the nature 1 
palatal dentition is unknown. The vertebrae are amphic 
and Dr Baur considers that intercentra were present. 

Family Homa:osaurid-e, — The type family is definitely! 
from the Kimeridgian Lithographic slates of the Continent, 
characterised as follows. The body is shaped like that of or 
Lizards ; the skull is comparatively broad and short, with oval 
and a complete postorbital bar ; there are no tusk-like teeth 
premaxillae or mandible ; and the pes is of normal structure. 



Sdgian genera are HomoMaurvs, Ardtosaurut, and Sapheo- 

all of which are represented by species of small or medium 

[The imperfectly known AphtJinaurus, from the Permian of 

wa^ regarded by the laic Professor (Icrvaus as allied, but it 

licate a distinct family, which does not belong to this group ; 

of phabngcaU is the same as in existing Lieards. 
MLV PlxUROSAURiD.«, — This family is typically represented 
<ys<ttifits, of the Kimeridgian of Havana, which is a medium- 
characte^i^ed by the extreme elongation of the body 
Ethere are a great number of presacral vertebrae), and the 
narrow skull, with blit-tike nares. An^isavrus and Aero- 
mt, of the same deposits, belong to this family; but it does 
fcfifii II cenain that the}- are teally distinct from the type 

BuLt Tklerpetid^ — The small Tekrpcton, of the Upper 
m of Elgin, differs from the Hemtrosaurida by the presence 
ttsk-Iikc teeth at the extremities of boih jaws, and the reduction 
he number of the phaLnngenU of the fifth digit of the pes to 
; in addition to which Professor Huxley considers that the skull 
: no pfMtorbital bar. The one species is of small size; and 
ongh the genus agrees with the HamtEcsattrxda: in its acrodont 
tition, its scnal position must be regarded as provisional 
wtsHmmm, from the Karoo system of South AMca, is not im- 
btbly an allied genus ; although it has t)een referred by Sir R. 
en to the Amphibian t^byrinthodonts under the name of 

'ioaUy, it should be mentioned that Professor Cope includes in 
pieseni order the genera T\ft>lhorax and Ailosaumsy which are 
\ praviuonally placed, on the authority of Dr Daur, among the 

)>l>Eii W\. Sqi;amata. — With this order we come to the con- 
tation of the one represented most numerously at the present 
, and containing the true Luaids, the Chama;leons, the extinct 
ta&aurians, and the Serpents. In this order the body may he 
*r short, with wcll-de\'elopcd limbs and distinct tail (tnccrti- 
a) ; Of it may be extremely clnrgated, without any external trace 
liinbs. and posaing gradually into the tail. As a rule, the whole 
ly and limbs arc covered with overlapping homy scales; and 
w nuiy be underlain by an armour of bony dermal scutes. The 
bs nuy be adapted cither for w.-ilking on land, or modified into 
die* for Fwimming. In the skull the proximal end of the qtiad- 
■ (5 more or less movably articulated ; the lower temporal arcade 
lanting ; the postorbital is generally fused with the poslfrontal ; 
palate is more or less open, (he pterygoids being nearly always 
ted by an iRter\-al from one another ; and the prcmaxillse are 

1 138 



Pig. 1040. — Part of ihe left 
pectonl girdle of IguaMa. i, 
Scmpula : m.ic, MexMopula : 
cur, Curacoid; f^cer, Precora- 
coid; m.ctr, MESOCoraccHd I^r, 
Foramen ; gl. Glenoid cavity. 

frequently united. The vertebrse are generally procoelous, a 
rarely amphicoelous ; their neurocentral suture is always oblit 
zygosphenal articulations may or may not be present ; an 
centra are always wanting. The dorsal ribs never have a 
processes; and true abdomiiuJ r 
likewise never developed. The 
has but a single centiale ; and t! 
coracoidal process (fig. 1040) is oA 

Three of the groups here r^a 
suborders of the Squamata are fre 
ranked as distinct orders, but their 
relations are so close that it app 
harmonise better with the classU 
adopted in other branches of the ( 
include the whole of them in a 
order. The Squamata may be n 
as occupying a position among 1 
somewhat similar to that held 
Teleostei among Fishes, and the 1 
among Birds. That is to say, t 
essentially typical Reptiles, whic 
attained to a considerable degree of specialisation ; and wh 
have lost, on the one hand, all signs of kinship with the Amp 
they exhibit, on the other, no traces of especial relationship \ 
Birds. That this order has originated from the Rhynchoi 
there seems but little doubt ; but we are very much in the 
to when or how the divergence took place. 

Suborder i. Lacertilia. — In the true Lizards the fot 
are usually well developed (fig. 1041) but in some cases one 
pairs are wanting. The ali- and orbitosphenoidal region: 
skull are imperfectly ossified ; the superior temporal ar 
generally present ; the quadrate articulates with the ptt 
the nasals form a part of the narial aperture ; and the ram 
mandible unite by suture. The vertebrae are in some ii 
amphicoelous ; they usually have no zygosphenes, and the 
in the cervical region does not exceed nine. When Ui 
present the pectoral girdle is complete ; and the terminal 
geals of the feet are clawed. Dermal scutes are sometimes ] 
and these may be developed on the upper surface of the : 
as to roof over the supratemporal fossae. Existing Lis 
divided into twenty families, but only those will be notice 
occur in the fossil state. 

We may commence our notice with a few Mesozoic U 
which the family position is not yet determined. The 



appenrs to be Matellodus (with which SaurHIus is 

ilical or very clascly allied), from the English Purbeck ; 

IJKard with pleurodont deniition, dermal scutes, and pro- 

tcbne. Adriotaurui « a larger form from the Ixiwcr Grccn- 

VAtntrio, also having deniul scutes and proccelous vcrtcbnc, 

vhich the dentition is unknown. An allied Contincnial 

fonii has receix^-d ihe name of Jiydrosaurus, which is, 

a synonym of Varanus. In the Cambridge tlrccnsand 

feci fcinur and a vertebra indicate a Lizard of the st^e of 

flbe existing Monitors, but of which the affinity cannot even 



P>(. )a)i.— Tlw < 

tSlatUiStiiKmnfliimtldi). RvdutwL 

utured In the English Clialk occur the imperfectly known 
savrus and Rafkiosaurni ; the former having expanded, 
laiia acute teeth. 

' also may be mentioned several genera from the Tertiarie:i 
America, of which ihe family position cannot yet be de- 
The Eocene foims have been namc^ TinQsaurus and 
It, and those from the Miocene Adprion, PJatyrhachis, 
ifMaurus. Notosaumtt from the Tertiary of Brazil, has 
loua vencbra, and may tw a Rhynchocephalian. 
LV AcAMiD^ — The Agamoid Lizards constitute a large 
[mainly characteristic of vVsia, but also occurring in Europe, 
[Australia, and Polynesia. The supratemporal fossa is not 
over; the dentition is acrodont; and the prcmaxillie arc 
SpecimenK referred to the type genus Agaaia have been 
ed from the Upper Eocene Phoaphorites of Ci.'iitral France; 
Pleistocene of Australia has yielded a &kuU indistinguish- 


able from that of the existing Frilled LiEard (Ch/amydasattru 

of the same country. 

Family lGUANiDi£. — The Iguanoids are readily distinguist 

the Agamida I 
pleurodont d 
(fig. 1042), a 
presence of zyg 
al articulations 
vertebrae (fig. 

Fi(. I04>.— Inner view of Icfi nmiu of ihc muidiblc of Nearly all the 

are now confii 
the New World. In the Upper Eocene (Oligocene) Phosphc 
Central France there occur, however, fragments of jaws whi< 

been referred to the typical Ai 

y^Mff I ^^ genus Iguana; that tenn bein 

YW~ I0s|fc"lfie5 '" * wider sense than in recent 

A» T^T r-Q^jjl ol(>gy> Vertebrx also occur 

T approximately equivalent depc 

Til. 104}- - Vjrtebr. of /««« Hordwell in Hampshire (of wh 

tnrvfra, viewed from ihc btemal, »n- . . ■ » ... 

tcrior, uid i*ieni upecH ; frain the IS represented m ng. 1043), WOIC 

Upper Eocene of Hampshire. «, Zy* ;_ .v_ ^r 11 

si^ene ; e, co.«i luilenrie. '" the presence of Small ZygO! 

with those of existing Iguana 
have been provisionally referred to the French species. Th 
Iguanavus has been applied to remains of a member of this 
from the Ex)cene of North America. 

Family Anguid*. — The Anguida are characterised by tl 
ence of scutes roofing over the supratemporal fossae ; by Uie 
tion of the premaxillEe and the nasals ; the more or less con 
pleurodont dentition ; and the presence of dermal scutes 1 
with minute tubercles. The genus Ophisaurus {Pseudopus), ii 
the limbs are either wanting or are reduced to rudiments of th' 
pair, is represented in the Middle Miocene of Gers, in Fran 
the Lower Miocene of Rott, near Bonn ; some of the species 
been originally described under the name of Anguts. Prapst 
from the Middle Miocene of Steinheim, in Bavaria, is distin] 
from Ophisaurus by its stronger dentition and the present 
double row of teeth on the vomer. The genus Peltosaurus 1 
allied Exostinus, of the Miocene of North America, may be it 
in this family ; to which we may also refer several Europ< 
American Upper Eocene genera which have been regarded 
stituting a distinct family under the names of Placosauri. 
Glyptosaurida. Among these Placosaurus (Palaouaranus 
the Upper Eocene of France and England, has teeth test 
those of the Slow-worm {Anguis\ which were originally rega 
belonging to a Varanoid ; the vertebrae (fig. 1044) are not 


of the existing DifilfghssMs, and ihe limbs were well de- 
Another Anguoid, with blunt cylindrical teeth, from llie 
Phcwphoriie*. was originally referred to the Scincoid genus 
{PUstivJam), but its gciKiTtc position must for the present 
[I Lirilccided. In Nonh 
i»e have dypiotau- 
from the Btidger Eocene 
"^V-r.ming, in which ihe 
feclT.c have rudtmcnta] 
eflet; the large Sa- 
of which the dermal 
are unknown ; and 

rig' i4pi«4. ^Vvrt^bFA af Ft*(^t*»t^t mfptrili' 

rnr _. 

ciM- *W>vl (nini the hamat, i.ouiior. una lawnl 
> i>' I ir . . vukU: (rwn Ike Uppit Evan* PlioiDtiiinla oT 

^iparcntly aJhcd Xtshpt Ccnml rmm. c. Co>ul lutwcle. 

9tints\. The vertebrae 

genera, like those of C3[istiiig Anguidtc^ present a strong 
BbUnce to those of Ihe next family, as may be seen by com- 
fig. 1 044 with fig. lo+s. 

iii-T Varaxiixc. — The Monitors include the largest known 

I ; one of the fosiiil species attaining a gigantic sise. They 

to Ihe Old World and Australia ; and appenr to be 

before the Pliocene. The skull has no postorbital bar; 

Qporal fo&sa* are not roofed over by bony scutes ; and 

premaxilke and the nasals arc united. The dentition is 

It ; and the teeth are large and pointed, and confined to 

The vertebrK arc characterised by their broad and flat 

(fig. 1045) ; and dermal scutes arc wanting. .\11 the known 

ouy be included in ^e single genus Varanus. 

la Fiittinif fomis the dorsal verlcbrar arc elongated, and have broad 

tual »iiincs: and the largest species attains a length of seven feet. In 

bnl state ibis group is represented 

[rouuds from the Pleistocene of Ma- 

M a|)parently rcrcr;iblc to ihc living 

\ttngatemis. Other vcrlcbrj?, again, 

IbA in the corrcupondint; cavc-dc- 

o( Queensland, not improbably 

lo one or nwrc of the species 

■biting that region. In a 

and extinct group the dorsal 

K XIX reliitively slwrter and 
bAn. and lui e narrower neural spines ; 
K two known species being of very 
'Wft sue. The smaller of the iwo is 

tF. mmifnat, from the Pliocene of the 
ftnlik Hills of India, nf which a dor- 
renebm it sliuwn in fig. 104$. Anuiher veilcbra is, however, larger 
1 the figured s{iecimen ; and the toial lengih of the ;inin:>.il was prob* 
Hf «i least twelve feet These dimensions were, however, );reatly ex- 
(Mied by the huge K priscut, of the Pleistocene of Australia, in which 

Kig. iw.— llciiul aipcci at 1 donal 
flioecne of Ibt StvalU, llillt. 


the vertebrae are three times the size of those of V. iival€mu,i 
total length did not probably fall short of thirty feet. This : 
originally described by Sir R. Owen under the name of Me^ 
tain remains which were referred to it having subsequently _ 
be Chctonian. An imperfectly known species, from the Lower 
of Attica, may perhaps belong to the former group. 

It may be well to mention here that Sir R. Owen has i 
two peculL-ir blunt and pleurodont teeth of a large lizard 1 
Pleistocene of Queensland under the name of NotiasoMna, \ 
is, however, preoccupied by the genus Notosaurus (p. 1139). 
just possible that these teeth may be referable to Varania^ 
in which event the generic term Megaiania would have to 
tained for that form. 

Family TeiidjE. — In this family, which is confined to , 
the suprntemporal fossae of the skull are not roofed over, thei 
tion is pleurodont or subacrodont ; the teeth, although 
form, are always solid at the base ; and there are no dennal 1 
An existing species of T^pinambis is represented in the Pleis 
cave-deposits of Brazil. 

Family Lacertid.^. — In t)\e Lacertida the supratempoialj 
are roofed over by bone ; the premaxillx are united ; the ' 
is pleurodont, the bases of the teeth being hollow ; and there aiei 
dermal scutes. All the genera belong to the Old Worid. Re 
of the existing Lacerta oeellata occur in the Pleistocene of Fn 
and extinct species referred to the same genus have been de 
from the Miocene and the Upper Eocene Phosphorites of the 1 

Fa-mily SciNCiDiE. — The Scincoid Lizards form a large coait; 
politan family, characterised by the bony scutes rooting over A^ 
supratemporal fossse ; the separate or imperfectly united premanDc; 
the pleurodont dentition ; and the pr^ence of dermal scutes. Dn- 
catiosaurus, of the Lower Miocene of France, is an extinct gem 
with moLiriform teeth, probably allied to the existing Scinaa (fi|. 
1041) or Chalddes. 

SuBOROKR 2. Rhiptoglossa. — The Cha/wleontida, or Chanfr 
Icons, differ from the Lacertilia in that the nasals do not enta into 
the borders of the nares ; the pterygoid does not articulate with tiie 
quadrate ; and, although limbs are present, there are no clavicles or 
intercla^'icle. The dentition is acrodont. All the existing fonni 
are Old World ; but Dr Leidy has described part of a mandiblt 
from the Upper (Bridger) Eocene of Wyoming, which he refers to 
the type genus Chameleon. 

SuuoRDER 3. DoLicHos.^URiA. — This group was originally formeJ 
for the reception of the genus DoHchesaurus, from the Englid 
Chalk ; which is a small snake-like Lizard, with more than nin 




in ihe neck, and welI-d«v«!oped limbs. The vcrtclirse 
. zyfCosphcful lubculatioru. Acteosaumi, from the Cretaceous 
is an allied form. 
iRitER 4. P^THOKOiloiiPHA. — The MosasauToids are car- 
tLrLs marine Repcilcs, freqiienily of large dimen-sionis and 
in lime from ihe Upper Grc-cnsand to the topmost Crcla- 
wiih a cosmopoliun di»tril)ution. The body \i much eton- 
The skull (fig. 104;} pretcnls a strong resemblance tn that 
VartMHiJa among the Laccrtilia, and has the iu&.tls and pre- 
welded togwher, the quadrates 
looirly articuUlcd, teeth on the 
J« as irell as in the jaws, and 
itly ossifications in the sclerotic 
eye. The teeth are large and 
jJkI anchrlosed by expanded 
to Ihe sammits of the jaws, 
may be xyjfosphcnal articuUliona 
»ertebr«, and the cervical r^ion 
tudude more than nine vertebn. 
davidcK are always, and the inter- 
and sacnun gmeially, wanting ; 
PtoAssor Marsh figures a sternum 
genus. The limbs arc modified 
lies (lig. 1046), with no claws to 
linal phalangeals, and no fora- 
I the horaenis. The development 
pelvis is, moreover, but imperfect, 
■ at least the majority of forms appear 
have been devoid of dermal scutes, 
gh Professor Marali has recorded 
preifence in one genu«. 
F&MiLY I'uori-ATvcARriD.e. — TTie 
■tt specialised form seems to be the 
, PiiapUtlycitrpHS, of the Upper Cre- 
pt Holland, in which both an inlcrclavicle and a sacrum 
' (ffcsent ; on which account its dcscribcr, M. Dollo, regards it 
type of a distinct family. 
kMILY VfosAMUktD.CL — 'hie whole of the remaining genera of 
sborder may l>e included in ihis family. One of the well- 
genera is CiiJastti (from which Edahiaurut \s, regarded 
Cope as inseparable), fioin the Cretaceous of N'orth 
His gcnos comprises numerous species, characterised 
the extreme elongation of the body, and their small or medium 
lice. In the skull the teeth, as in the next genus (lig. 1047} are 
(mtinued to the extremity of the premaxills; the Tcrtcbrse arc 




rf Piatycttrf^t I from th* CratA' 
DCrUh 'jf Nurth Ain<nc%. Ort<- 
iHlfih uuinl ttv. ji, Scapula; 
^. Cnricoid ; f, Humvni* ; d. Ha- 


long, and have zygosphenal articulations ; the humenu is 
and the chevrons are anchylosed to the centra. SirMuiMl 
allied genus from the same deposits, in which the cherromi 
anchylosed to the vertebra. Flatycarpus^ {Lestoseuirtts, 
saurus), -ft-hich is found in the Cretaceous of North America ] 
Zealand, and perhaps Europe, differs from both the 


rie> 1047.— Fronlsl aipccc ot <bc cranitim of fJaljcmrptu airtimtrii ; fion ttM Dir 
lafCBiu ef North Anuna. Greuly nduccd- Juijr. PwnusLUk ; mlt, MmnUtl/r, ' 
prf, PnlVoDtal, (AfiuCspe.) 

genera by the absence of zygosphenes ; the chevrons being i 
and the humerus (fig. T046) short and broad. With the 
Licden, in which Professor Cope includes Tyhiaurus (J 
saurus) of Professor Marsh, we come upon forms attaining gig 
dimensions, in which the body is proportionately shorter 
in the above-mentioned genera, and the vertebrae are 1 
without zygosphenes. In Liodon itself the extremity of the 
maxilla; is devoid of teeth, and farms a cylindrical lostnim; 
teeth are smooth aiid more Or less laterally tompressed; the' 
merus is long ; and the" chevrons are free. It occurs in the Cnli- 
ceous of Europe, North America, and New Zealand. HainoiOMm, 
again, of the Upper Chalk of Mons, agrees with Uodon in its edfli* 
tulous rostrum and free chevrons, but has teeth of three typd 
Some of the teeth are subcylindrical, and others compressed, with 
serrated cutting edges like those of Megalosaurus. The total length 
of this huge Reptile b estimated at about 40 feet The tj'pial 
genus Mosasaurtts is definitely known from the topmost CretaceoiB 
of Maastricht in Holland, and the Cretaceous of North America; 
the type species from Maastricht having been made known to 
science in the last century. It is characterised by the prenuxiUB 
l>eing toothed to their extremity ; by the teeth having their crowtu 
faceted and more or less compressed ; as well as by the great* 
number of the chevrons being anchylosed to the vertebras. The 
fine skull of M. Camptri represented in the accompanj-ing woodoit 
was obtained from Maastricht previously to 17S5, and is presen'ed 

' Amended from Platecarfm. 

1 146 


The very imperfectly known Cetartkrosaurus, of the Camb 
Greensand, which Mr Hulke refers to this order, is noticed 1 
the Ichthyopterygia. 

Suborder 5. Ophidia. — ^The Serpents and Snakes ccHistit 
last division of the Squamata present the following distinctivel 
tures: The body is greatly elongated (fig. iosi,*/>), andthei 
column divisible only into a trunk and a caudal region. Hki 
phenoidal region of the skull is fully ossified ; there is no \ 
arcade, parietal foramen, or columella ; the quadrUe and thel 
of the palatal and maxillary regions are loosely ^tacbed to thci 
(fig. 1049); the premaxillse are more or less aborted and 
edentulous; and the two rami of the mandible are connecttdi 
by ligament. The vertebne (fig. 1050) havezygosphenes; ball 
is no sacrum, and chevrons are also wanting. There is, 
no sternum, nor any trace of the pectoral girdle or limbs ; tatJ 

some cases there are 
of the pelvic girdle and 
Dermal scutes are inv 
wanting. This suborder ii | 
vided into three sections; 
since its pala»)ntolt^cal 
is but imperfectly known, 
very brief mention will be 1 
of those families represented id 
a fossil state. With the ap^ 
tion of an imperfectly known form from the Chalk, described nad*' 
the name of Cimoliophis, all the known fossil forms are of TttiMJ 
or Post-tertiary age. The next earliest genus is Helagrvi^ of dc 
Lowest (Puerco) Eocene of North America, in which the impofta 
development of the zygantrum of the vertebrae indicates veiy gffl» 
alised affinities. 

Family ColubridjE. — The first existing family of the section 
Colubriformes, or Innocuous Snakes, contains the great bulk of the 
suborder ; and, with the exception of Australia, is representtd in 
nearly all temperate and warm regions. The Indian genus Pt}^ 
is probably represented in the Pleistocene of Madras by the eadsting 
P. mutosus. In the Middle Miocene of France we have the extinO 
Piltmophis closely related to the mcxlem Tropidonatui, ot codiukhi 
English Snake ; while the existing genus Elapkis occurs in the 
Upper and Lower Miocene of various parts of the Continoit ^ 
species of Periops closely allied to one now living in ^ypt ocean 
in the Pleistocene of Coudes, in the south of France ; and TXa* 
nophis, from the Middle Miocene of the latter country, is said to be 
allied to ElaphU. 

Family Pythonid^ — The Pjthons, or Rock Snakes, are no« 

Fig. low. — PoMcrior (a) and hsnal (■) view* 
oT ■ trunk vencbra of PjtAem mttanu; fhxn 
the Plciilocene oT Indw. 



M> Africa, Asia, or Australia ; and, wUh ihe next family, 

the Untcst existing rcprcsteniatives of the sutwrder. They 

th in the premajulla ; and all of them are good swimmers. 

of lh« existing Tndi^in Python tnelitnis (fig. 1050) arc 

the Plirisioct-ne of SUdnu ; and not improbaliiy also in 

tn« of the I'unjah; while in the Pleistocene of Australia 

chri: probably referable either to Narhoa or IJasis, which 

ihit that continent. From the Upper Kocene (Lower Oligo- 

iwater deposits of Hampshire, and the ef|uiv.-ilcnl Phos- 

af Central France, we have the genus Palet^x {Pal^o- 

which is apparently nKirly related to Pylhun. Finally, 

ktt, from the Quert7 Phosphorites, is said 10 connect the 

with the Tortriada, 

ItLV Botixx. — Tlie Boas are at the present day eunfined to 

■ World, atKl differ from the Pylhonida by the absence of 

iUary teeth (fig. 1049). The genera Beavvi, Lithapkis, and 

{JJmm^n\ of ihc Upper F.occnc of North America, arc 

fenble to this family. Frota^^i, from ihe same de- 

'also be proviMonally included in this family; to which 

suggested that BotrapAis, of the French Miocene, may 


iiLY Eavcro^ — The membent of this family are small Snalcex 

\\o the Botdu, but having a much shorter and non-prchcnsilc 

\Stafioph$s, of the Middle Miocene of I'Vancc, is rcgardwl as 

^lo ^c existing African Eryx ; while in the Miocene of North 

we have Aphelophts, Caiamagms, and O^mophis, all of 

appear to be moic or less closely related to the genus 

now inhabitii^ die same regions. 
1LV pAUTXJPHit)*. — Here we may provisionally place the 
formed for the reception of the extinct genus Paiaophh. 

Lfty*. — Vimb»» of Pmln^Ui Iffkrtu I (ttttn i)u Midille Ko«nf of Knfluoi. *i'h* 
ORial ivai ■■ v*ming. <, Caul ubcrclc ; ai, Zyiinvbiiw- 

[the Lower and Middle Eocene of Europe, comprising very 
Its, which were probably of uiiuine habits. The vcr> 


tebne (fig. 1051) differ from those of the Pythonida (fig. 1051 
their much taller neural spines, the lower position of the c 
articular surfaces, the less divergence of the zygapophyses, a 
the stronger development of the haemal ridge on the inferior ■ 
of the centrum, which often carries well-marked processei ■ 
two extremities. The type genus Palaophis is represented by I 
English species, of which the largest is estimated to han in 
a length of 20 feet In the Eocene of North America we 
closely allied forms reaching to a length of 30 feet, whidi n 
ferred by Professor Marsh to a distinct genus under the nas 
Titanophis {Dtnopkis), but which Professor Cope regards as i 
tinguishable from Palaephis. By Sir R. Owen these snake 
regarded as allied to the existing marine Sea-snakes or Jlydrcf 

Family ELAPiDiC. — The first existing family of the se 
Colubriformes Venenosi includes the Cobras (JVaia, fig. 1051 
and Coral-snakes {Elaps). The former genus probably occo 
the Pleistocene of Madras ; and perhaps also in the Middle Mk 
of Steinheim, in Bavaria. 

Family ViPERiDiE. — The present and following fomilies c 
tute the section Vjperiformes ; characterised, among other fea 
by the perfect development of the poison-apparatus. A snake 
the Upper Miocene of Switzerland has be^ referred l^ M. R 
brune to the existing genus JBitis {EcMdnd) ; having been orig 
described under the name of Coluber Kargi. 

Family C«otalid«. — The Pit- Vipers and Rattlesnakes are 
lined at the present day to Asia and America. The genus A 
dromicius, from the Miocene of North America, is provisic 
referred by Professor Cope to this family ; while iMipkis, 
the Tertiary of Salonica, has also been regarded as a memb 
the same group. 

Ordinal Position Uncertain. — It will be convenient to 
tion here two imperfectly known genera from the English Purl 
of which the ordinal position cannot at present be detenn 
They were regarded by Sir R. Owen as belonging to the Lace 
but their teeth are much more of a Dinosaurian type. The 
genus, Nytheks (Nuiheies), is represented by a species of the si 
some of the existing Varam'da, but has teeth closely resembling ' 
of the Megaiosaurida, although it is said that they were not impk 
in distinct sockets, and were anchylosed to the bone. The se 
genus, Rckinodon, is a smaller form, in which the teeth pres- 
striking resemblance to the much larger ones of the Dinosa 
genus Scelidosaurus ; they were implanted in imperfect sockt 

Here also may be mentioned the remarkable genus Atoposc 
from the Kimeridgian lithographic limestones of Bavaria, 1 
includes two species of small Lizard-like Reptiles, presentinj 



iving peculiar fcatorcs. The (nanus seems to l>c of B Rtt)7)- 
srptalion tfpe; but the pes has unl)- Tour digits, in which the 
■aggalft number 2, j, 4, 4, or the some as in the Crocodilia. 
bones of the carpus are also elongated, as in that 
the ra<liiu and ulna in Uie pcctoiat, and the libia and 
pelvic limb, are respectively in close appo).iiiDn. The 
symphysis h long ; and the dentition is ^id to resemble 
ot the Get^nidit atnoug the LacertilJa. lite above cbaiactera 
It to a curious blinding of Squfuaaiinc and Crocodilian features. 




Orders Dinosauru, Crocodima, and Ornithosauku 

Archosaurian Branch. — The three orders constituting 
branch comprise the most highly developed of all Reptilei 
those which make the nearest approach in their oiganinti 
the Avian type. They also include the largest forms jret ki 
not only among Reptiles, but also among all Vertetvatcs id 
for a life on land. The following features are common t 
entire branch. The teeth are very generally implanted in di 
sockets ; are never anchylosed to the bones of the jaws ; u 
exclusively confined to the premaxilla, maxilla, and dentaty t 
Both the pectoral and pelvic limbs are always well developed 
cranium has no parietal foramen, but is furnished with b 
superior and an inferior temporal arcade (as is shown in the 
of the Crocodilian skull on p. ii8i); the quadrate is firmlj 
among the adjacent bones ; and there is frequently no cola 
The anterior ribs have a distinct capitulum and tuberculum 
dorsal vertebne carry long transverse processes, which in 
placed entirely on the arch ; and there may be more than tw 
tebrje in the sacrum. There is never a T-shaped interclavicb 
the only indication of the precoracoid is afforded by the font 
in the coracoid, which indicates its original duality. The ha 
has no foramen, but an ectepicondylar groove may occur, 
proximal row of the tarsus comprises two bones, representii 
astragalus and calcaneum. Abdominal ribs are generally [ff 
The number of phalangeals in the limbs approximates more ( 
closely to that obtaining in the Streptostylic branch, althou 
some cases there is a reduction. The lateial surface of the n 
bular ramus may have a vacuity. As a rule the centrum ( 
atlas vertebra forms an odontoid process more or less closi 
tached to the centrum of the axis ; the arch of the former 



ported by the crcsccm-shapcd firs! inlercemnim. Generally the 
Bpd inierccntnim, like that of Bi^d)^ was likewise fused with 
ocntnim of the axis ; but in one Dinosaur (? Mega/esaums) 
I cxh.\% is a distinct wedge-like bone, 
pchris the pubis and ischium nct-cr fonn tbc broad and 
piftics found in the Synapiosnurian branch, and ihcy fre- 
umime a more or less rod-like form with expanded cx- 
(fig. 1073). The ilium (rlMtf.) frequently presents an 
fonn ; and the obturator notch is never converted into 

lt»eii VIII. DiNOSAURlA. — The Dinosaurs comprise the largest 
Reptiles ; »z%6 vhile some of thrm appro:i:ima(e closely to ihc 
of structure (Attaining in Birds, others come so near to the 
generalised Crocodilians that it is almost impossible to give 
!deSnitioa of the order that will sepAntte it Trom the latter. It 
probable that in the I-owtr Trias there lived the common 
of the two orders, and the forms from the upper di^-ision 
same period indicate not only the close connection lictwceti 
two groupi, but also show some signs of affinity with the 
recently been proposed to divide the Dinosauria into two 
from the structure of the pclris, for which the names Gr- 
and Saurischia have been proposed. If, however, this 
be cventuallj accepted, it would he advisable to adopt the 
Omithopoda for the first division, and to reaiict the term 
luria to the second, which would include the two groups here 
Theropoda and Snuropoda. In regard to the names of the 
il ^ould be mentioned that I*rofessor Cope first proposed 
Orthopoda and Goniopoda for the groups hi^re termed 
and Theropoda, on the ^ound that the relations of 
. ai)d fibula were ewcentially difyt-reiit in the two. According, 
, to Professor Huxley, this alleged difference docs not exist, 
; these t\»xae& are therefore rejeacd by him. 
I This order is entirely extinct, and may lie regarded as chatacter- 
: of the Mcsoioic period ; since it ranges from the 'frias to the 
aoit Cretaceous of Maaslricht, and the [.aramie beds of the 
States, and apfieara to have attained its maximum dcvelop- 
.in the Jurassic and Wealden. In space this order was widely 
favibuted over Europe and \uitli America ; and it has abo been 
iKt with tn India and Africa. 
Dinosaurs *-ary exceedingly in the contour of the body and 1 tmhs ; 
in sonoe instances were more or less of a crocodilian tyjw, hut 
en approach very markedly to the avian structure ; the latter 
being most marked in the pelvis and hind limbs. The hind 
cither tnoderately or excessively long ; while the body 


was sometimes defended by a bony dermal annour, w 
carry long spines, but apparently was never composed of ii 
pitted scutes. The centra of the vertebrse are veiy generally i 
ccelous ; but are not unfrequently opisthocoelous in the 
more rarely in the anterior dorsal region ; and occasionally i 
the caudals are proccelous. The neuro-central suture was 
persistent till an advanced period of life. As a general 
sacrum includes from three to six vertebrae, but 
number is reduced to two. The cervical ribs are not produced I 
spines directed antero-posteriorly ; and there are no undnale { 
cesses to the ribs. The rib-facets of the middle dmsal 
may either form a " step " on the transverse process, as in tbe < 
codilia, or may be placed on the lamina of the arch. The skull 1 
many features of that of the earlier Crocodilia, but also 
approximate in some cases to the Rhynchocephalian type. The] 
maxillie were but rarely fused together ; and the union of tbe 
dibular rami in the symphysis is cartilaginous. The teeth are | 
erally more or less laterally compressed, frequently having 
edges, and may be of complex structure; they were not 
implanted in distinct sockets. The sternal region is imp 
known, but it frequently comprised two paired bones^ which 
represent parts of the sternum. The limb bones may be 
solid or furnished with a medullary cavity. The contcoid hn i 
fontanelle, and is always short and rounded In the pelvis (I 
1060) the ilium has both the pubic and preacetabular 
well developed, the latter being in some cases greatly elongatiBdi 
the pubis always takes a share in the formation of the acet^uM 
(of which the inner wall is unossiBed), and may be directed ddj 
forwards or backwards. The femur may have its head placed dtha 
obliquely (as in the Crocodilia) or at right angles to the cond;r^] 
and may or may not be furnished with an inner trochanter. Tfel 
tibia, as in Birds, had a cnemial crest ; and the astragalus wu fi» 
quently flattened, and more or less closely applied to the lowtt Cfli 
of the tibia. 

Suborder i. Ornithopoda. — This suborder is taken 10 inchidl 
the Stegosauria of Professor Marsh, and embraces the most spedA 
ised forms. In the skull (figs. 1059, 1062) the anterior part of tfal 
premaxilla is devoid of teeth; there is no preorbital vacuity; th( 
nares are placed at the extremity of the skull ; and the teeth 
more or less complex, and are frequently not set in distinct socketl 
The vertebrse are solid throughout. The pectoral limb is considff 
ably shorter than the pelvic ; the limb bones may be either sdii 
or hollow. The ilium generally has its preacetabular process mud 
elongated (fig. 1060), although this is not the case in the type 
Campiosaurus (fig. 1052), the ischium has an obturator process 



Hu>st Striking and remarkable feature in Uic group, and one 
it difTent frum all others except Birds, i» that the sliaCt of 
b directed backwards more oc less nearly parallel with the 
, while a shorter and thicker portion in advance of tlie 
lorn projects forwards. How remarkably this pelvic stmc- 
nmmatcs to that of Dirds may be seen by comparing figs. 
id 1060 with that of the pelris of a Katitc Bird given in the 
Thus the large preaceubular process of the pubis of the 
correspondg with the much smaller but similarly situated 

-Tte to* ida nt iht pdntof OwAM'iinrt Jb/«r: from tilt Ujipei tina»(c a\ 
A 0D*4wilKb kuonl *ue. Tht uptir tviit u ihc ilium, ibai un iue left iKi 
M Ml Ik* fidu Am whiam. ( Vi*r Manh.) 

ll process in the pelvis of the Katite Birds (fig. 1 107). In 
Kscs the pubis bad no median symphysis. All the mote 
led members of the nuhorder appear to have walked habitu- 
the hind limbs alone. In thiit and ihe following suborder 
ph of ihe tibia is often nearly equal to that of the femur \ 
tiyahph^don and Cemfso- 
the former Iwnr, as in 
\ the longer of the two. 
Lv Trachodoktiu-»u — 
K1 specialised, as well as 
be most recent, familieit of 

brdcr, seems to be that typ- 

leprewnicd by the genus 

mom {HaJrvsaurus), This family, although closely related 
next, is diittinguibhrd by having tliv teeih ananged in a 
of vertical columns and articulating together so as to 

Fijc. 1013.— Tooih of Tr^cktiifH ^wmtiJi 
tram tha IJliwr OdiiCBCiut uf N«w Jenrrv. 
(Aft«c Uiay.f 


form n kind of pai'emenL The t/pe genus, of which a 
shown in fig. 1053, was first described from the Upper 
of the United Stales^ but has been subsequently recorded 

Upper Greenland of Comt 
All the dorsal vertel>ne are ofk 
coclous. five other gencra^Ti^ 
fhrn'us, Cioftodom, Jfaneclema, 
MKut, and AgathaHfuas — fitm 
I^ramic beds of North Ameno: 
sent a similar type of dendtMA 
have been referred to the ^ 
family ; the first being probably I 
lical with Trachodon. The Luq 
beds, it should be ohKerved, ^ 
to be tninsitional between ike V 
tflceous and Eocene although ■! 
In the soolled Didonius the A 
allhoiigh presenttnj{ many of the features of that of If^aMi 
is much more elongated and depressed, and has the cdcnuli 
prcmaxilltc produced in ad%'.ince of the large oares. Onhemo 

titm in i)k i*m. Rcducad. 

able to the former epoch. 

Pif^ nS}-—IaM> Mud eulcr{n)vpccu«ra Iowa lomh o( //vawiAii 
fiijin Ibe Wuldon irf Sums. 

of the Upper Cretaceous of Maastricht in the Nethertaatl 
probably be also referred to this family, although it shows H 
signs of connection with the next. 

Family Iguanobovtid*. — 'i'he characteristic features of thk 
markabic family arc to be found in the hollow hmb booes; 



and Tclaiive length of the pubis and ischium (figs. 1057, 
I'; die avion cturacteis of the Temur, which has a Inrge inner 
the digiltgrade hind-foot, rumishcd with cither three 
digits and more or lew elongnted metatarsals ; and the 
limbs. The buncs usually referred to the sternal 
ate tjpicalljr hatchet-shaped, and have been regarded by 
I autbontics as clavicles, althoi^h it seems more prohahic that 
ife cotutecied with the xiphistemuni. I'he cervical v'enebrx 
■stullj- opisthoccclous ; and the rib-facets in the middle dor&al 
vcfe pbced on the arch. I'he teeth (lig<:. 1054, 1055) were 
in a tingle row, and are very peculiar and characteiiatic. 
.they have flattened, diatnond-^liaped crowns, bearing slrongly- 
serratiiMis on tbe anterior and posterior borders, and one 

I Af>*i(uryriui(. Thf iltlal«n ; tram lh« WwMmi vf Dflfiim. 

verticad ridges, some of which nuy themselves be serrated, 
Ftfadr outer aspect. The mandible, again, presents the peculiar 
of having a horsc-shoc-lilcc prcdcniary bone at the extremity 
symphysis (fig. 1059). This prcdcniary ossification 1% devoid 
h ; while the mandibular symphysis itself is deeply chan- 
Tbc family ranges from the Middle Jurassic to the Weal- 
■od Neocomian of Europe ; its members generally showing a 
loal increase in sij< from the lower to the higher horizons. 
like the members of the preceding family, the Iguanodonts liabitu- 
■tUf nif^rted themselves on Che hind limbs, as in the accompany- 



In the imperfectly known Sphenoipeniylus of the English Wed 
the anterior dorsal vertebrse were opisthoccelous, and all the da 
had very low neural arches. In the type genui Igtiamedsu, ri 
includes the largest forms, the skull (fig. 1059) is compaoti 
short, with large and terminal nares and no teeth in the pRi 
ilia. In the typical forms, constituting the Euiguanodont p. 
the dorsal vertebrEC (fig. 1058) are amphicoelous, those b 
anterior part of the series having very tall neural aicfaea 
compressed centra ; while the sacral vertebrse are anchjioHd 
gether, and have rounded inferior, or hsmal, surfaces. Ii 
pelvis (fig. 1056) the ilium is shallow, with a sharply pointed| 
acetabular process ; while the pubis is much shorter thin 

Fij[. 1057.— Inner ufMct of ihe left femur 
of I^anoxlon ttmiisarttitsis (a), and of 
C^mftosaurut Lttdsi {a% Reduced to ume 
Ei», d, Hcid; b, Lcuer trochanter; c, 
iDuer do. ; d, Ectocondyle ; i, EnlociHidyle. 

fig. lOfS.— PsMcriorupanofuHn 
ul venebra of Igu am t d t n ttraltmrt 
fniiii ihe Wealdm of (he Ide sT WighL 
■ixth DUural lia. (After Seder-) 

ischium (which is twisted on itself), and does not form a sympl 
The femur (fig. 1057, a) is characterised by the inner trodu 
(c) forming a crest directed almost immediately backwards, 1 
its shaft is nearly straight. The foot had only three functi 
digits, of which the metatarsals were short and thick, and 
phalangeals broad and flat ; while in the manus the one phalai 
of the first digit, or pollex, was modified into a stout conical si 
As in the other members of the family, the astragalus, althougt 



to the tibia, was doady applied to its distal extremity, 
rendered the Ktmciure of the ankle-joint (as in msny 
|_lKsiibcn of the ordci) essentially similar to that of a Bird. 

I observed that the inner trochanter of ttic femur corresponds 

1 o«ie foand in some Birds, which gives ait.ichment to the 

r1 and tschio-femoral muscles, and it is interesting to find, 

ob te fvaiioM of M. DoUo, that its form in Jguano^Mt is that 

with the Avian type. 

ntnges from the Wealden to the Lower Greensand, 
lu has only been described from Europe. The two 
itivcs of the typical or Euijituanodont group occur in the 
^Wcalden and Lower Grecnsand, and comprise /. MantclHt 
Ffte Urger /. t^rmifuirtentis (fig. 1056) ; these two species being 
itinguishcd from one another by the number of the sacral 
vid the contour of the ilium and femur. 

kngth of the entire body of the larger /. ^rmstarltntis is esli- 

[ ■( abooi 33 fwt The histor>- of the j^radual reconstruction of the 

I of ibu sentu affords an instructive instance of the results u hich 

: aitai&ea by careful and patient study of ftaKmcniury remains. 

!. « J 


K-Lrtlanolatpeci oriUi)i>llar/rMHiABJ#r>uBit>V.^. !. In 'V.:,U#n 

MadindBMd. Tlu Miurior *p«1H« u ib« obo : ibc ni <!i ■■ , w\ 

.^ th* iahunfieal lona. TVc pr«il»Ui]l boa4 b *uii ;.i m, . .1 .,1 iIm 

tAfta tMhi.> 

_j labours of the late Dr G. Mantell of Lewes, in the first lialf of 
_i centiirv. a considerable knowledge was acquired of tlie Kre»'«T pf^rt 
TAc skdcion, ahbou^'h ihc structure of the pectoral and pelvic girdles 

a puule. The structure of the latter was, huwcrcr, aflct more 

oac tncfTcctual attempt, tinally solved by the kboiua of I'tofcMor 
and Mr J. W. Kullce. 

In the Wadhurst CI.iy. or Lower Wcaldcn, of Sussex, wc meet with 
ics of JguanoJoH which connect the preceding typical 

1 158 


fonns with the undermentioned genus Cam^tesaurms. I. Dmm 
a form intermediate in size between /. ManielH and I. ienum^ 
characterised by the less compressed centra, and lower ucbta4 
dorsal vertebra ; and also by the form of the ilium {6§. 10C4 
which is of great depth, and has a deep and romided posticeli 
process, while the preacetabular process has a horizontal ■ 
plate. In the smaller / Fittoni the ilium is equally deep^ h 
the postacetabular portion narrowed to a point laterally, widi 
tinct inferior horizontal plate ; while the sacrum has latenUj 

"^ ---"' 

Fig. 1060. — Lert peine bona of Lower Wulden Iciunodom. Onc-iuttcailb natml ■ 
ItiuiD oT Iguaiadim DimaeHi; F, Imperfect pnbn of do. ; /i, Itdiloe* tt I. ktlliailt 
a, Obcunior proceu of iichium. 

pressed vertebrse like those of /. ManleUt. The posterior p 
of the ilium of this species is indeed almost indistinguishable 
that of Camptosaurus (fig. 1052). Finally, in / kollingUm 
which agrees approximately in the proportions of the limbi 
J. Mantelli, the femur had a curved shaft and pendant trocl 
as in Camptosaurus (fig. 1057, b) ; while the sacrum had tfa 
tebrse remaining separate from one another, with their it 
surfaces flattened, as in that genus, with which this specie 
agreed in the contour of the ischium (fig. 1060, Tjr), which 
stout shaft not twisted upon itself. This species agreed, hot 


: t\-piral speries of the genus in having the pollex modified 
canical apiitc, and tliereby diflcrcd from CamfitosaMntt^ 
b indicating how the one genus passed into the other. It 
improbable th&t the Urge Iguanodont from (he Upper 
c oC the United Slates, deMribed as CamptosaurMs ampins, 
be referred to this group of Jguaned&M, since it has but 
bncuonal digits in the pes. 

Ii the ifeniu CamphMurui ( Ca>Hpt^>t/>ltis\ wc come to foims 

arc u&tiaUy of tnullcr size than ihc preceding. Ie occurs 

ly in ibc Upper Jurassic of North America ; but certain 

{rom the English Oxford and Kimeridge Oays, to the latter 

d) tbe name CmmHoria hu been applied, as well as a Wealden 

^ do not appear lo Ixr gencrically separable. This genus has 

afsotnewhat simpler structure than those of fguan/idou, nnd is 

r characlcriscd by the Battened hxmal surfaces of the centra 

: sacral vertel)ta!, which appear to have remained separate 

Itout life; by the short prcacctabular process of the ilium 

o^i) of the type species ; by the equality in the length of the 

Knd ischium *. b)- the pendant, or downwardl)- directed, inner 

Her of the curved femur (tig. 1057, ») : and the presence of 

iu of narmal structure in the nianus, and typically of four 

The length of the femur of the type specien is some 

The English forms arc not fully known ; but so far as 

he case they agree in essential characters with the type : the 

of the manus and pes is, howcvirr, unfortunately unknown. 

^si, of the Oxford Clay, is known only by the ft'inur {fig. 

1), which ineuures a little over one foot in length \ while in 

idgian C, PmtU'icMi (the tyjje of Cnmnoria) wc are ac- 

d with the greater part of the vertebral column, in which the 

have tall neural arches, while the ilium ha<> a long preace- 

pnwess like that of Jguanvdon, The V\'ealden C". valdensU 

ed by a femur, Laosaurus h an allied form from tlie Upper 

of the United States, di.'.tinguished by \x% amphicoelous 

veRcbrc Tbc imperfectly Vnuwn Cryptodnuo {Crypto- 

from the Oxford Clay of England, is by the 

lootncu of the femur, which exceeds a foot in length, and 

Baif^ ^taSt, with an inner trochanter like that of SgusHodcn 

irttMtii, but differs from the corresponding bone of all oihcr 

the family in the absence of a groove on the anterior 

lower erwd between the condyle*. 

ly, wo come to the consideration of ffyps//«pfjodaHf the 

and least specialised representative of the family, which 

veil known to us through the labours of Mr J. W. Hulke; 

genus, which occurs in the Upper Wealden of En^jland, the 

ttrOiCtiire of the pelvis, as well as the pendant inner tro- 

Ik s 


chanter of the femur, resemble the corresponding den 
Camptosaurus ; and there are Ukewise four functional digi 
pes, of which the metatarsals are elongated ; while the teet 
also occur in the hinder half of the premaxilla, are somewhat 
than those of Iguanodon. In conclusion, we may mentim 
perfectly known forms which are evidently related to thi 
Of these, Moeklodon, from the Upper Greensand of Ai 
characterised by the absence of the channelled mandibc 
physis characteristic of typical forms ; while Craipedodtm, 
Upper Cretaceous of Belgium, is a very small Dinosaui 
known by its teeth, which are more complex than I 

Family ScELioosAURiDiE. — This family includes a groi 
markable Dinosaurs of medium dimensions, characterised 
clad in a stout dermal armour, usually consisting of detachc 
and long spines, but sometimes taking the form of a solid 
completely covering the lumbar region. The rami of the i 
are slender and tapering, but it is not known whether a pi 
bone was present. The teeth (fig. 1061) have laterally flatti 
subtriangular crowns, with the borders carrying serrations 
or less obliquely or parallel to the long axis of the tool 
anterior and middle dorsal vertebrae differ from those 
IguoHodoniidtE in having the articulation for the head ol 
forming a "step" on the transverse process, as in Crococi 
not a facet on the arch ; while there is also no fossa beti 
transverse process and the postzygapophysis. The !imb-l 
solid and massive ; the pre- and posta< 
processes of the ilium very long ; and t 
and ischium comparatively short TTi 
has an inner trochanter ; the metatai 
short and thick ; and the pes, which was 
plantigrade, always has four functions 
This family connects the Iguanodontida 
Sttgosauridte, and appears to be confine 
Old \\'orld, ranging from the Lias to tl 

Fig- lofij.— Tooih of CGOUS. 

iZi; 7™™he L^™r The type genus Salidosaurus, of whici 
T^LtL.^IIlf"- is shown in fig. 1061, occurs in the Lo 
of Dorsetshire ; and is well characterise* 
strongly-marked serrations of the teeth, and the short an( 
dermal spines or scutes. In the Wealden, the well-know 
saurus was a Dinosaur of considerable dimensions, chai 
by its enormous and laterally<ompressed dermal spine; 
which the skull and teeth are unfortunately still unknowi 
detached teeth which have been provisionally referred to 



Icnown to be Sauropodous. The anterior dorsal vcrttbnt hire 
'kcd hienul hdgc, which di&appcars in iho5C later in the 

ataitrtdt is founded upon a lower jaw from the WcAlden, 

imay pertain cither \o the last or ihc following genus. In 

wmJkus, of the same bed<i, we have a remark-ible form, in 

the dcmul armour constilutcs a completely solid carapace 

whole of the dorsal aspect of the lumbar region, some of 

nponent scutes being tuherculated, and others ridged ; while 

I was al^o a number of detached flattened spines somewhat like 

of Hylaaaurui, which probably formed a line in ihc dor&nl 

"ITiti ptrcuiiar type of carapace forcibly recalls that of the 

dont Kdcntatcs. In the Cbalk-Mail and Cambridge Oreen- 

&nd Acanlhnfikoiit with a dermal armour somewh;it similar 

of Salidffttuirvi, but with rather more complex teeth. The 

• Ait^lasatirus, of the Cambridge Greensand, was probably a 

allied, if not generically identical, form ; while the equally 

I Sipi^mesayrut and Euareotaurut, of the same depoiiits, are 

by their compressed and carinated dorsal vcriebrse, 

resemble those of Hylitesattrus. Apparently allied to these 

n t-Wtisauruff of the ^V'caldcn of the Isle of Wight, origi- 

'tefencd to the /fcuaitodontida, but showing the "step" on the 

processes of the dorsal vertebra; characteristic of the 

family. PrioJoniogHathus^ founded on a fra^meni of jaw 

■ntnovn age, is prol^bly referable to one of the preceding 

LV STKCoaAumn*. — This family is typically represented by 

(eiuu Siexosaunts ; but before mentioning that form it will be 

ieni to refer 10 two apparently allied Dinosaurs from the 

system of ihc Cape, both of which arc very imperfectly 

and may indicate a family coiiiieciing the present with the 

<mtuite. In Ensct/tsaurMs (more correctly Eusce/i'tfosaurits), 

ftrst of these forms is named, the liml>-bone5 were solid, and 

fcntDT has a large inner trochanter, and approximates somewhat 

duf of the Igiianodonts. The tibia and fibula seem, however, 

hme been more like those of the S/egosourida ; being anchyloscd 

their two extremities, and closely joined to tlie astra^;iius and 

'Ilic caudal vertebra;, belonging either tu this or the 

hm genttt, are of an Iguanodont Type. The genus Orinosaums 

WrrmMna) was (bunded upon a bone of a larger reptile, regarded 

If iia desoriber as the distal end of a femur, but which is really the 

Imiiiial end of a tibia. Although solid throughout, (his bone re* 

MritAes the ubia of ipusnedon in its expanded extremity, and thus 

iVHs a tiactsttMn between the Tguaiiedentidat and Sitgosaurida, 

IW genus SUgoMMms was originally described from the Upper 


;ass reptilia. 

Jurassic of North Americi ; but it a|>pe3r» that certain (oma I 
Oxford and Kimcrid^c Clap of EngUiKl, described at an culi 
under the preoccupied name of Omosauiys, are not entHlol I 
eric distinction. These Dinosaurs agree with the Saiiitta 
the geneml structure of their teeth, and the possession of a i 
aitnouT of scutes and spines, ha vrcW as in their solid Xa. 
but differ by the great height of the neural arches of the 
as well as b/ the circumstance that in the bacrum cacb 
either chielly or entirely itupported hy a ungle centrum, io 
by the adjacent portions of two centra, as in the preceding ! 
The skult ^fig. 1062) shows many points of teseinblince to 
/guanodon, especially in the presence of a predentary booe'J 
is lower and narrower, and thereby approsimate» to the 


ato«>.-Laft Uiinl Hew oT skull of SUftmitru anuH; turn «k« V|ipw 
Aiunoi. One<tiunM namiwl ■!«. *, Nkro: *. Orbii; /, IbA 
" lit; -.NuvliM Pnfivnuli /^.Sufnorblul; A. 

; «, Ancufar ; i, Spknk) ; W, UtnUrj i /</, Pndt 

lB(m««nanl la 
,. PiMi/raaial: 1 

PndtolBiy, <JUtarl 


m \wM 


sAurian type. The Iguanodont resemblance is, howevfcr, so 
as to forbid the reference of the forms wiih dennal annour 
separate sutiordcr. The two rami of the caudal chevronJ 
not unite superiorly. In the pelvis (fig. 1065) the ilium 
enormous preacetabular process, and a very ^lort postacec 
portion ; while the ischium and pubis arc relatively short. In tb 
femur (fi^. 1063) the inner trochanter is either very small D 
absent; the mctataruls arc very short and stout; and the if 
di)|iits of the plantigrade pes approximate in contour to Ibosc C 
tile Elephant. The tibia and fibula arc suturatly united together I 
their extremities ; and the former is completely joined to the asot 
galus, and the latter to the calcaneum. Other peculiar fcatuns 
these remarkable reptiles are to be found in the stmctore of [h 




nal in the sacnim. In the solid structure and gencTa) 
pr the liml>-borKS, as vel] as in the relatione of the 
Cia to ihcir jrchcs, Sieg^sairrui approximalts to the sub- 
Mpoda ; and thtis shows how imposstlilc it i« to bring out 
relationships of aninuils In a linear c1a»:^iBcation. The 
ies indicjics a reptile about two-thirds the dimensions of 
B teririsjar/ensis. The American species have no inner 
r to the femur. The 
]fCM4oH has been ap- 
an American Jurassic 
•cterised by some modi- 

in the sinicture of the 

'fyttrvpMaus, from the ' ^fc^BTl K\.--ts 

Ariiorta, appcare to be 
ku closely allied to the 

._ f 
CsitAToriDA. — This 
been proposed for 
ihle Cretaceoos 
•Hied to the pfeccd- 
|r« but can onljr be pro- 
adopted. The type 

(according to " ^^^^ /\ M ./'' 

atical with his JWy- 
if the I-animic Crcta- 
ir the United States, 

'pair of large hflm-cotes /-d&flr t- 

kull, which are curiously 
•e of ihe Bvvida, and "»■ •°!>:-,'i^» ij" iwo"! •^ p*/* 

mtably Sbcathcd with from itii Lppcr Junme of Nortb AmcritB. 

rhe body was cm«r«l d-r.!". Ti^dUTrri.:-. W^J-*: 
rmal scutes, which not ;-i?;„;i,''rr;J:;';Sn"u,^/.-. w 
biy had overlying homy ii»lu»;t'. Caiamum. (Afirt M«nh.i 

ike those of ChcIoni3n». 

Upper (ireensand of Austria there occur similar horn-htce 
lace thought to have been attached to the body, and 
I 05 S/n//it(uat/nts {Crattramus), and it has yet to be 
(at the American types are Kencrically distinct A gigantic 
rm, from the Laramie, has been christened Triarxitops, 
ihancterised by the presence of an additional nasal hom- 
ppoited by the coalcMrcd prcmaxilla: and an additional 
woe The skull is stated to he upwards of six feet in 
the frontal bonxorcs measuring some ai inches. A 
Hike bone in the British Museum, from the Wealden, 
indicates an allied type. 



jVaJosaurus., of the LaTamie, is a form more or less iw 
to Slegosaurui, characterised by ihe completeness of ihel 
armour, which consists of tows of rounded knobs, b«comiB|i 
and (juadrangutar near the head. The skull rs unVnovm. 

Finally, the remarkslilt^ genus S/enaptlix, from the 
Wcaldcn, lihould {icrhaps find a position somewhere in this I 
bourhood, although it differs from all other membets of ilrit| 
order hy the presence of cavities in the centra of ibe 

Suborder z. Thebopoda-^Hic suborder Theropoda 
some respects a position intermediate between the Omith 

f\tt i<6|. — ~I*hc lift ildc oT Iht pclvli at AUuaur^i /hifiSti liMn lite UfVCt Jb 
Monh Americ*. Otie-twtlfih namral else- i7. Ilinmi f; itMam;^ hbii;^ ' 
(AAer Muth.) 

and the Sauropoda, although its members are more nearly: 
to the latter; with which, o^ already mentioned, it has b«ii 
posed Id group them, under the name of Saurischia. In 
respects, such as the structure of the teeth, the form of the b 
mur, the occasional prcaencc of only two sacral vcrtcbnc. "^ 
the fonn of the qtiadr&tc bone, certain members of this suberiw 



Fig. id6}.— Utcnl 
tipttt oS a UMh 4f 
Myai>uu>nu iP>ct- 
jntirff ( rrom ihn 
SUDMllald Sbi«. 
Out • iUkJ nalural 

The limh-bonea 

appro^im.iiton iHan the Sauropoda to the genenl> 

odilia, although in iheir hollow limb-bones diey agTM 

hanker Omithopoda. All (he forms trcrc cariiivoroui. 

call (fig. 1070) th« ptemaxillic are tur- 

tDogbout with Iccth, which aic laterally 

(fig. 1065) and backK-ardly- curved, 

:faant edges, of which the ]M»tcnor, and 

also the onienor, Ijordcr Ls sermt4Ml ; 

tions of these serratiotti being ^vnciaJly 

ri^t angles to the axiR of the crown. 

seifa XK always implanted in distinct 

and the tIcuU has a Urge preorbital va- 

1070). The cenira of all the veriehne 

intenully ; and their neural arches 

by xygDcplwncs (diplospbenc^) conc- 

10 those of the Squamata ; while the 

. 1067) arc much comprcttcd laterally. 

ire medullary cavities ; and since the pectoral limb \f> much 

than the pelvic, it it probable thai many forms were of 

habits, although .-iomc may have been quadrup«bl. In 

(fi^. 1064) the ilium is of great vertical depll). and lias a 

proves ; while the pubis is dircrtcd downwards 

and unties with its fellow in a lon^f bony symphysis, 

generally extended up the anterior face of the two liones ; 

ibem to have the appearance of an elongated V\ when 

tfaia aspect Both the pubis and ischium are of a coni- 

j short and slerxlei type of structure, and the latter ii>iu.illy 

tutator process like that of the Ornilhopoda. The astra- 

usually clowly applied to the tibia, and not unfrequently 

a long flattened proceiis lying on the anterior face of the 

nne, and thu& resemblet the condition found in young 

Birds before the anchylosis of the two bones has taken 

The metatarsals are elongattnl, and the feet digitigradc. In 

iras the number of the digits varies from four to five ; while 

jies there may be either three or five. And iii all cases their 

phalangeals are furnished with curved claws, which in the 

arc very long and prehcn-silc, and were apparently adapted 

•cizure arvd retention of the living prey. 

y further be observed that while the cervical vertebTW arc 

ly shoner than the dorsals, at in the Omithopoda, yet they 

imea longer, like those of the Saiiropwia ; while the ver- 

prches in the sacrum are occasionally sup]X»rted by single 

[as in the latter. Finally, the femur, which may be cither 

hat lon^ or shorter than the tibia, is generally of a more 

filian type — especially shown in its flattened head — than in 



Fif. toM — Ool«r 
»ma at ibc mmn 
of • WMih of y^tew 

•^ I fnm the Lower 
K^uptr nun UiitioU 

any other Dinosaurs, although tt has a distinct inner ti 
In all cases the dorsal vertebra: arc amphicuclaus. 

Kamii.v AxcHiRAVBiu-*.. — The least specialised (onas 
ing ihis family, of which ihc name must he conftidercd 

are mainly known from beds of Triaswc 

^k are typicitly represented by the Nonh 

^■^ genus Anchisaurui (^AmfiAisatims). ttw 

^H^k i-ertchne axe amphktKlDus; the pubca ore 

^^■B rodlilce bon«ti ; and there are Cnt cligitt 

^^P^H the manus and the pes. The teeth ( 

^^Lj^P usually have cumparaiiTely thicic and «hoTt| 

with a slightly convex posterior boi 

inoie or less oblique serrations, which 

whole or both borders. Th^fodanfataun 

in the Txjwer Kcupcr near Rn'stol ; the 

the tv'pe species being much smaller than 

Rgvied in the woodcut. Aiuhisaurtis (which, according U 

sor Marsh, is identical with a form described at an earlief 

der the preoccupied name Megitdaetyl«t) occurs in the repti 

of the United States ; but no 
has yet been presented to 
it differs from the Euro; 
1'he writer lasl quoted inci 
family the .\mencan Tn 
liaihy^atfiHS and Clefisjfsiuii 
of which are stiU very imj 
known. Here may be meniil 
meruit. Moiiospondyius, from tU 
system of South Africa and lU 
st.ige of the (iundwana system) 
of which a tooth is shown in < 
tp. 1 049), and the ccntnim oli 
vertebra in %. 1067. Thcse| 
approximate to those of Mtgn 
and have a fusiform internal ti 
I-inally, i[ Is highly proba 
an amphiC4£ious cervical ven 
tained from beds of unknowf 

V'pt- i<«T.— Lawral and inliru* &•■ 20tc 

Ct> bf till. f«iiiriini of ■ dona) vvna. 

th* KimaiiiKmaf BkiDioLind. Un*- 
)utt Baiuni tin. 

age in Bathurst Island 

America, and described on 

name of Arctotaurus, indi 

Dinosaur more or less closet 

to the present or following family. This vcrlelna agrees \ 

mateiy in »i«: with that of Caiamos/vndylHS {f\^. io;i^ bull 

cer^-ical ribs and a neural spine ; ihe lenj,tfa and cunratur* 



ang indicative of a long and arched neck. The occur- 
k Dinosaur to far nonh is of extreme interest, its serving to 
t gcnem common lo ihc Old and New \Vorlds may have 
ffrocn the one hemisphere to the other. 
^ MeCALOSAUKiD.t:. — In this Family ihc ccmral vcncbne 
n known) shorter than the dorsals, and may he either 
r opislhoocdous ; while the dorsal-s often have prominent 
^tbe haoit of the arches, hetvreen which are deep pitK ; the 
1 oeniia, as in the last family, liaving a fusiform internal 
fin the pelvic hmb of the type genus, the femiir is of a re- 
f Crocodilian tjpe, and is longer than the tibia. The crowns 
lUh (fig. 1065) ate more or less L1II. and much compressed, 
1 po^erior border distinctly concave, and the serrations, 
kc rtearly at ri^ht angles to the axis of the crown, usually 
,M the lower part of the anterior border. In the Lower 
of England we meet with Palamaurui (with which C/ad^-odon 
Bne deposits may be identical), of which only detached teeth 
m. In the Up|)cr Kcuper of the Coiilincnt there occurs 
paratfv'cly generalised genus Zandodon (liralosaums)^ which 
iprobaldy idcn- 

y Ptaleoiaurvs, A ^-^ B 

case the latter 
ould be adopt- 
c serraiioos on 
nor borders of 
ps of the teeth 
\ nearly to the 
)ft cervical rcr- 
|re amphios- 
Icre arc but two 
tin the sacrum; 
•fttiagalus does 
I off a process 

Ig on to the anterior surface of the tibia. One of the 
|Vaft fully an large a.i Mtgahsaurus, and the gcmis was 
f represented in the Lower l.ia.s of IJorsctshirc. 'the small 
yoden {Ankistrodon\ of the Panchet stage of the Indian 
Dfts, has teeth (fig. 1068) of a Megalosauroid type, in which 
Itions arc totally absent from ihi: anterior border, and do 
ltd to the base of the posterior border. 

^rpe genus Mtgahsanms has been tendered classic tluough 
DRof the late Prnfes-iors Buckland and Phillips, and affords 
lent and instructive instance of the gradiinl restoration of 
letoa of SJI extinct and uncouth iomi from more or le^s 
ricmainSi. In Europe this genus raiigc» from the Stoncs- 

ric kM.— Fncncot of ihc nunillble at F.fltmmtoivm 
{mdStmii (ma the PwKrhtt bdli of ihc Gumlwanu. Thr«« 

TnaivcMe BMtlon o< tt»ih. (Artw Hiuky.) 



field Slate (Lower Jurassic) of England to the topmost Cre 
of Maastricht in Holland ; and has also been reoxded fti 
Upper Cretaceous of Southern India. There is good evid 
the existence of two species in the Wealden. An attemj 
restoration of the skeleton of the type species is shown 
accompanying woodcut The North American Aiiosatina{o 
the pelvis is represented in fig. 1064) appears to be > 
allied form of Upper Jurassic age. The nearly entire sk 
of the latter and other American forms have shown thai 
very excusable errors were made in the restoratioo 
imperfect elements of the skeleton of the English gem 
is quite evident that the skull of Mcgaiasaurus mu 

had an inferior temporal 
and that it approximatf 
or less closely lo th«t ol 
SOUTHS (fig. 1070), althc 
bony processes of tfa 
may have been wanting. 
teeth (fig. 1065) the > 
on the anterior bordc 

Fig. 1069.— Restoration a( the ikelclon or Mr^Maitna BueHamJi; fnui tbc Si 
Slate. Gruily reductd. 

some cases confined to the upper half of the crown, but ii 
extended nearly throughout. Typically there are five vert 
the sacrum j the cer\-ical vertebrae are opisthocoelous ; the a: 
has a process ascending on to the tibia ; and there were thn 
in the pes and probably four in the manus. In the North A 
Creosaurus we have an allied Upper Jurassic genus, in wl 
postcervical vertebrs have very deep depressions on the sides 
centra ; and there are but two sacral vetebne. In Ceral 
of the Upper Jurassic of North America, the skull (fig. i 
remarkable for carrying a single bony protuberance beh 
terminal nares, and a pair of similar protuberances direct); 
vance of the orbits ; while the mandible has a lateral vao 
that of the Crocodilia, Professor Marsh believes that I 
tuberances on the skull supported horns. In the type specii 
three pelvic bones and the metatarsals were respectively an< 



but this may perhaps be due to a patliological peculbnty 

iuSdual. 'lilts genus U regarded by its describer as the 

dtsUDct family — ihe Ceratosaurida, Here may be men- 

nnali and imperfectly known Dinosaur Uon\ the Weiildcn 

tfac name Aritlvsuc/tut tios been proposed. The type 

comprises the sacrum and pari of the pelv-is. It is cun- 

ly its cle*crib«r that the sacrum includes five vcitcbrac, 

pelvic boiKx, which have a ion^j vcntiat Hj-mphyws, were 

t described as pubes, although it has been eubiicqucntty 

B that they may be ischix The dor.sal vertebne referred 

brm by Sir R. Owen ha^'c a fusiform inleinal cavity in Ihe 

but it has been recently liu^c^led that [his reference is 

r/ i 


-t^ Utvnt «i(v cf ikotl of Cir*ietAinti mtukernii : fn/m ihe Vpp<r JnniMitc 
'a. Omdiaih nMonl tin. «. NarMi *. Roiiy iiruminenr* ; r. Prrviblial 
1 1 f. lAA^itatponl (om^i y^ Mudabuku vacuity ; /, Travi'i^'prta IjQufl, (Xricr 

thai this genus may have had dorsal vertebra: like 
the C<t/uride; in which case this form may be lefcrahle 
ntf, in which, indeed, it has been placed by Professor 

,T CompsognathidjK. — This family is represented by the 

wmfje!p*afAM of Ihe Lower Kimeridgian Limestone of 

laod the allied or identical Haliopui of the Upper Jurassic 

R America. In the typical genus — known only by a t^inglc 

cervical vcrtcbise are opistlioccelous, and much longer 

amphiooclous dorsals, and have free cen-ical ribfi. In the 

)« shorter than the tibia, and both the tiianus and 


pes have only three functional digits ; the astragalus being d| 
applied to the tibia. The teeth and pelvis approximate l«j 
Megalosaurian type. Compsognathus was about two feet n i 
length ; and undoubtedly hopped on its pelvic limbs, ata^ 
manner of a bird. Hallopui is one of the few Dinosaun wilkl 
two sacral vertebrae ; the number of those of Comps^giiaiiaa M 
unknown. '< 

Family C<ELURiDiE. — This family is represented by companll 
small forms, ranging in size from five or six to about twehel 
in length. They are characterised by the vertebne and ribt t| 
completely honeycombed by cavities, communicating with a i 
aperture on the side of the centrum. The cervical vertd»«J 
1071) are typically longer than the dorsals, and have thei 
anchylosed to the arch and centrum, and the neural spine ledl 
to a mere ridge. The limb-bones are hollow, and the pdriii 

Fig. 1071.— Anlertor and [eft lateral atpKii of ■ cervical vencbnof fa Immmftmijlm 
fram the Wealden of the Isle of Wight. Two-thinl> iiatml liie. frt, ' P n rjtf fafkfm 
Po9tiy);apoph]nis; r, Rib (resiored) ;/, Pticumaiic (bmnen; i, Neuiml qnne. 

the ordinary Theropodous type, with a very long pubic sympt 
The type genus Calurus, typically from the Upper Juiassi 
America, but also occurring in the English Wealden, has 
cervical vertebrje greatly elongated, the first few being opi 
and the remainder amphictetous. In Calamospondylus (fig. i 
the cervicais were shorter, and were probably all opisthond 
the one known species being from the English Wealden. 
fessor Cope includes in this family other small Dinosaurs 
the Trias of North America, which he identifies with Tiii) 
pfueus, originally described on the evidence of extremely elon( 
caudal vertebra from the German Muschelkalk, once refern 
the Sauropter>-gia, In these forms all the cervical ^rtebt* 
amphiccelous ; and the femur has an inner trochanter. 1 
dacty/us, which Professor Cope regards as identical with 
forms, is, as already mentioned, identified by Professor Marsh 

Suborder 3. Sauropoda. — AVith the third and last subord 




TJ-— IhitAr i»i<l* of 
rnaiiiil (turn ibc U|vsr J<^ 
Oorloll'nBtunl tkat. (Allii Marth.) 

Jurawit of tfcnih Aiiiulia. 

the conakleratioi) of Lhe Urs^t known Dinosaurs, and 
make so many marked approxiiimtiunii lo the more 
CrocodiUa as to &hiivr how close is the relationship 
I order and the Dinosauria.* The skull (ftg. 1076) 
thai of the Thcropoda in having the prcmaxillae com- 
Dthixl, and also in tht ijrcscncc of a large preorhilal 
»ut the external narcs formed long slits in the fore part of 
Bs in Omithosaurs and Bird!i. 1'he teeth are invariably 
inct sockets and are of a spatulatc form, n-itliout marginal 
<fig. 1072). The sternal hones are ovate. All the vcr- 
kdvance of the sacrum, and sometimes those of that region 

a large racuity 

»de of the cen- 

■nunkating with 

iotemal cavities, 

|re a hontrycomb- 

to the whole 

This affords a 

ion of strength 

in the massive supports necessarily required for the 

limbs ^^^ muscles, which could not have lieen attained 

manner. All the anterior vertebrae are opisthocwlous ; 

cen-ical region the lihs are anchylosed 10 the vertebra;, 

kn no neural spines, and arc longer than Lhe dorsals ; while 

fi» of the latter are laterally expanded : and in the sacium 

supports its own arch, The limb bones are solid ; 

the pectoral limb is not much shorter than the pelvic, it 

1 these Reptiles were liabitually quadrupedal. All the 

plant^rade, and furnislied with live digits ; those of the 

terminated hy large curved claws. The ilium (fig. 1073) 

border only moderaicly arched and its postaceiabubr 

; while the piibi? (fig. 1073}, which is directed down- 

fonwards, is stout with n comparatively small distal expaii- 

it unites by a oartila^iitiious t;ymphy8is with its fellow of 

itc side The ischium (fig. 1073) is likewise a stout bone 

with that of the Crocodilia in the absence of an obturator 

and the two ischia have a peculiar incurv-ing of thcii distal 

where they meet in a symphysis. 

be observed that in the lateral views of ihc pelvis shown in the 
befc the bones arc drawn more or less nearly in a vertical 
impossible to give a true id» of the peirulinr contour of the 
mily of the pubii and ischium. The-»e hones arc really con- 
above downwards on the outer (figured) aspect, and convex on 

\ Cop« wottUl include ths S«ui«|wdous UiiwMuirt in tlu Ciocodilia. 


the inner upect : and owin^ to a cnnfuiion bciween the 
and outer sides of itie Kn);li»h »ipeciincnii. only recently de 
lias been considered by some pabeontologists iltai then was la t 
Oiffttpticc between the structure of the pelvis of the English aodj 
■can forms. 

The femur in its straight shaft and absence of an inneri 
likewise resembles to a considerable extent that of the 

.* y 



_ftt- ia}3'— Tho Iril >rde of tht ptlvii of iJntaMMurM titi/nu: fnta i^ L'pM J* 
/, KoniRicn in da (Alicr AUnh.) 

although its head is not laterally compressed to the samei 
placed aa obliquely to the distal condyles. 

In titne this suborder ranges from the Upper Trim to tbe < 
Uceoii-t, and it is especially well represented in the Kijueridge 
and VVealden of Europe. 


Sf Atuuttosau urn*. — This nime may be provisionally 

for a family which is represented by tlic genera Aiianta- 

^fiatosanrus, and Brontotaurus^ of ihe Upper Jurassic of 

icrica, as wdl as by allied European types. In these liugc 

isdttutn (fig. 1073) IS directed dou'nward<>, and il!> shaft ii> 

not bcni upon itself; while the humerus is comparatively 

and ihe chevrons of the «iudal vcrtebric have their superior 

united, 'j'hc teeth have the summit of the crown not 

■ka IJtvcJiMaMc of Nartta Amnio. Rcdund. (AficcCopc 

Apparently closely allied to, if not identical with, 
the abore-mentioned genera arc Amphicaelias and Camara- 
]"of the same formations. An anterior doriiaC vertebra of 
|er genui; is represented in the accom|ianying n-oodciit ; this 
ii b transitional between the cervicalh and later dorsals, the 
■pine of the latter bein^ absent. 

bu>r Manhestinuiet the total length r>i Bronlostturus M. upwards 
;fc«i, and Jii weight at more than twenty tons ; and Peiofosaun/s, 
En^'liih Wealden, must have been fully e<]ual to these dimensions. 

Kn ai the former, the learned American palwontologist obsen-cs, 
« animal at times assumed a more ervci pnniiion than is ri^prc- 
I in the restoration is probable, but locomotion on the posterior 



limbs aJofic was hardljr possible. The head was rcmarlutblv trnJi 
neck vfta long, nnd, considering its proportions flexible : ^m 
lightest portion of the vcrlcbml column. The bod)- wm quite 
tlic abdominal cavity of inodenile si«, . . . Each fooLpritil i 
been about u iti^uare yurd in extent. The tail wns lar^c, .ind 
the bones wcic bolid. The diminutive head will fir^t attnci *iv 
it ifr smaller in proportion to the body than lu luiy vertebrate 
known. The entire skull is less in diBRictcr or actual weight i 
fourth or fifth cervical \-vrtcbnt. . . . The vco' snudl hcAdM 
nnd alcndcr ncuml cord, indic:iti! a >tupid. s]ovr-inD%-iitg rcpnllj| 
beast was wholly without otTcn^ivc or defensive weapoos, cr i^ 
armature. In habiu. lironiosaums was more or less amphibious, i 
food was probably aquatic plants or other succulent veKCtatioo.^ 
remains arc usually found in localities where the animals liad i 
become mired." 

Of still more stu[>endous bulk is AtiaHtotaurus immuuty tlwl 
of which has the enormous length of six feet tvo inches, »^ 

rrvmtbcWuldaiofthcltleafWicbi. (Aha Wrttbl.) 

indicates one of the largest land animals yet known; the only Tot 
which could possibly have exceeded it being the Cretaceous 7>m 
aiops mcnlioiied above. It is by no means clear that all tlM 


' tf peii are genericolly disUnct Trom those lo be now 

now proceed to the coriMderalion of certain Eitro|x>an 

vfaich boai ihetr tnorc or less close alliance to the )>rc- 

appear to liclong lo the same ramily, although opinions 

have \xxn expressed. 1 1 may be observed in this 

that the study of oil the European S.-iiir(i|K>da is t)eset 

>t iruuimountable ditSculties owing to the drcuinstunce 

By all the !>pccimcn!> urc di»a»sociated, and thai genera and 

»ve been named on the evidence of angle teeth, vertcbrsc, 

of the limlM or limb-girdles, so as not to admit of cum- 

rith one another. Moreover, (he unwieldy bulk of the 

t thenudvcs is a bir to an exact comparison, even when 

Itrictly comjurable one nnth another. 

Doth from the Wealdcn of the Isle of \V'ight, represented 

ptural size in fig. 1073, has t>ccn made the type of the 

i^irMW/Kj {Ofilasaunis), itrtd it appears almost certain that 

jmd dorsal vertebra, and a pelvis from the ttame deposits, 

S under the nanw^ of Omifhfpm ffulkei and O. ewtimrrotiii, 

■Me to the iiame form. These remains indicite a Reptile 

Bcrmbly smaller dimensions than Braalostiiirus, havint; a 

i»ch approximates in structure to that of AtUintmaums. 

ram, which has the downKurd direction characteristic of 

nt fxmily, has a length of 27 inches, and is com))arntively 

raportion to the pubin. The genus PtlonnaurHi i» typified 

! hoiDenis from the ^V'ealden of SuKsex, meosiiring some 

\ IB length, which would appear far too large for the type 

f ffirfioiayrHiy A slightly L-uger humerus from the Kim- 

hjr, origiiuJIy described as Ctthtaunu humcrtKristatus, 

genericnlly inseparable from Ptioroiaurtts, and its owner 

II probabiUly very closely allied to an equally large form 

Oxford Clay, described upon llie evidence of tlie pelvis as 

us Letdii. Of the latter the lumbar and caudal vertebra; 

movn, and approxinute closely, both in size and contour, 

Oi Jir<>ntesaurtt!, the lumbars having a diameter of nearly 

E across the centrum. In the pelvis the ischium mcxsuroi 

I inches in length, and is also narrower in proportion to the 

Kn in Hopt&taurui — differences which, coupled with others, 

ubly be reganled ^is, of generic value. Referring all the^e 

least provisionally, to Ptittrmaurus, it would appear that this 

dudes TCiy large Dimotaurs closely allied both in vertehml 

c diaiaciers, as well as in point of size, 10 BroHliiiaurus.^ al- 

m lli« profMrtiooi of ihU lionc and of the tMliiuin iDDnl)one<l Lclowr 
DfTopoAdb^ dinMuioai of other Dlnouun giTCD in ihc ubie on the 



:ho^^h it is probable that the humenis was reladTcIy longer, 
from :he KL-neiid^un of BoMlogne. upon the evidence of i 
Zcr.cs -W/a^w '»ai foimded. bur which were subsequeniljr id 
wiih .ViE'eHcan forms de^ribed as Camlffdan, are probablf t 
to *he 5iine OTcies. Small rertebne from the Kimcndge I 
Swi^ndon. upon which Ihe genus Botkriespondylus Taa 
mav perhap* be referable to a ver\- young indi^'idual of Ftk 
humir-^^!:tat4i ; while a coinpara.Qvdr small humerus 
same depos-Ls. orijinallj- described under the preoccupied : 
lichyrciiMurus. may indicate a smaller species of the same 
Finally, of rhe Kimendgian remains described a& Gigat 
while <om.e may hi referable to P. hxmerocrista/us, & sacral 
may belony to the smaller /*. Afamseli above mentioned. 

Thcfi'lo-iing table ;jivei thedimensionsof someof thebonHofc 
of the a'a.jve-mentioned ioTTn«, tojfe the r with thoseofothen noticed t 
under !he head of ".he CH:\ijjurUiP : — 

^ w It ti C 

~ « "- "• ■ ' 

Lenjlii of *.apuU . 


' 6a 

II Huincni! . 


i< ischium . 



II feitiur 




Widih of doisal ctntnim . 


M lumlur II 


1' caudal n 


54 57 
■■■ 3S-5 


Family Diplodocid.e. — The genus Difilodoats, from the Uppt 
Jurassic of North America, is typically represented by a species ( 
smaller size than many of the preceding forms, and is regarded b 
I'rofessor Marsh as the type of a family. It was originally cw 
sidorcd that the external nares were single, and situated at (he W 
of the skull between the orbits (fig. 1076) ; but it now appears thi 
they really formed long narrow slits between the premaxillx, nasal 

' Caudal vertcbite of a lai^er individual are nearly equal in sue to thoK 


the fanhion tibwining in PieTodact>'Ics and IJirds. 
prolnl>t« that a biniiUr arrangcmcni exists in the other 
of ihe suborder. It will be obscn-cd from the figure 
ti^ail bifurcates posteriori}' to form the anterior and inferior 
of the oriHt; while the quadraiojugal joins the maxilla 
be intervention of the jugal, which is tliruRt up. These 
features arc repeated, is will be noticed below, in one 
the Omitbo&aurii. The pelvis is of the general type of 

Om «ulh own! tin. Tha poMtion of ih« nam ai Ihe ion oT int ikuU If 

he last biDiljr; but the diiital extremity of the ischium is 

r CiCTrosAURiDie. — This family b t)-ptcal1y represented by 
sh genus Cetiosattrut, whieh, so far as can be determined 
chancten of ihc scapula and pelvis, appears lo lie so 
ated to the American Marotaurus that there seems every 
If including the latter in the same family. Cctiosauryt 
pically in the Lower Jurassic Great Oolite, and Forest- 
( Oifordshire and Northaniptonahire, where we meet with 
C. ox^muHtii. Comparati%vly small teeth from Ihe same 
desailred under the earlier name of Cardi&d^n ruguUau, 
■ same general type as those of Hophiaumt, but have rela- 
zller crowns, with a more incuncd summit, and are clearly 
[rom the last-named genus. Professor I'liitlips referred 
this type to C. oxomcnsis, but from their small size they 

1 1/8 


would agree better with a dorsal vertebra from the same 
subsequently described as Botkriospottdylus ro6us/us. This 
is somewhat longer than the dorsals of C. exom'ensis, bat 
be generically distinct In Cetiosaurus the caudal vertebne 
no distinct postzygapophyses, and articulate by two fimti wA. 
chevrons, of which the upper extremities are not tmited, ai in 
codiles. The scapula is much expanded at its distal extremi^, 
the humerus, though long, is wider and shorter than in the ~ 
pean forms mentioned among the Atlantosaurida. The 
appears to be of the same general type as that of Moroumna 

Y\a. 1077.— The left udc oT ihc pelvis of Maresmmna grtmMf, from the Uppcr^Jnoi 
North America- One-sixteenth iiAtund tue> d. Acetabulum; i7. Ilium; it, Isdinim; 
Pubis. (Aftei Marsh.) 

1077), in which the shaft is bent backwards and has no di 
expansion, while the symphysis does not extend to the extrem 
thus causing the middle of the acetabular part to be far above 
axis of the shaft. 

The typical species of Cetiosaurus is comparatively well known thro 
the labours of the late Professor Phillips. This huge reptile was peril 
somewhat smaller than Peiorosaurus ; and it was inferred from the » 
ossified extremities of the limb-bones, the free projection of the heai 
the femur into the acetabulum, and the lar^e terminal claws, (hat < 
creature was of terrestrial or subaquatic habits, and that it proha 
dwelt on the banks of lakes or rivers among brakes of (ems, cycads, i 
conifers. From the structure of a tooth {Car^odon) found in the si 



[Cctiosaur was inferred lo have been of herbivorous habits 
irkibly these coDclusions bat c been coiilirmcd by the discovery 
td Ainchcaa forms, is now a maitcr of history. 

iWejUden sacra! and caud:il verteVjrK of the general tyjie 
lOf CtHcsawus, dcMTilxrd under the nnmc of Celiosanrus 
OtCBie a smaller Torm thnn Hitphsaums. ll is prol^Kible 
tssl renebra from the same dt-posits, dcscriUi-d as Bothrio- 
\fifft^ants, as veil as a bumenis and otbn bones of the 
linttK to which the name MoroMUrus Btckltsi hs& been 
trc referable to the same form, which would appear to he 
ly disUiKt from Cilioiaurus, and n>ay l)c known as Moro- 
tvit. The dimenitions of sonic of the bones arc ^ivcn in 
I on p. 1176. An ilium, from the same beds, clos^^ly 
I that of the American j^enus, and probably belongs to 
t form. In North America, Mcresaurvi and the small 
fctr are diaiactcristic of the Upper Jurassic. The dorsal 

of ihe latter arc rtlatively elongated, and do not exceed 
ic* in length ; those of one sjjccics being remarkable for 
fy low neural arches. Small teeth 
Wealden (fig. 1038), once referred to 
ma, probably belong to a species of 
^. i'hese teeth are less spoon-iiiaped, 
poaeimate more to a compressed cone 
ic of Morosaums (Rg. 1073); and the 
kuroaxitts includes the smallest repre- 
B of the suborder. The pelvis of 
PBLT is shown in fig. 1077; the toeth 
(a)i although considerably larger than 
f Cardiodon and PUurocalus, exhibit 
t incurving of the crown, 
e preceding forms the centra of the 
enebne are amphiccelous ; but Titancsaunts, originally de- 
iFrom ilic Cretaceous of India, and subst-quenily found in 
bsh Wealden and Upper Oreensand, has precocious centra 
i-ertcbtse. The femur of this genus indicatL's an animal as 

tins of oilier Cretaceous Sauropoda have received distinct 
Dames, but some of them may be identical with the aWvo- 
cd types. Thus we have /?/««ibi-«j based on Woken bones 
J Lower Oreensand of Kent ; and ^pyutuna founded on a 
I from the French Cretaceous. MacrHretauruf, From the 
Ige Greensand, is a smaller form, with imperfectly procudous 
vertcbrx. I-'inally, the name Thecospondylui has been ap- 
' a specimen from the Wcilden, of which e%'en the subordinal 

cmiiot be determined, although it has been auggotcd, 

\*t. iotS. — Oittcr and 

frofili i-itwi cFn inoih af 

tram Iha Wnlilcn. 



without sufficient foundation, that it indicates a fora 

Order IX. Crooodilia. — The Crocodiles of the 
arc well-Vnown, bccrtiform Reptilcii tnhabitin}( the I 
and marshes of the warmei regions of the globe; and] 
largest exiting representatives of the entire claw. U we] 
these existing and specialis>ed forms to deal with, we 
no difficulty in i^ivtng a concise dcfmition which wxiuld 
order to which they belong from the Oinosauria, 
howcfcr, in the I'rias a number of generalised forms 
proximate so elusi-ly to ibt; latter order as lo render such i 
extremely dilficult; and it is quite posiblc that same of 
mentioned characters are not applicable lu the first sul 
order is sometimes known as the Emydosauna. 

In b]I the forms the limits and body (fig. 1079) are of 
form type, Uic former bcin^ vcrj- short, and the latter 


Fit- lOM.— Hml and roTfpan af iti* body(a)iitdhIn<l(iiMt*)('rrwaA.'hi/Mi 
fraoUuliu. Mitch mduc*!. {AlWr CAmlw.) 

carried close to the ground ; while the tail is relatively long, 
the exception of a few later JurasMc forms, the dorsal aspect \ 
tx)dy carries a dermal iiimour of articulating ur imbricating 
scutes, arranged in two or more longitudinal rows (fig. 1079 
marked on iheir outer surface by a scries of deep pits. In ( 
cases there may also be an armour of similar type developed 
ventral surface of the body. The centra of the vencbrae atr 
amphi- or proccelous, and the neuro^rentral suture is p«n 


TOtcbne have double pedunculate cosul tubercles, 
one on tlie centrum and [he other on the arch ; and the 
nbs ha\c long proee^io projeeting anteriorly and pos- 
wbich completrfy prevent the liead from being turned 
iy&. In the dor&aJ region usually the four anterior vertebrse 
tmuvene process for the articulation of the tubercle of 
placed on the arch, while there is & lower process, or rib- 
, on the centrum for the capitulum of the rib ; but posteriori)' 
-Cicet rises on to the arch, and in the midillu cIor}>nlt> forms 
kiod of *' step " on the transverse process, while still more pos- 
ii«tly it tnctgcs with the lubcreular facet, 'Ihe dorsal ribs have 
lie processes, lilte those of Birds ; and ihc chc\Ton-l)onea of 
region usually have the upper limbs of the V not united 


_ lUk btml airf 

I of nbkh H* ■)■ 4rtiB cUBBHaHatJDji pvutr 

arf Hipiriin vicv ottiit liiiW c\t Crtxntti/ui falmfrfi ; IniKa, 
1 a|wniin« lo Ihi rifht art ihc tu[jriieni[ornl lauB. in ul- 
iBDHatinji puutr lolly uriiii ilic •u(r*icui(ivi<l fuuc 

_' btme- Nonnallj" the sacrum has but two vertebrae.^ The skull 
(Ig. to8o) is rebtiveljr large in proponion to the body, and is 
Mvally much depressecl ; its eompontrnt bones are firmly united, 
M>d generally have a chaiarterislic sculpture on their externa] 
tefacei The palatines and pterygoids unite in the middle line, 
ibd thus cknc the palate : and very frequently one or both of these 
bones develop inferior plates, which meet beneath the narial 
(fig. 1089). The quadrate is lightly wedged in among 
■ adjacent borws ; the tympanic cAvitie!> u»>ually communicate with 
mouth by thrcx eustachian canals j the mandibular symphysis 
by suture; and tliere arc generally no o^ificatiuns in the 
derocic of the cycbalL There is almost invariably a lateral vacuity 
>e mandil>le (fig. 1093). The teeth are always either pointed 

* Ai an atinoinialiiy ihiM ucraU inny be prcMnL 



Fig. loSi.— Left sideofihe pelvis of Bvoung 
.Alligmtor. //.Ilium: /ijlacbium; /", Pubis; 
a. ^. i, Accubulum, wiih iu mcutfy: F, 
Obiumor notch : t, *, Cutila(iiKNU pro- 
cc^»0' of die ischiam And DimiL 

and subconical, or laterally compressed. In the sternal n 
sternum itself is canilaginous ; and there is a bony intercla^ 

generally no clavicle, 
pelvis (fig. 1081) the 
^OTt and deep, withoui 
preacetabular or pubic p 
the ischium is stout, aiu 
of obturator process ; 1 
pubis* is directed do 
and forwards, and is fr 
excluded from the acel 
In r^aid to the pector 
and limbs, it will suffio 
that all the bones an 
that the coracoid has 
nelle, and may be eith 
or long ; that in the 
and femur the heads 
perfectly differentiated, 
the latter being OHnpre! 
placed verj' obliquely to the plane of the condyles ; while 
of the femur has no inner trochanter. Moreover, the tibi 
cnemial crest at its proximal extremity. The habits of 
members of the order are quadrupedal ; and the feet (Jij 
are plantigrade, those of the hind limbs being partially web 
The existing Crocodiles present many peculiarities in 1 
the soft parts ; but since we do not know whether the same 
occurred in the generalised fossil forms, and cannot compi 
with extinct orders, it is unnecessary to allude to them furtl 
This order is peculiarly interesting, not only as contai 
on Y existing members of the Archosaurian branch ; but a 
it affords a beautiful example of the gradual e\-o1ution of sf 
characters as we ascend in the geological scale 

Suborder i. AEtosauria. — This provisional suborder 
hut a single family, which Dr Baur places in the Crocodilia, 
Professor Cope regards it as more nearly related to the I 
cephalia, to which it perhaps belongs. 

Family AgTO.sAURiD.«. — This family is t)-pically represi 
the genus Aeiosaurus, of the Upper Trias of Wiirtemberg 
form with Crocodilian armour and limbs, but with the mi 
much elongated, and approaching in many points of its org; 
to the Theropodous Dinosauria, to which Professor Marsl 

' It has been su^estcd that Ihe bone termed 'pubis id the Eusuchia 



jf rdaied. TSfpotherax, with pitted scutes adherent to the 
I allied form f^rotn the reputed Triaft of North .Vmcrica ; 
tfe^AOf Co|>c rcj^rds xs forcshndowinj; in its dermal skeleton 
Bcc of thu Chelonia. 

PKR 2. pARAsrcHi*. — ^Thiiextremely generalised suborder 
Bd to the Trias, or kitdU of approximntely eqiiiv-alem age. 
Bctcmcd by the absence of descending palatal pLites de- 
rom the roof of the mouth, so that the posterior nnres (ti^. 
CO directl)' into the latter, without the intervention of a 
rjiaaBt^. The vomers are seen on the palate; the middle 
w thrvc eustachian canals appearii to he wanting ^ ; the 
nans are placed in the middle of the cranium ; and the 
Bc have typically sonic iwcmy-onc teeth, and arc produced 
3g rostrum. A clavicle was prohahly present, the coracoid 
: and rounded like that of the Dinoaauria ; while the pubis, 
SaxuSy takes a share in the formation of the acetabulum j 
I foot was probably furnished with five digits. The centra 
enebtx are amphicwluus : :ind the dorsal scutes have a 
ridge snd form only two longitudinal rows ; while those of 
al buckler (when present) are arranged in not more than 
HKh rows, and each scute consists of a single bone. This 
Scis »-cr)- widely from the true Crocodiles, and Dr Baur 
cars 10 rvgard it as a distinct order, under the name of 

tv pHii"TCKAtiinD.«. — This family is best known by the type 
ftoMums {Btlod«M) : originally described rrom the Keupcr, 


-KctKkMnl v{«w«f iam •kail or n^tninf^ ijll-^-TtfJ*^ ; (mm ih( KeurcroT 
Math nitnnid, Ittt vwniilio In lb> ciBiiiiim *n Ibc prewbilBl, Eht orbtl, and 
■K fAnaHcys.) 

Trias of Wurtcmbcrg, but subsequently found in beds of 

aldy equivalent age in both India and North .America. 

itull (figs. 1083, 1083) the orbit is separated by a bony bar 

inlraiemporal fossa ; there is 1 large prcorbilal vacuity, 

Accofding to Dt Kuken. 



and the supnuemponil fos&a is exceedingly smalL The : 
reached the premaxillx and completely suiroiuided the 
pre^nting a feature unknoim in any other vertebrate^ 
0rbii5 were soraewhat irrtgular in contour, and directed 
laterally, and in pan frontally. The teeth are shaip and 

Fij;. loSij.— Fniniil ■nd paLaial upccu of ihe cnniinn of FkylttaMnti cjHm 

ahicnor v^icuiiirft an Ihc upper ^i^urr ajv fbr Bntcrior rurci, «f^ ibc ijiis in ibc- lo«f I 
arc ihr po^ifrior lurcih (After Meyer.) 

with serrated an tero- posterior ridges (carins) ; and in the 
part of ihe jaw (fig. 1084) are subcircular in transverse 
i)ut po>tortorly are laterally compressed. There was no ten 
armuur. In the apparently nearly allied StagonoUpis, of the I'pfd 

Trias (Keuper) of Elgin&hire, 
was, however, a ■well-developed 
dermal armour; the teeth were 
and swollen ; and the pattern of 
sculpture on the dorsal scutes 
different. This genus was 
founded upon the evidence of th<t 
scutes, which were thought to have l;elonged to a Ganoid Fisl 
The name Episcoposaurus has been applied to a North Amaka 
Triassic form which is regarded by Professor Cope as allied « 

Family Parasuchip^ic. — The single genus Parasudius occm 
in the same lower Mcsozoic horizon (Maleri beds) in India whic 

Mb- 10S4.— Tcwch of /'Aj/rjaiifj(j 
cttnlinsrvit ', from the Trijks of 

-inauic LTocoauia. incK wtrc, indeed, origtnall] 

two suborders, but subsequent researcheti have shown 

arc K> closely connected »s to render such dMsion inad- 

In all ihe&e forms the premaxill:e, inaxillie, and palatines 

mrcnor pnbtal plates meeting in the midJIe line beiiciich 

at (Kusage, and thus complctelj'' separating the latter from 

tfa, and causing the farmation of secondary posterior niircs, 

come instances are situated immediately lichind the paU- 

il in others (as in the figure of Crwedi/ut given on page 

Nring to the dcvelojxnent of similar plates hy the pterygoids, 

he bttcf bones. The object of this peculiar arrangememt 

Ide these animab lo drown their prey hy holding it in their 

Miths under water, which is thus entiiely prevented from 

the air pasuges. A gradual evolution of this structural 

be traced from the Ust suborder, where it is entirely 

to the gcneraliiied, and thence to the most specialised, 

of this division. Other characteristic features arc found 

terminally -situated, and usually undivided, naics ; in the 

earatncc of the vomers on the palate ; in the bony middle 

an canal ; and the presence of not more than four or 

b in the prenaxillae. 'f'hcre is no clavicle; the coracoid 

\ elongated ; the pubis is entirely excluded from the nce- 

(fig. loSt); and there are five digits in the mnniis .nnd four 

DCS {fig. I079). 'I*his suborder may be dividtit iiuo two 

Ecordtng to the development or non-develojiment of palatal 

n ibe pterygoids, and t)tc fonn of the vcrtcbm. 

ccurrcncc m all the groups of the Etisuchi;! of long-jawed and 
red forms is so suggestive of ihc direct on^in of the c>i&iing 
. Creon long-jau-ed Mcsozoiciypcs, andof lheCr<KO<lili-stird Alli- 

Elbon-javtcd form* of the iamc epoch, thai I)r Kokcn iidopis 


portion, but may be single ; while in the transverse rowi a 
buckler the scutes always imbricate anteriorly, but in the pa 
part usually articulate by suture ; each scute being invaiiablf 
posed of a single piece of bone. 

It may be incidentally mentioned here that while in the M 
region of the ventral buckler of all Crocodiles the componeoti 
of each transverse row articulate together by suture with dn 
either side, yet, as will be gathered from the foregoing dma 
in the posterior portion of the same buckler in the present 
the articulation of the different transverse rows with one ■ 
may be either by suture, or by imbricating like the tiles on 1 1 

The present series ranges in time from the Lias to the 1 
and Middle Cretaceous, and is especially characteristic ( 
European strata. 

Family TeleosauridjE. — The members of this family are i 
distinguished from the more specialised forms by the arcnm 
that the supratemporal fossae are always much superior in i 
the orbits, and that the latter are completely separated by i 
bar from the infratemporal fossse ; both these features bein 
shown in the accompanying figure of the cranium of Stenm 
In front of the orbit there is always a well-marked vacuil 

Fig. 1085.— Upper view of Lhe crsnium of Sinu«tatinu ilthtrii; fnm the OxfciK 
France- Much reduced. The bones on Lhe neht side of lhe ratniiD uv inperfKC, 
should have been a line connecting the apex of the frontMli viih the soture divtdiDC tbi 
The larEC vacuities behind the orbi« are lhe wpntempon] fouK, below vhich m 
temporal imsx. 

shown in the figure) ; the dorsal scutes, when present, are ro 
and arranged in two longitudinal rows ; while the ventral bw 
divided, and the component scutes of the posterior transven 
are united by suture. The axis vertebra carries two facets 
rib, as in Dinosaurs. The members of this family were of 
habits, and range throughout the Lias and Jurassic sysl 
Europe. They may be divided into two subfamilies. 

In the subfamily Teleosaurina the skull is generally produc 
a long slender rostrum, like that of the existing Gharial ; th< 



a long interval from ihc premaxillae ; the orbits are 
mtour. and directed more or k&s completely frontally ; 
look more or less anieriurly. Tlie dermal armour 
lOped, and sclerotic plates were not present in the eye. 
us Tr/r^iairus comprists small or medium-sized species, 
,Iy characterised by the Iccth being inclined horizontally 
Knd cxtrciuciy numcroiisi. It is ronfincd to the Ijowcr 
Od is abundant in the Stoncsfietd slate of Oxfordithire, 
Bttly equivalent beds of Caen, in Normandy. The mosl 
genus Is, however, StftKorauna {lij{. 1085), in which 
trut may be included, cliaracierised by the elongated 
nearly vertical direction of the teeth, and the large size 
pntcmporal foK»c, which in some species attain enormous 
In the Liassic fonns, i^pstratcd ^fnerically by Eome 
Myitriosaurus, the orbits are somewhat uhliijuc, and the 
iofsx arc nevtr excessively large; but in the numer- 
of the l-ower and Middle (Oxford Clay) Jurassic the 
of the orhitK is entirely frontal, and the supratemporal 
vcfy \as%e. Ill the hjjured A. Hebtrti, of the Lower part 
ord Clay, the skull is somewhat iniennedtate in these 
the orbits being slightly oblique, and the supntlcmiwral 
This genus does not appear to have survived above 
Day. In PtUgosnurtti (fig. 10S6) we have an 


\. — It^^ taMr»l view of iIm •knll af Pfimftiuir*i O'/w -' ^r^n the L'ppcr Uju tt 
Tim— Mil KadDMd. T, Sopnumporal fvN ; O, OrlnL 

.US, represented only by two species of Uassic age ; it is 

(third from the precedirtg by several characters ; but more 

■y the font! of the posterior nares, and the smaller and 

supratcmporal fos&jc. The remains of tlic small P. typus 

nally abtindant in the Upper white Li;is of Nomtandy, and 

rcllously perfect preservation of iome of the skeletons bu 

the Iwny anatomy of this sixrcics lo bt: as Lompletely 

as in the case of an cxislinj^ form. In M<i<himoiaurus, of 

lerklge Clay (Upper Jurassic) of both England and the 

at, and TikiJoMurus, of the Kullcrs' l-:;arth (Lower Jurassic) 

andy, we have two genera in which the sicull becomes much 

broader, the (eelh stouter and less numerous, and the 

oblique; and which thus connect the present with the 


next subfamily. MaehimosaunUy which occurs both in Fn 
England, is the largest member of the order, the length 
mandible exceeding 50 inches ; the skuU has been 
Pliosaurus. The teeth closely resemble those of 
having conical and deeply fluted crowns. 

With the second subfamily, or Mttriorhynefuna, we 
very remarkable group of Crocodiles, presenting certain , 
features unknown elsewhere in the entire order. The 
10S7) is either of moderate length or comparatively ihortl 
nasals are either in contact with the premaxillie, or sepoiated 
from by a short interval ; the nares are directed frontally ; 
are of very irregular contour, and placed completely cm the 1 
the skull ; and the teeth, which are never very numefoo^l 
always of considerable size, and directed more w lea 

Fig. 1087.— The r^imoaaol Miirierhntktithattijtri Anm the Klmertdsc Cbyef Xa 

One-Kixlh natural size, /"iJ^. PremaxiUa ; mj-, MuiilLa ; j^ Nual ; «, m^f—mul ;^, fn 

tr, Orliii. (Afiet Ueslongchampt.) 

vertically, while there is no vacuity in the mandible. Toit 
remarkable features of the group are, however, the developn 
of a ring of bony plates in the sclerotic of the eye, and : 
general or universal absence of a dermal armour. It is, indfi 
very curious to notice the correlation of these two features, fl 
there is no known instance of the presence of both sclerotic pll 
and of dermal scutes in any reptile. The pelvis of this subfiuailj 
also worthy of notice. Thus the ilia are very small subtriangi 
bones articulating with long and downwardly curved sacral iM 
while the ischia are enormously large, with the shape of an isosodl 
triangle. This presents a remarkable contrast to Steruasamnu, «liB 
the sacral ribs are straight and directed outwards, while the iliSB' 
larger, with a considerable portion projecting above the costal irti 
ulation. In the genus Mtiriorhynckus the skull (fig. 1087) ti 1 
moderate length, and frequently somewhat slender, with the frod 
region slightly sculptured ; there is a more or less well-marked pi 
maxillary expansion ; the prefrontals (a) are very large, and 0* 
bang the orbits ; while the teeth are curved and caiinated, with t 



ly fluted at the base of The crovn, but without ser- 

thc cariruc The pectoral limbs arc extremely small. 

ranges from the Oxford Cby to the Portland Oolite. 

Oxford Clay and Kclloway rock, both of the Continent 

md, we meet with J/. ikfieroliosHS and M. morelt, which 

iy diMin^ishcd by the sculpturing of the froni^ls nntl 

and relations of the prelroniaU. Beautiful examples 

lus occur in the Oxford Clay near Peterborough, which 

absence of dermal scutes : a skull from this locality, 

the rume Sttmesaurus darycephalus has been applied, 

fcrred to M. su/ifra/ieius. The Portlandian form was de- 

Sttfutfjtnnts gnalii. It is pri>l»ilile thai Gnathosatrus 

tAtasamnts are synonyms. Hie most s[)ecialised genus is 

(Crifsaurus or Dawsaurui), in which the sltul! is com- 

shon. and is do-oid of frontal !>culpiure, and also of ihc 

expanston ; while the teeth (fig. 1088) arc compressed, 

^and orinated, with distinct serrations on the carinse. The 

uttich occurs in the Lower Kimcridgian of Bavaria, 


MiL— Crv*ma(MKhefC«Ma«rw KnaxiJinM; 'ram ituKirovmlM Clay arEIr- 
! <AIWr Wsod-MMon.) 

of comparatively small size, was long thought to belong 
Mosaiixurida. A much lai^er form, occurring in the 
and perhaps the Oxford, Clay of England and the 
ing Iwds of the Continent, has been deiciibed under 
of Dati'saMrtiSt but can be only specifically di»titigui.ihcd 
type; A tooth is shown in the accompanying woodcut. 
K improbable that vertebne from the Lower Grecnsand of 
jr, described under the name of EnalioiuekHs, indicate a 
^iad form. 

embcn of thb family, of which the serial position is imcer- 
y be mentioned small forms respectix'ely from the Kimerid- 
Bavoria and Fraiicc, to which the names jEolodcn and 
tmMs have been applied, both of which haxx a dermal 
the ventral shield consisting of a number of small and 
[lined scutes. In the type of y£o/tfdt>n these scutes arc only 
pitted, and the teeth alternate in sire. 
.V CoKiopHOUDiD^ — The members of this family resemble 



existing Crocodiles in having an open channel connecting dwj 
temporal fossa with the orbit. The orbits themselves m,i 
over, usually but slightly smaller than the supratempofal 
only exceed them but little in size ; while there is do 
vacuity. The dorsal scutes are rectangular, and may be i 
either in two or in several longitudinal rows ; while the 
armour may form either a single or a double buckler, in ' 
posterior transverse rows of scutes may either imbricate or i 
by suture. The members of this family inhabited freshi 
they range in time from the Purbeclc (Upper Jurassic) to thel 
den (Lower Cretaceous), and not improbably also to the 
Greensand. Nearly all the known forms are European, 
genus which may belong to this family is American ; and the I 
is probably also represented in the Cretaceous of India, 
subfamily divisions have been proposed, but before discussiDgl 
we may allude to the genera Suchoseiurus of the English Wa 
and Hyposaurus of the Cretaceous of North America and 
which not improbably belong to this family, although their 
affinities are not yet satisfactorily determined. The type 
former genus is of very large size, and has the teeth greal^< 
pressed ; while in the latter the posterior teeth are of this i 
the anterior ones are rounded like those of GcniofkoHs, 
symphysis of the mandible is elongated. 

The first subfamily, or Petrosucfuna, is very imperfectly 
and is represented only by the genus Petrosuekus, of the '. 
I'urbeck beds, in which the posterior nares are placed 
middle of the skull, as in many of the Teltosaurida, and thei 
are considerably smaller than the supratemporal fossae, 
cranium itself is of moderate length. 

The members of the second subfamily, or Gonioph^iditiA, I 
the posterior nares placed more posteriorly than in the pr 
group ; while there are two longitudinal rows of dorsal scutes; i 
the ventral buckler is divided, with the transverse rows of 
the posterior portion articulating by suture. In the first, or 
rostrine, section of this subfamily the skull is elongated like i 
the existing Gharial ; the nasals do not reach the nares, the 
hone enters into the mandibular symphysis, and the teeth \ 
numerous, and all nearly similar in size. It is represented ' 
Pholidosaurus {Afacrorhynchus) of the German and English 
den. In the Brevirostrine section, on the other hand, the . 
short, like that of the true Crocodiles, the nasals sometimes 
the nares, the splenial enters but very slightly into the symphy 
the mandible, and some of the teeth are much larger thaa < 
others. The dorsal scutes present the peculiarity of artic 
with one another by means of a peg at one angle which 6ts ii 



tn the adjacent scute ; an anangcmcnt vcr>- similar to that 
in il»c scales of certain Canuid lishes. The genus Gonio- 
T«ry characteristic of the WciUdcn and Furhcck, and has a 
of moderate length, with the nxsals not reaching the nn.res, 
otbtt^ rather sonaller than the >upra[cmporaI ru».t;e. The 
ies attained very large dimensions, and vis. lonjj since 
3wn \o the world by the late Or Mantell, under the name 
Crorodile ; its blunt and grooved teeth, and chaiac- 
I scutes, being comparatively common in the ^^''ealdcn stone 
of Sussex, This genus has been recently recorded from 
sic of North America, whore it had been prcviousEy dc- 
as Amphiiotyltts. Allied but considerably smaller forms 
Uorsctihire Purbeck constitute the genera Ntinnosxtchut 
niastteJats {Bnuhjfdedes). The most specialised genus, 
appears to be the minute Theriesuihui of the Purbeck, 
in h.iving the orbits slightly larger than the supra teniporsi 
ximates to the next subfamily, althoujjh retaining the 
donni scuics of GtfHtofihoJis. The nasals in this genus reach 
tly divide the nares, as in the true Crocodiles; and we thus 
comparatively close approximation to existing forms, which 
fttill more manifest by the members of the next group. 
genus Bemissartia, of the Belgian Wcaldcn, which forms 
of the subfamily Bemissarttinte, the &kull is comparatively 
ind broad, and h.ts the posterior nares placed very close to 
ipital condyle ; while ihe oibit* are decidedly larger than the 
iporal fossae. Like existing C'rucodilcs, these reptiles were 
with more than two longitudinal rows of dorsal seuies 
have no pcg-and-sockct) ; white their ventral buckler is un- 
and has the transverse rows of scutes imbricating throughouL 
iblancc to existing forms being complclcd by the pectoral 
[being considerably shorter than the pelvic; pair. The verte- 
still retained the primitive amphicceloui character, 
probable, from the position of the poMcrior nares, that the 
ly known genus UyittoeAampsa, from the English Wealden, 
1y allied form of rather larger dimensions ; and it is not 
that certain proccclous verlebne from the same formation 
I have been described under the name a{ HfUrosuchus may l)c- 
this form ; while others from the Cambndge lirccnsand and 
isand of Austria, which ha^-e been referred to Crocodilus, 
indicate albcd reptiles, although there i-t a possibility that 
of these vcricbrse belonged to the next scries. It will 
seen that '\K Hjlaockamfta, or an allied form, had such pro- 
vcnebrx, it would only require the development of palittal 
to the pterygoids to convert it into a Crocodilian of the 
type ; and it is highly probable that such a form once ex- 
it, u 

1 192 


i&ted, since it n most unlikely that the change frotn 
to procoelou» vertebrae look place precisely at the &ame tinW 
pcerygoidt dereluiwd palatal plates. On thu other hand tbetti 
equal firimA fade probability that these two chaoges 
occurred in the reverse order to that indicated above. 

B. ProcieuaN Seriic*;, — In this series the vcrtcbnc. 
exccptioD of live, ore procoelous ; the pterygoids d<n-elop 
plates to prolong the narial passage (6g. 1 0S9) ; and 

eustachian canals are 
bone. The dorsal scutes . 
over, always arranged in 
two longituilinal rows : 
there is a ventral buckler 
divided, and invariably 
Rtore than eight rom of i 
scutes, in which each scute] 
posed of two separate piecejl 
The axis vertebra differs I 
the TeUmaurid4t, in having I 
articulations : its rib 
shifted forward on to the ' 
ihc atlafi, or odontoid 

Family CKOCOuiurtc- 
sufficicnlly known ineint 
series niay be incJtided in 
family, whieh agreo with the I 
pktiiidida in the free come 
of the infratemporal fossa 
orbit, which is considerab^'l 
than the suprateroponl fo 
family is first definitely knc 
the Upper Cretaceous, and 
lies to the present day, 
sented in the fi^eshwaten 
of the warmer regions of tbc { 
If, however, the proca:)ous 
mentioned 111 the lait series from the CrcenMuxI and ' 
really indicate members of the present family, it will dale 
latter horiion. The form of the skull affords grounds for 
tional divisions. 

The I,ongirr>strinc section is represented at the present dif 
by the true Oharial {Garlalis) of the (langes, and Schlegel'i (^ 
of Borneo. I'hc skull i» produced into a long turrow rottrua 
1091), with the denial borders nearly straight: the natdi 
extend to the iiiiterior aarcs, and are frequently scpamted 


FIi- iiAu.— faUulMiwctor Ihc (Tini um 

nia: M, matilU; PI. Vt.\»X\w. Ti, Tiuii. 
virw; I'l. Huryfloiil : ^f JoabI: Qj,. 
QiuuiniiajuEal : ^, Quadrate \ Of, Kn-i- 
OMiraial : CdV, Ocriplcal ooQilyl*: Ort, 
Orbil i C4, Porttrtar nank 



the supratcmporal fossae are or somewhat large sijie, 

exceed the orbits in this rcspcci ; while the mandi- 

liysis is very long, and embraces the anterior portion of 

bone. The teeth are always numerous, and are only 

' onerjual in size ; and neither of ihe existing species is pro- 

veiiiral armour. In this gruup Jikamphosuchus, of the 

of the Siwalilc Hills of India, was a gigantic form }>ru1>ab]y 

fifty and sixty feet in length, and characterised by ihc stout- 

ihc tectli, of which llie uppei series biles on the outer side 

r, a& in the Alligators, instead of interlocking with ihcm 

-r members of this group. In ThoracDsaurus, again, of 

kccous of North America, we have a genua teinarliablc for 

ibe preorhital vacuity of the Teleoiatirida, and in Having 

itemporal fos^a larger than the orbit; 

features apparently pointing to the r^J 
It of this genMS from the last-named 
without having passed through the inter- 
R«gc of the Gonhpkelidida- The exist- 
Garialis x>^ngetiats, of which the den- 
ts shown in lig. 1090, is one of the 
spedcs of Reptiles of which remains are 
the Pliocene of the Siwalik Hills. The 
(its have also yielded two more or tc!i!9 
lallsrd extinct species; while in the somc- 
Jcr beds of Sind there occur two other 
differing considerably in the form of the 
other cranial characters from the existing 
iiiTe of the genus. One of these extinct 
>(tf./«MKrA)'BM«*) appears to have attained 
fully equal 10 those of RhamfAosuchui. 
Ghamloid provbionaliy referred to the 
genus occurs in the Middle Eocene of 
in Sussex ', while species from the 
of North America, which have been 
under the generic name of Hoiops, may 
be allied either to the present or to the 
1US. In Tamis//rma, typically represented 
c-nisiiiig T. Sch/fgtJi of Borneo, may be in- 
thc fossil forms describe*! under the names 
klitoiourvi and Gariahsuihui (fig. totji). This genus is 
distinguished from Gariaih by the circumstance that the 
leasKl forwards to articulate with the prcmaxilla; (fig. logi), 
of being separated from them by a long interval. A Large 
ftjmi (MdilosMirtts) occurs in the Miocene of Malta, and a 
lOaria/asuiJuis) in that of Austria. Another Charialuid, 

■ion ol g«ni» (i, tt 
httncdfh iKt toQth ID 
u« (a). <Aftw Owm.) 



from the Upper Cretaceous of France and Maastrichl^ 
scribed as Gariaiis maa^rhymhus, has also been 
eluded in the same genus. This form is, however, 



fig- lost. — Fionuil aspni of ihe cianiuni or Tmmiiltm^ vtaAmrttiai I froalht 1 
Austria. One-ieiitbnaiunil tiie. pfi. I'renu'ilk ; mr, Mizilla : jm, Naul: ik,T 
■Vi FronlKl \ rr, OrtnL. Tbe vacuiiy tiebind ibe orlni ii ilie infralemiHWa) To^ 

and Ksil. ) 

Dr Kolten to Thoraeasaurus, although it has no preorbilal 
ard the nasals reach the premaxillae. It appears to connedl 
typical Thsramsaurus with the existing TomUloma. FituDy, | 








Fie. 1092. — Ohiiquc Icfk Inlersl and luperior view of Akull of Crp^dOmt pmhutfitl !■ 
Much reduced, Ine two ftcna]! vacuities id the right are the »upr>t em p jj l *"■'—' itaH 
advance arc ihe orbiiii, while the sinj{le vacuity to the left ii the nwa ; ihc bOHs in adv^iB 
the latter are the prcmahii^K, and those behind the nB^ab. 

imperfectly known Thecachampsa^ from the Miocene of No 
America, should probably be placed in the present group. 

The second^ or Bre\'irostrine section, includes the true Crocodi 
and Alligators, and is characterised by having the skuH short, 



Ij- «Iongat«d, with its dtnlal borders thrown into distinct 
I (fig. loga) ; by the nasals alwa)'s reaching the pretnaxillje. 
Illy enending down to the anterior rures (as in fig. 1 0(>3) ; 
[IbeorljiL* being alwaj-s larger than the >ii|3ralcri|)oraI 
le slwn *ym|>li>^is of the mandihie, from which the 
[dement is entirely excluded {fig. 1093), is another sltiWing 
The teeth, inorco\-«y, vary in siie tn different parts of the 
»nd Usually the third and ninth in the upptr, and the fourth, 
t«)iKntly also the fint and eleventh, in the lower jaw (fig. 

mricvor iIh ItA cwDui oT ihc mudiUe uT* Cn>(«tl1e. Kodund. an. AnguUr; 
liar: M^Canaaid: A Dcnury: «, ipkniali nc.Suranfulat; 17, bymphy^i. 

t considembly larger than any of the rest. In the type 
^vcPjiAit the upper and lower teeth mutually interlock ; the 
tr tooth bites into a perforation or a pit in the cranium, 
fourth into a lateral notch ; while the third lower luuih is 
TT»erc is, moreover, no ventral armour. This genua is now 
ed orcr nearly all the warmer regions of the globe, and it 
to have had an equally extensive disiribuiion in Tertiary 

irlVesI rcpiescnt-itive of lliia genus seem* lo be C. SprtKeri, of 
r toccne of both Enxland .ind Iiuly. vhicli waii a ipccics with 
rxtively long muule like that of the livlnt; American C. in/t'rme- 
be genas ia abo represented in the Mi^idlc Ternaries of Euiopc 
ifa America. In toe Pleistocene of yuecnsland «c meet with 
nt the existing C. ficrcsuj (%. io79)> which now tanj^cs from 
1 to Eastern India ; while in the Pliocene of the Siwnlik Hilb of 
ire occur spccici eUnely allied to the ihi^rt-snuulexl (.'. palustris 
1) of that country, which makes iht nearest approacli in cianial 
rs to the Alligators and their allies. 

<€ytio4tm is an extinct genus found in the Tertinries of both 
and North America, which presents characters intermediate 
Crocodifus and A/figalor. Thus tlie cranium (fig, io(j4) is 
t and hroad ; ihe upper teeth bite on the outer side of the 
the fourth lower tooth is nomially received into a notch 
aaiorally into a pit) in the ci»niuni ; the third lower tooth 
tsthefounh; and there is a complete ventral amiour, 
pe this genus nuigcs from the Upper Eocene to the Lower 
(Upper Oltgocenc), and is common in the Tertiaries of the 



south of England, and also in those of France and Germany;] 
of [he !ipecic<i from ihc latter deposits having been 
Atiipilor. The genm has also been recorded from the 
Eocene of North America. The last, and in respect ofi 
characiers the most specialised, group of Crocodiles U nowi 
into the (wo genera Caiman and AUigatar. In these 
the first and the fourth lower tooth are received into 
upper jaw, so as to be concealed when the moutli is 
upper tireth bite on the miter side of the lower ones ; the 1 
pixal fossa: arc vcr>- small, and are occasionally oblitc 
the third lower tooth is smaller than the fourth. Com 
tinguished by the presence of a \-cntral armour, and abo 
circumstance that the nasals do not extend across the nssj 

Tig. lOM- — OtiliqDt ItA Iklertl •nd p^Ulal view vT ih* (itcUl pan of ik« cnonn of J 
HuttPiufiuii : tnm eh* Upper Eocana of Hanp^hin. Radxvd. 

is now confined to Central and South America. In A/Jifi 
the other hand, which occurs at the present day in North 
and China, the ventral armour is absent or extremel;^ thin, 
nasak extend forwards so as lo divide the nanal aperture, 
which arc probably referable to Caiman occur in the PIctt 
cave-deposits or Brazil : but it does not appear that there it < 
certain evidence of fossil species of AUigaier; the European,' 
probably some of the North American forms which have been I 
feired to that ^'cnus, belonging to Difl&cyntrdfiH. Filially, the Ml 
IsuhnaHrui has Iiten recently applied to Crocodilian remains fit) 
the Middle Eoocnc of Krancc which may really belong to one of d 
above-mentioned genera. 

Ohper X. Ornithosauria. — The Pterodactyles, as the laa 
ben of this extinct order are commonly termed, are among i 
most remarkable and strange Reptilian forms that PaJjcontokgy * 

' Accoriling lo Ihe smngi^ment adopted by Mr Boiil«fer in bb Brit 
MuKam (.aiiici|^c of this oniec. 



to Us. So Ktrangv, indeed, are they that some autho- 
considered that Ibc 

Brorn arc mtitltd to rank 
tiiKt class ; but they are 
ly Reptiles, and agree in 
thtuic characters with the 
■o orders placed in the 
now under consideration. 
U organtsjtion h, however, 
d for the purpose of flight 

tbc air. Thin the txidy 
iponcd during flight by a 
mous expansion, or /a/a- 
iliich w%xA mainly l>ome by 
uljr elongated phalangeals 
ibnr, or outermost digit of 
■OS (fig. 1 095, maritcd iv) ; 
dh aUo inctcniled along the 
' tbc I»ody lo cminacc the 
mis and tail (fig. 1098). 
ncbra; are proccelous, and 
letr neuTo-ceniml suture 
the precaudol &criea 
lumerous : the cervicab are 
the dorsals ; and rroiu 
&ix t-erieb[;e are an^rhy- 
pgetbcr to form the sacrum. 

."ical ribs in tbosie cases 
tbcy haw been observed 
Crocodilian lypc. The 
1096, 1097) is rela- 
; and although more 

btrd-Iike in general con- 

t mainlaim the reptilian 

the presence of the supra- 
bI tosat, bounded by the 
n of the poetorhital with the 
MaJ bone Bird-lilcc resem- 

I are, however, shown by ^ ,^^ _ ^_ r;^. p^,„„, ,;„,, „r 
cunoitance that the bones of Srvl.v»-rt« (««.."«f™; ^ Si.™«B, 

„ . 1 J 1 "iJ "inKold* of I'Urrdnctyfnt. h, Hu- 

Sill anctiylosed together at lueru.i ^. r«i>u>: ■. iiii»; t. Cxtpu*: 
ly age ; and that the two l;^';*"- "«" '"'^'^ *"" '"" """"■ 
f the mandible were cotn- 

wcldcd tO£cihcr at ihcir syniph>-s]s. Moreover, as in Birds, 
■ater portion of the upper jaws is formed by the premoxillo* ; 



:he nares are similarly approximated to the orbits, with lb 
rention of a preorbital vacuity, which (fig. 1096) may b 
large ; while the occipital condyle is situated on the base 
:ranium ; and the orbits are large, and there is generally an > 
nng in the sclerotic The teeth are invariably simple and p 
ind are always implanted in separate sockets. In the i 
>irdle the scapula and coracoid are long and bird-like, and th 
tias no fontanelle ; there were no clavicles ; but there was i 
[leart-shaped sternum (fig. 1095, b), carrying a median keel 
Drly. The angle of junction of the scapula and coracoid, 
IS the keeled sternum, curiously resemble the correspondin 
in the skeleton of the Carinate Birds, and are consequendy 
unlike those of the Ratitse. The carpus consists of two main 
one distal and the other proximal ; ^ while on its radial side 
a small styliform ossification, regarded by Professors Ow> 
Marsh as the representative of the poUex — an identification 
if correct, will make the four remaining digits which are 
present the zd, 3d, 4th, and 5th of the t3rpical series, and 
ist, 2d, 3d, and 4th as they are regarded by some writt 
1095). The phalangeals of the ulnar digit, as already mer 
are enormously elongated, and the tenninal joint has n> 
The pelvis is relatively weak ; and although the ilium is o 
on both sides of the acetabulum, the structure of both thu 
and of the pelvic limbs is far removed from the aviai 
Thus the pubis (or prepubis) is directed forwards, and the 
is short and wide ; while the pelvic limbs are relatively short 
fibula is, however, always fused with the tibia ; and the as 
may also unite with the latter bone ; although the metatarsals 
remain distinct both from one another, and also from thi 
row of the tarsus. The greater number of the bones are 
and are frequently provided with pneumatic foramina, like t 
Birds. The brain was bird-like, and the body was probably 
In time this order ranges from the Lias to the Upper Chal 
was especially abundant in the Upper Jurassic and Cre 
strata of both the Old and New Worlds. Although the s 
presents many remarkable resemblances to the Carinate Bi 
these must be regarded as mainly due to adaptation for a 
mode of life ; since it seems clear that the Pterodacty 
altogether off the direct line of the Avian pedigree. 

Suborder i. Pteranodontia. — In this suborder te 
totally wanting, and the jaws were probably completely ensi 
in horn, like those of Birds. The skull (fig. 1096) has an en 
suprnoccipital crest, projecting far behind the occiput ; and tt 

' The carpus is enoneous in 6g. 1095. 



ipletrly conftucm with lh« prcorbital vacuities. According 

Marsh, thtse reptiles were mostly of gigantic size ; some 

■prrad of wing of nearl>- or quiw 25 feet And in order 

)c powerful pktagium in flight, the pectoral girdle was 

Uretigthcned by the anchylosis of several vcrtcbnc, and 

: «cq>ute articulating to the spines of these anchylosed 

— t^ti laiml new of ihc ikull ot PUramnim fougictfi: (tom >lic CmHROui ti 
IMA, OrM-mlDb natnnl mm. ■>. Nuu uid prverbiul ■-».-iiti)' : i. Otlni; t, 
uui; A Aack of ■audiMa; r. V"*'''*'*! '■ !>TB>ph)iijt (After Manb-J 

1; this peculiar fe;iture being vinually a repetition of the 
irdle arKl sacnim on a much larger scale. 
V PriiRA'toikiMli)-*. — Thf type gL-nus Pleranodon (!ig. 
ccurs in the Cretaceous of Noflh America ; and altliouj^h 
bers arc generally of large size, it i.s also represented by one 
ecics — I', nanus. The coracoid and scapula were united, 
oral aspects of the jaws have not the ridge and groove 
Omithfithirus. Ornitheitoma, of the Cambridge Urcen- 
ly have been an allied form. In Nyft»tla{/y/ns, of the North 
Cretaceous, Professor Marsh thinks that none of the 
cnebra; were anchylosed; and on ibis account ihc genus 

rhaps form the type of a di;>tinct family. 
UilH 2. pTKKfJSAUKlA. — In this, the typical, suborder teeth 
nt in both jaws ; the cranium (fig. 109;) has no long supra- 
crest directed backwards, and generally has the narcs more 
completely separated from the prenrbiial vacuities. The 
is (at least usually) not nnchyloscii 10 the neural spines, of 
TOftebne, which are distinct from one another. Thi-t sub- 
mainly European. 

LV Pterodactvud*. — In the typical family the tail i» 
J. 1097); the jaws are toothed to iheir cxtmnitiL-s ; and 
of the metacarpus con<ttderul>ly cxceed<> half that of the 
1097). The sliull, which is extremely bird-like, may he 
or short, and has the nares imperfecily separated from 
bital vacuities : while in the pelvic limb the astragalus is 
istinct firom the nhia. In Europe this family is especially 
ic of the Upper Jura-»<ic, and is abundantly represented 
Kiineridgian iiiliographic limestones of Bavaria, which, 


from their fine stnicture^ have preserved not only the smallest 1 
but not unftequently also the impresam of the VKWtbamom 
gium. All the forms which can be certainly referred to dm 
are of small or moderate size. In Jtemodraar, of the Itthag 
limestones, we have a small Pterodac±yle not larger than a ^^ 
with a skull of very much the same contour as that of tbe Id 

Fig. ingi.— Nearly enlire skeleton of FIrredatlyllt iftcliMlii: fttna tbc KJMO 
Havana. U'he ventral aspect U tbown ; and on ibe right aide tbc ilium, and on tbc left 
<d) it eipowd. 

which the teeth are confined to the extremities of the jaws, s 
nares do not appear to be separated from the preorbital va 
Pttrodactylus itself (of which Omithoaphalus'^ and Diopu. 

' It has been proposed to take Ihe name Omiihoetplialtu in place c 
draco, a suggestion which is entirely opposed to all ine tulet of nome 
The Rrst use of the former name in this Mtise was made by Fiu 



iifm&), on the other hand, has the sliull produced into a 

or rostrum (fig. 1097), in which the teeth extend over a 

le space, and Ih« large nares are slightly si-paniifd from 

■taJ %-acuities. The scapub is not anchyloscd to ihecora- 

the pubes are shon and laundcrd (fig. 1097), This genus 

to be oonBt>c<l to the Uthographic limestones ; the typical 

us Iteing about the dimensions of a woodcock. 

lUy allied to this genus is Dermodaclylus, of the Jurnasic 

th America, in which the bones are said to have thicker wnlls. 

ftype !)pecies is e»limated to have had a itproid of wing uf from 

to MX fe«t. Firully, in Cymorkam^aa, of the Solcnliofcn 

s, w« have a genus with a broad expanded beak, like that 

Swan, with the teeth confined to the anterior cxETc-mity. This 

is represented by a single species of comparatively large si2c, 

as C. Skeviois. 

Here we may conveniently notice some gigantic Ptcrcnlact)-les 
the Cfctaceou-s system of Europe, of which, owing to the cx- 
lii\^iy imperfect remains hitheno discovered, the family position 
be Icl'l undeieinimcd. Remains of some of those fonivs were 
ily r^arded ax Wlonging 10 BirdR, and described under the 
of Pa&rvmis (Wealden) and Cimttlwmis (Chalk). .A.11 these 
may be provisionally included under the generic name of 
vrvr, although it is highly probable that some of them may 
be distinct ; and there does not at present appear any very 
reasons for separating a Purlicck form for which ihc name 
icrkymJttis bas been proposed. Many of these i'terodactyles 
of gigantic siie ; the spread of wing of some of the larger 
being estimated at as much as 35 feet. Pmhahly the 
long; the jaws were toothed to their extremities, and 
Jy the upper anterior teeth cur\'cd forwardfi to project 
advance of the muzzle, llie oral surfaces of the uppvr and 
jaws were marked by a longitudinal ridge and groove; 
&kull was either »hort and stout, or much elongated ; the 
was often ar>chylo«ed to the coracoid ; and in some 
ices the astragalus united with the tibia. It has also been 
ted that OmilhtKhirus had but three digits in the manus, 
this statement requires confirmaiion. The name Crelomis 
been applied to the remains of Ornithodiinis from the Chalk 
t Bohemix 

: Family RMAMPHORHvr<cHiiJA. — In this family the tail wa-s at 
HKt tisually long (as in fig. 109$); the extremities of the jaws were 
■ many instances edentulous; and the length of the metacarpus 
iii much less than half that of chL- ulna (fig. 101J5). The fi.kull 
Ig. 1099) waa Ics^ bird-like than in the type family, with the narcs 
lied by a distinct bar from the preorbilal vacuity, and uras 



often comparatively short and stout j while in some ii 
astragalus united with the tibia. This family certainly 
the Lias to the Upper Jurassic, but if Omitkotkirus belong 
its range must be extended to include the Cretaceous. In l 
genus Siapkngnatkus (fig. 1099) the teeth extend to the 
of the jaws of the massive skull, in which the nares are sef 
by a broad bar from the larger preorbital vacuities. The Oil I 
the type species is unknown, and in Goldfuss' restoration 
1099) it was made like that ol Pttrodactyius ; but Professor ! 
considers that it was elongated like that of RhampherhynAti 
Dlmorplwdon (fig. iioi). The type species, which attains 
iiiderabile dimensions, Occurs in the Kitlteridgiian limestood 



Fig. ID9S, — ReKoratian uf Dimorptuidoii Hacrenyx. Rnjucml. (Afler Owmli 

Bavaria, hut the genus is also represented in the Upper Lias 
Whitby. It is noteworthy that the peculiar form and relatioM' 
of the jugal and quadratojugai found in the Dinosaurian gram 
DipfodiHiis (fig. 1076) also obtain in Scaphagtiathui. In RhamphB- 
rhyn^hits, of the Kimeridgian of Bavaria, the extremities of the 
jaws are usually devoid of teeth ; while in the hinder region the 
teeth inclinu forwarda instead ^f having the nearly vertical direction 
of those of Scaphognathiis. The scapula and coracoid were some- 
times anchylosed ; the astragalus was generally distinct from dx 
tibia ; the pes had either four or five digits ; the pubes were slender, 
bent, and joined by a bony symphysis ; while the long tail »» 
strengthened by the ossification of its tendons. The membranous 
]iatagium developed a leaf-like expansion at the extremity of the tail, 



have been an allied genus. In the Lower Liassic genus D, 
odon (fig. iioi) the jaws are toothed to their anterior extn 

and the hinder teeth of thi 
dible are much smaller than 
in front. Both the nares ar 
orbital vacuities are of en 
size, and are separated by a 
bar. The coracoid is and 
to the scapula; and the ast 
united to the tibia. Dimm 
is thus the earliest known re 
tative of the order ; and tl 
species attained considerable 
sions. Its remains occur in the 
shales of Lyme Regis in Docs 
and were fiist brought to nc 
1 8 3 3 by the indefatigable Dear 

Ordinal FosmoN Unceri 
Here may be noticed a genus o 
the serial position must for the 
remain undecided. It is kn> 
Omiihodesmus, and was founde 
an imperfect sacrum from the 1 
Wealden, which has been rega: 
Avian, although its right to 
tion from the Omtthosauria : 
by no means certain. It i 
observed that the so-called C 
pterus, of the Upper Jurassii 
stones of Bavaria, said to b 
acterised by the presence of oi 
digits in the ulnar digit of the 
and which has been regan 
Avian, appears to have been 
ed upon an imperfect specie 
Rhamphorhynckus ; and it m; 
be mentioned that the name 
thopterus is preoccupied by th 
dopterous genus Omithoptera. 
ally, it has been suggested 
tooth from the Trias of lb 

scribed under the name of TribeUiodon may indicate an 

saurian at that early period, but the evidence in support 

view is at present wholly insufhcient. 

Fig. IIOI. — ResloreU slt«letoii of 
lyimorphedpK maCTVKjx ', from the 
Ixjwer Lia«. Rtduced. /. Ulnar digit : 
m. Other digits; /, Mciatunuii. (After 




. (G.) — " On ihe Phytwcnctic Arrangcinent of the Sauropsida." 
'Journal of MoTphoIO(n'? v**!- i- (i8H7)- 

ILXNGEK iC. A.)— "CaaJogue of Clieloiiians, Sc, in the British 
ram' (1SS9). 

ksn (W.)—" Geology and Mineralogy" (1837). 
: CK- I).) — ■■ RKtinci Batrachb, Repiilia, and Avra of North 
America.* 'Trunv Amer Phil Soc.,' vol. xiv. p, 100 (1871). 

" Homolo^e^ of the Cntninl Hones nf the Kcptilia, and the 
Systematic Arrangement of the Class." 'Proc. Amer. .'\&snc.' 
for tSTOL 

'The Rehlions between the Theromorphous Reptiles and the 
Mnnotreine Mamnuilia." ' Proc, Amor Assoc." for 1884. 

"The Vertebraia of the Creiaccotis KortnatJons of the West.'' 
'Rep. U.S. GeoL Surv. Terrs.,' vol. it. (iS;;!. 

See al&n ntimerous memoin in variou-; Amcriean serials. 
IKKUNKR (H.>— " Die SlCBoccphalen iind Saurier aus dcm Roihlie- 
nsiden dcs Plauen'schen Grundes bei Dresden." *Z«itschr. 
deotscfa. geol. Ge«dl«chaft' ) ItX i -88. 
SLONRCHAUPS (E.)— " Note* Pa!*ontoiogiques"(i«63-6q). 
»U.O<L-)— " Note jur le* Dinosauiiens de liemissart," 5 parts. 
•BulL M«s. R. d'Hisi. Nat. BelR.' t8«i-84. 

' Note sor I'Osteologic dcs Moiasaurid:p." /hJ. 1S83. 
Se« also other memoirs in same and other serials, 
StfKTHER (A.) and MiVAHT <St t;,)— " Kcptilia." ■ Encyclopedia 

Britaonica,' 9(b ed. 
HCLKB 0- W.>— " Preudential Addreue* to the Geological Society." 

'Supplemental Note on PofacaHlhut F&xii^ 'Phil. Trans.' 
18K7. pp. I67-I7J, [J- viii, 

" Conribution* to ihe Skeletal .Anatomy of ihc Mcsoiuchia, 
based on fossil remains from the clayc near PeierborouKh, in the 
eoUeaion of A. N. Leeds, Esq." ' Proc. Zool, Soc' 1888. 

See also other memoirs in ' Phil Trans,' and ' Quart. Joum. 
G«oL Soc' 

HCXLEV (T. H.J— •* A Manual of the Anatomy of Vertebrated Ani- 
ixtal*,' 3d ed. London. 187T. 

— "Cki ihe Affinities between Dinotaurian Reptiles and Uinb." 
•Quart- Journ. Geo). Soc,' vol. xxvi. (rS?©). 

"On the Classification of the Dinoisauria." torn. at. 

"On Sta^imt^epit and the Evolution of the Crocodilia," ibid.. 

voL xxxi. f'1875). 

" On lirfxrhdaptdfit^ IbiJ., vol. xtiii. { 1 887). 

See al>o numerous memoirs in the Mmc and other serial a. 
KlPRiJANOFrlW.)— "Studicn uber die Ptfssilcn Reptilicn Russlands." 

' M6n. Ac Imp. St Petersburg.' voU. jtwiii.-sxxi. (1881-83). 
KOKXN (E.>~"Uic Uinosauricr, Crticodilidcn. und Sauroptcrygicr 

do Norddeutschcn Wealdcn." 'Pal. AbhandL,' voL iii., pt. 5 

LCJPV {)•>— "Cretaceous Reptiles of the United Stales." ' Smiths. 

Cootrib. KnowL,' vol. xiv. (1864)- 


23. Leidv (J.)~" Contributions to the Extinct Vertebrate Fuiu 

Western Territories." ' Rep. U.S. Geol. Surv. Terrs.,' vol i(l 

24. LyDEKKER{R.)—" Catalogue of Fossil Reptilia and Amphibii 

British Museum." Parts i., ii., iiL (1888-89). 

25. " Indian Pretertiary Vcrtebrata." ' Patseontologia India,' 

4, vol. i.. pts. 3-s (1879-85). 

26. " Indian Teniary and Post-Tertiary Vcrtebrata." iSuC, m 

vol. iii., pts. 6, 7(1885-86). 

27. " On Eocene and Mesoioic Chelonta and a tooth of (?) 

iifisit." ' Quart. Joum. GeoL See,' voL xlv. (1889)1 

28. Marsh (O. C.)—" Classification of the Dinosauria." 

Journal of Science,' ser. 3, voL xxiii.(i882), and 'Rep. Brit 
for 1884. 

29. " Principal Characters of American Jurassic Dinosamt.' 

parts, /eiii., vols, xvi.-xxvii. (l878-84>. 

3a " Principal Characters of American Cretaceous Pte; 

/6iW.,vo\. xxviL(i884). 

31. ■ "Structure of the Skull and Umbs in Mosasauroid 

Mid., vol. iii. (1872). 

32. Meyer (H. von).— " Saurier des Muschelkallces." (1847-55-) 

33. "Saurier aus dem Kupfer-Schiefcr der Zechstein-foi 


34. Owen (R.) — " Descriptive Catalogue of the Fossil Repliba 

Fishes in the Museum of the Royal College of Surgeons of 
land" (1854.) 

35. " Catalogue of the Fossil Reptiles of South Africa in the 

lection of the British Museum." (i876l) 

36. " Monograph on the Fossil Reptilia of the London Clay." 

i., Chelonia (Owen and Bell). ' Palfeontographical ',' 
(1849.) Part ii., Crocodilia and Ophidia. /^V. (1850.] 

37. " Monograph on the Fossil Reptilia of the Wealden and Purfaed 

Formations." /did, (1853-59, ^^^ Supplements extending 

38. " Monograph on the British Fossil Reptilia of the Ooliuc F» 

mations." Wd. (1861.) 

39. " Monograph on the Fossil Reptiles of the Cretaceous Fomii- 

tion." /Aid. (1851-64.) 

40. " Monograph of the Fossil Reptilia of the Liassic FormaiioDS.* 1 

/h'd. (1865 and 187a) j 

41- " Monograph of the British Fossil Reptilia of the Kimmeridiei 

Clay." lifd. (1869.) : 

42. " Monograph of the British Mesozoic Reptilia." /ii'd. (1873, ; 

1875, and 1877.) 

43. "Odontography." (1840-45.) 

44. "Report on Fossil Reptiles." 'Rep. BriL Assoc' for 18]) ' 

and 1841. ' 

4;. "On Uicynodon." 'Trans. GeoL Soc.,' ser. 2, vol. vii. (1845). 

46. "History of British Fossil Reptiles." (1849-84.) 

47- " On Parts of the Skeleton of Meialania platycfPs." • PhiL 

Trans.' 1888. 

48. Phillips (J.)— "Geology of Oxford and the Valley of the Thames.' 

Oxford. 1871. 

49. Riley (H.) and Stutchborv (S.)—" Description of Remains of 

.Saurian Animals from the Magnesian Conglomerate of Bristol'' 
' Trans. Geol. Soc.,' ser. 2, vol. v. (1840). 


EV (H. G.) — ** On the Dinosauria of the Cambridge Greensand." 
' 'Qiuut. Joum. GeoL Soc,' vol. XXXV. (1879). 
—~ •"On the Reptile Fauna of the Gosau Formation." Hid, vol. 
xxxviL (iS8i> 

— ** On ttkc Dinosaurs of the Maastricht Beds." /did., vol. xxxix. 

m— ** On the Mode of Reproduction of certain species of Icktkyo- 

saitrtts/' ' Rep. Brit Assoc' for i8Sa 
— ^ " Researches on the Structure, Organisation, and Classification 

of the Fossil Replilia." Van '\., Protorosaurut Speneri ; parti!., 

PareiasoMrtts ^tnbidetu. ' Phil. Trans.' 1887-88. 
^_ " Index to the Fossil Remains of Aves, Omithosauria, and 

Reptilia in the Woodwardian Museum." Cambridge. 1869. 

"The Omithosauria." Cambridge. 187a 

See also numerous other memoirs m the above and other serials. 
SOLLAS (W". J.) — *' On a new Species of PUsiosaurus, accompanied 

by a Supplement on the Distribution of the Genus." ' Quart 

Touro. GeoL Soc.,' vol xxxvii. (1881). 
PTOODWARD (A. S.)— "The History of Fossil Crocodiles." ' Proc. 

GeoL Assoc/ vol. ix. (1886.) 

•>!_ II. 




General Structure. 

The fifth class of the Vertebrata is that of Aves, or Birds, 
as we have already mentioned under the description of tb 
Rcptilia, presents a number of characters in common t 
latter. Birds, indeed, as Professor Huxley remarks, are : 
so similar to Reptiles in all the most essential features c 
organisation, that they may be said to be merely an exi 
modified and aberrant Reptilian type. Their diflferentiat 
however, so great as to indicate without doubt their right t 
a distinct class. It will be unnecessary to recapitulate thi 
acters common to Birds and Reptiles — together constituti 
province Sauropsida — and we may accordingly proceed t 
the distinctive features of the former class. It may be wd 
ever, to observe before proceeding further that, according 
arrangement proposed by Professor A. Newton, Birds are i 
into three primary divisions or orders, respectively kno 
Saururfe, Ratitte, and Carinatje ; the fossil representatives of 
will be noticed in the next chapter. 

In the first place, all Birds, so far as can be ascertainei 
provided with the peculiar epidermal covering known as f< 
which are totally unknown among the Reptiles ; while ossifi 
in the dermis are extremely rare, and never take the form o 
scutes. No Bird, again, has procoilous vertebrae ; while in 
isting forms the centra of the cervicals have cylindroidal, 
shaped, articular surfaces, although these are amphicoelous in 
Mesozoic forms. In no cases are there sacral ribs for atta 
of the ilia in the proper sacral vertebne. The sternum 
backwardly-produced median processes for the ribs ; all of 
arc attached superiorly to its lateral borders. If an inter 
ever exists, it is fused with the clavicles into a compoun 

the fiitcuU (fig. 1106). N'o Bird has more tliaii three 

in the nunut; aU of which may be furnished wilh claws. 

three elemenu of the pelvu sua nearly always anchylosed 

lODcihcr in the adult ^fig. iio}); the ilium being produced in 

»<iv-n'-e of the acetabulum (of which the inner wall 1* unossilied), 

'.' itchtum :in(t pubt« dirccttrd backwards, in a mure or lcs<i 

1 dtrcctioo, and only very rarely meeting In a vcntial sj-m- 

Thc proumal row of the tarsus is always united with the 

i.) funn a tibio-larsus ; while the distal row coalesces wilh the 

-imlil btvnl new of lb* piMt »nil rawld vinnhni of ■ ^owl (c.WJHi). //, Ilhin ; 
iami /V, PuUii ■it", AwuVulimi M, DothpIiuiiIxu V'tcMtiic. Cd, CBuitof do. 

median metatanals to constitute a tarso-metatarsus. In all 

tt Bud» only the rij^ht aortic arch is present ; the arterial and 

circulations meeting only in the capillaries ; and the blood 

ae of the above characten are common to certain Kepdles; 

it is only the whole of them collectively which can be regarded 

tchjjactenstic of Birds iu a cliiss. 

lotking in rather more detail certain features of the osteology, 

ay be observed that the skeleton is usually remarkable for its 

>iiution of compactness and lightness, and also by Che per- 

aon of ihe greater number of the bones by air<aviiieM. The 

(fig. 1 103) is formed on the general reptilian t>pe, but is 

able for the greaier relative development of llic brain-case, 

gh this feature is nearly [laralleled in the Omithos-nuria, The 

lent bofKS have a great lendcnry to unite in the adult by 

obliteration of their sutures, and their texture is delicate and 

»ng>-, and totally unlike the irory-like structure so generally obscrv* 

among Reptiles. The single occipiinl condyle, which in mainly 

hy the bosioccipital, is not plncctl at the hinder extremity 

the cramum. hut becomes shifted forwards and downwards, so 

basal oiia of ihc tatter forms an angle with tlie axU of the 



Column. In Reptiles a simtL-ir feature ocmn 
ria. 'I'hc inferior tempoTal arcade, formed by 
quzdratojugnl (tig. 1103), and connecting the masJIa ' 
Quadrate, is invariably ])resenl, hut the superior temporal ar 
always wanting; and there are ne\(rr distinct poslorhilals or; 
frontals. The complete inclu»an of the paricul-t in ihc roof 1 
Ijrain-case prevt-nts the funnation of tht po^ttemporsl fosae, ' 
constitute such x charactcrbtic feature in the skulls of the 
of Reptiles ; and no Bird has a parietal foramen. The hue at \ 
cranium is formed by the baniocdpiul and basispbcnoid, fron 
latter of which proceeds ihe rod-hke iphenoidal rostrum, np 
ing the anterior pan of the parasphenoid ; while the posterior ] 









ji^d^kkMml view of (he ilcufl of Ihe Fowl. <b. Denun fonioB oTdH ■ 
or, Ania4l'|WftP>'<'°-< /■• Qiudnu i f, SquinOHil; m Km«Tf itil ; tn, Sapa 

«, pBttanni /H'^MUli /■, L*tfaty<nal; iH, ^aul; tv, VwKr; fmi. l^mmti 
BilIU: JiM; ]li0l : 1)/, lJmUln.toiuBil : fl. Puryf aid : y^. PilUtM; u, launrUuli 

of the taiter pertiitls In the basitemporal plate underlying the 
oecipitiil and Ijosisphcnoid. There is always a prcorhital (1 
nasal) vacuity bciwecii the nasal, lachrymal, and maxilla (the 
angular space immediately behind the nasals in fig. iio^)k«til 
many Mtinct Reptiles; and the imcrorbital septum Is always 
or less ossified. 'I'lie narial aperture (tig. 1103) is LUcial, 
nearly always placed a short distance in ad\-ance of the orhili 
the root of Ihe beak. The jjreater portion of the latter i« 1 
as in the Oriiiihosauri-i, by the prei»a\ill«, which coalesce at^ 
cady period in the middle line, and thus form a irirodiatc Mt*] 
giving off a median nasal and a pair of lateral maxillary pr 
The pterygoids (fig. 1103) ne\-er unite together in the middle Im 
to form a completely closed palate ; and neither those bones not (l*j 
palatines ever develop inferior palatal plates to separate the nin^ 



from the mouih. In ihis respect, therefore, the de\'eIop- 

: of the Bird's skull docs not nttain Mich a special Lsnt Ion as that 

existing Crocodiles, The quadrale is usually iiiovably at- 

to the -^qiLamoul ; and the ftniculation of ihe pa]atuptcr>'goi(l 

the iKisipwrygmd proce»es of the spheiioidol rostrum h also 

one, by which rntans the premaxil)ur>- beak can be moved 

lin extent upon Ihe rest of ihe skull. The vomtrs arc 

to great variation. Tbcy underlie the ethmosphcnoidal 

and when present .■m; connt'cted posienorly with the [Mla- 

^ except in tbc Ostrich. The relations of lhc»c find the oUicr 

of tiic pahte form importan: features in Professor Huxley's 

Scalion of Birds; but since this i» a subject to which the 

lion of the Paleontologist is bul seldom directed, the reader 

of fvirthcr iflfonnation must refer to other works. The 

rl and the quadratojugat arc slender, rod-Iike bont-s. of which 
fonner atniculates with the equally slender nioMlla, and the latter 
' B hollow KUrfare with the quadmlc. In all exL<iting llirds the 
alary elements of the two rami of the mandible arc always found 
at the symphysis into a single bone ; but in the CrctaccoU!> 
M>, and perhaps in other Mesoioic forms, this union ta 
fe«:t. There is frequently a laicral vacuity between the den- 
and splenial, like that of the Croeodilia. The angle of the 
liblc may be cither truncated, or produced into a long recun'ed 
OM in the Fowls (fig. no.'}), Ducks, and Geese, In exist- 
and Tertiary Birds the beak is cnshcalhcd in hum, and is 
ly devoid of teeth ; but rudiments of teeth ha\'e been found in 
DC Parrots. And in certain Mesoioic forms the ])rcmaxilla, 
ilia, and dentary bones were furnished with a complete series 
Iccth. A ring of bones is always developed in the sclerotic 
I some McsoBotc Birds ihc vertebral centra were amphicoelotjs, 
lin all others the %'ertebnE exhibit certain well-marked peculiori- 
Thus the neural articulations are always well do'eloixrd, and 
Mtrh a invariably ariicuhled to the centrum. The neck is 
Ry ^xry long ; the number of its vtrtt-bra: ranging from ei^hl 
Itttciuy-three. The atlas vertebra forms a thin ring, in which the 
iwrse ligament may be ossified ; and the axis nlw.iys has the 
niaid process anchylustrd to it. The sucx'ccding ccn'icals liavc 
' short neural spines, or no spines at all ; the anterior surfaces 
[ihdr centra arc cylindroidal and convex from above downwards, 
IcMicavc from side to side, the reverse condition obtaining pos- 
r|y (fig. 1 104), These surfaces arc usually dc5cril»ed as soddlc- 
1;^ and there may be a haemal spine inferiorly. In the imma- 

n« tetM it/ermwlmt baa been proposed for this tf^ of vcrklira] »tiuclurc. 



turc Ratiia; the ccrricals Kavc ribs ailjculaltng with an opperj 
lover process, as in the Oocodilia ; and in the adult {6g. 
these ribs anvliylosc to the vertcbta:, and thus rcsctnlric pafd^ 
transverse proeewes, in which the canal (/) staves for the |ir 
of the vertebral artery.' In adult Caimalie thvsc lateral ^c 
lh*y may be termed, become further modified, and derckfl] 
longations for the piotc-ction of other pana of the va 
system. 'ITie dorsal vertebra; arc liable to variation m ni 
their centra usually resemble those of the cervical rcfpoo, 
the Penguitw the articular surfaces of sonie may be spheroid 


ris. ii"*.^*} Aiiitiior aiiil(R)praterie> vlewi«f Rfcniot •intUioT /li4firm-*u i 
"^ 'W C«t»<»o«n» of Nonh Air-— — - " ■ — '-- - - "^ -»-- - -■ ■»-- 

ftgai ilx C«t«-m« of Nonh Ktanntm. i, Nesnl tp«iw ; *, PiunaOfdiTH* < ■ 
phnM: ^, rnniTtri* p«ix(u.(W(tiipDpAji^i A Rflf ruw, or MraponliVKt* ; mt, N 
/, CODal uiuJ. (AflarUuvh.) 

teriorly.' They usually have well-marked neural, and may or 
not have inferior median spines; and they ore in some 
anchylosed together, but in others arc susceptible of a 
amount of motion. Tliroughout the whole dorsal series the 
well-devt'loped transverse process from the arch for the tut 
of the rib : while the ccntnim has a lateral facet for the lapit 
The nielhud of costal articulation resembles, therefore, that < 
in the first two dorsal vcrtebne of the Crocodilia. These 
are characteristic of all Dirds. The dorsal vertrbne are sv 
posteriorly by a number of anchylosed vertebra: forming the : 
According, however, to the researches of l>r Gadow, only 
three of these vertcbne are truly sacral ; those in front hcloti 
the lumbar, and those behind to the caudal region. Of the 
sacrals the two hindmost correspond to those of the Crocodiliit^ 
the second of these to the single sacral of the Amphibia. The! 
tcbrx articulating with the ilia do not develop ribs, but < 

I TkU *muic«incnt or ibe rihi ti prectMly limilftr Ift Ihftt nffiiin wi 
Dlnounn, u t« ihd* n in fij;. 1071 (p. 1170). 

*Ctr[iitn Waiff and Wading Kiidi sx well sc Partoit, and ike fcmnt 
StfU^rtiii h.itc opiithocfclous donal vcilcbnc Tbc I'utots alio hate 1 
to tliCK vntcLutc. 



r'uf. iw).— Sicnut mAoti ol the 
Scipnb ; t, Coracold. 

p»oce»»es placed on the aich. The anchyloscd &«ri« 
which «i« sometimes Icrmed uro-sacml, are in some in- 
in Archacpteryx and Rhtay followed by a coniid«nible 
>r free vcrtcbnc, but more usually by only a few, succeeded 
riaiifjular icrminal bone, known as the pyioHyle (fig- i io6), 
carries the tail feathers and ^Linds, and reprwenis several 
«d vertchrx. In no known Birds are ossi^ed intcrcenlral 
tts developed, wiih the exception of tlie infciior bar uf the 
ertebra. Neaity all the dorsal ribs 
p tubercular and capitular pro- 
, and Mime have also uncirute pro- 
, (fig. 1 1 06, w/). The sternum has 
>f gTOiivcs superiorly for the tt- 
llic corawids ; and in the 
1105) it is rhonilMjidal and 
jt trace of a median kcd, 
Ijmcut taking place froni tu-o 
ES. In the majority of Cari- 
, jXcfnum (6g. 1 106, si) '\%, how- 
, and has a strong median 
altachincni of the pectoral 
In this type two membranous 
fTe<|Ueni]y eutl in the posterior portion, whii'h in the dry 
form holei or notches, separated by bony processes, which 
at dimions of the Mammalian xiphiMemuni. In many 
IdUK, and especially the Passerines, there are als.0 developed a 
^m manubrium stcmi, and lateral co»ul procesMi^s for the attach- 
" the ribs. The coracoid (fig. 1 106, e) in the Carinatx is an 
led hone more like that of Crocodiles than that of Oinosaurt ; 
^no fontanelle, and articulates at an acute angle uHth the scap- 
which it UMially remains distinct, ti takes an equal share 
latter in the fomutinn of the f;1enoid cavity for the head of 
lerus, and at its dialal end may overlap its fellow. In the 
It* the coracoid (fig. iiii) is, however, generally shorter .ind 
K Dinoaaurian-like, and may have a fontanelle, while its long 
■ dlbcr coincident or parallel with that of the adjacent portion 
p scapula, with which it is invariably anchylosed In the ndull. 
■Capula in Carinatc Birds ^fig. tio6, f) consists of a thin and 
ttw plate of bone, often <\tcnding backwards to a considerable 

tand without any suprascapula. Occasionally an additional 
xl is developed on the outer side of the scapula. The 
il crxb of both the scapula and coracoid -ire di\idcd into a 
9idal ar>d a clavicular procciu in this order. In tlie same order 
datide is nearly always well developed, and fuses with its fellow 
jnn the U-shaped furcula (fig. iio6,yiv); but in the Ratitie 

Ucs of ihc :^-- ^-r tt.. ■-. : 
■hu !ura.i»:-: .4r.- ':.' /:;- ■- - 
. : \i-i ihc rr.2r.j- ;":.■-■.- :-""-.7 
■.iih the f.r?: ...":-. i : . :•. 

..langeals: ;r.c :v~.:r..; ■ .-- .r. 

I.1W, In ex>::r_- < :.:::-.::-.- j ri- 

ire more or !e>s ■•::.■.: II:-. :j-.i 

■ .itssor Wi;in;>, ;■,-:■. :"..:.:::-- 

I yf phalanjrcals in :r.i.- :>-: 'iij:: ^,» 

■ hearing a ciaw : whiir four :'..:.■.:!:-.■> 

:■! the second dijiit. like Ar:/:.,- rr-:r- x. 

■y.--^ is the terminal phabn^i-al ;iro\:':tol 

_ farinates the third di^'it ha-; i';i:v ,i 

.■ claw. Among the Rniit.e there are 

K/ua ; but W//m'.v and t'asuariu.'i\\M\: 

■ lawed. There is usually an inler>|iaiv 

■ i and third metacarpals whii:li ni;iy l)e 

■ I lime. 

■clvis have been already hriefly alluded to. 

. hvays produced cunsideralily im liotli sides 

-1 some cases, as in the .'\pteryx, the anterior 

.•X : it articulates with the long sacrum, of 

..s been already notice^l^^he ilium arches 



over itic greater part of ihc aoetabulum, oF which, as ^itaAfi 
tioned, the cenUe is unassified, and has an exterrul aniaibri 
for ihe greai tiochanter of the femur termed the axti 
The ischium foniu. the hinder half of the inferior joft of Ifacj 
bulum, and is a moderately broad bar of bone directed i 
and backwards, or occasionally backwards (fig. ittz). 
Carinatic (Bg. ttos) it generally uaiics infcriorly with tlw tin 
which mcon^ the upper pan of the ilio-ischialic notch is < 
into a foramen; but iti tbe Kalitx (figs. 1107, iitz)lhae 
such union. In A'Ara alone, among existing Birds, tbe ischial 
in a ventral syiuphysis. 'Die pubis is generally a long ondj 

/, Publi:/. ]'«caB»l|iroc«<«(<lo.: d. A«eiatalan. (Alter Mank) [ 

bone running parallel to the ischiunt, and entering into the aniai 
part of the lower border of the acetabulum ; it freqiimtly gitw 
a pectineal prucess (fig. 1107)1 which is apparently homoli 
the preacetabtilar process of the pubis of the Otnith' 
sauria. The Ostrich (S/ntMo) is peculiar among hving 
having a iymphysis pubis ; while in Arc^trvpferyx alone arciiir 
pelvic bones separate. The femur is a short thick boncrilfci, 
head plictd at right angles to the shaft, as in certain Uij)UHaa( 
lis condyles arc large, and anlero-posieriorly elongated. A ;"*^ 
is frequently present, .ind may be double, llie fibula is al*»y»'4 
perfect discally, and may be completely anchylosed to the tw 
The latter, or tibio-Ur«us as it should be correctly termed, ''"H 
characteristic bone ; .ind is .tlways longer than the fciniir- W 
tnally this bone is expanded and produced into an anterior Cff^ 
process, like that of the Ihnosauria, which may extend abow Ot 
knee-joint; and the distal extremity (fig. 1108, a) has a uc^^ 
like surface, and consists of the astragalus of the tarsus. rhJchM 
been completely fused with the tibia. On the anterior siiitot * 



bove the Bstragaluii, there is frequently (as in fig. 
my bridge over the channel for the extensor tendons of 
Ik tarAO-nicUitarsus (fig. 1106, tm) immediately sue* 
bia : and consists proicinutlljr of the diMa) portion 
i, -with which the 

mc'tataisals of the 

arc usually com- Bl ^ MXi 

«1 to form a rod- 
lencrally tcmninal- 

dtsul pulley-like 
tbc Articulation of 
als. In the Pen- 
-cr, intcnuls exist 

three cnntponcDt 
le shaft ; and io 
orij two metatar- 
>lo the compound 
rcnerally the mc- 
jsal at tu distal 
t somewhat in ad- 
e other two (fig. 
The fomi of the 
■les of thi« liotK- 
icterislic of differ- 
if Birds. In cases 
aIIux, or first di)^i, 
, It is only the dis* 

of its mctatarisus 
[ached to the posterior aspect of the tarso-metatarsus. 
ifd has any liace of a fifth digit, and tlic number of 
raiy from two {0«lrich) lo four (Tanots). In four-ioed 
)talangeal& generally number a, 3, 4, 5, reckoning from 
lliu) to the fcMinh digit. 

ue in an arithmciical ratio of ihc phalangeals «f the toes, in 
rant ihc inner to the outer tide of the fool, obtains in almost 
I etiablei us readily (o dclert nhicTidiKil ■* supiirtHMrcI, wlicn 
Nir are not all present. VariDiiima uf diffcrvni ktn<U exist, 
(be number and dis{K»ition of tht iocs. In many Birds — 
'arrots— iheouteriiiosi tot is lumtd backwAids, so ihnt ilicre 
in front and Ino bt-Iiind : whilst in ihcTrogons the inner toe 
k with the hallux, and the ouicnrost one i^ tiimcd forwards, 
ain, the ouier toe is normally dlrrrtrd forwiinls, but r»n he 
rards at the will nf the uniiiul. 1 n the Swirih, on the other 
ur toes arc present, but iticy are all turned forwards. In 
-especially amongst the Anserine birds — ihc hallux is wholly 
udimeniai)-. In the Emeu. Cassowary-, Bustards, and other 
invariably absent, and the foot is ihree-tocd. In the 

poriMn <f ihc rigki libuyiiRai (a) uxt ltd tana. 

Ostrich both the hallux .tndthe second toe are wanting, undlllc 
silts simply of the third and fourth di^ts. 

Ill regnrd to their geological distribution it should be 
that the rcm>iins v( Birds are generally by no means so 
OfisiferouN OejHJ^its :i8 those of M.-iminals. This sc^irrit}' is p 
in part due to tlic cotnparativcl)' small sixe and fragile nanu 
boim of a large nunihcr of members of the class ; and ala 
C. Lycll h&s observed, to the circumstance that "the pa 
flight possessed by most birds would ensure them againM 
by numerous c:isualties to which quadrupeds arc cxi>ONd 
floods ; " so that, " if ihcy chance to be drowned, or to (fl 
swimming on water, it will scarcely ever happen that they 
submerged so as to become preserved in scdimentaiy 
since, from the lightness of the bones, the carcass wocW 
long afloat, and would be liable to be devoured by piedacet 
mals." To these con^idernttons must be added the ab$enc« < 
in the great majority of Birds, whereby we arc dt-privcd of 
which in the case of Mammak has thrown most imporui 
upon the nature and alUniiy of fossil forms. 

The earliest suggestion of the occurrence of Birds is 
by impressions of huge Sauropsidan feet (6g. 1109) fouod' 



Fig. iiag.— S*iiro|iHdaa f«oiprini, and iinpmslom oF rain-^ren ; Iran ibclnwi 
CooDKticui VkIIft. RtducML 

reputed Triune MOdstone of the Connecticut Valley in the 
States. These Impressions were exndcntly made either by ' 
podous Dinosaurs, or by Ratitc Birds ; and the oeciurcn« ' 
tain reptilian bones in the same deposits indicates thai at 
of them are probably of Dmosautian origin. The absence i 

y a Rrptflc! Of the extremely generalised nature o' 
pition is made in the next chapter. In ihc Cretaceous 
kth America we lind not only the Tcniarknhle toothed 
rcre already difTeremiated into the KAliie and Carinate 
5 but there were apparently others more nearly allied 
||>o. Bind-rvinains also occur rarely, and in a very 
^le, in the Upper Cretaceous of Europe. With the 
Rcr upon an Avion fauna of a dvcidctlly modem type ; 
rriod of the I-owcr Miocene the greater number of 
tden were wdl represented. 

Bresent lo a great eXI^nt in the dark as to the manner 
k branched oflT from the primitive Sauropi^idan stock ; 
t evident that the Dintxiuna arc tho?>c Reptiles most 
no Birds, and that the C>milhusauna arc totally out of 
^inttral line, — the cuiious rcscmbbncc which iht-y pre- 
jte Birds being Apparently solely due to their somewhat 


h the mode of origin of the Ratite and Carin.ite moditica- 
Inicture, we may quote from an admirable anicle by Pro- 
on, who observes that—" First of ;(1l w« find that while 
lessrd the teeth tliey had inherited fnmi their Keplitian 
remarkable and verydi&tinct types of the class had already 
rararce, and v^ must note iKal these two types are those 
I the present day, and even now divide the class into ihe 
rinato^ Furthenrtiorc, while the Kaiitc type {Htsptromii) 
ind of teeth, arrayed m ({Touves, which indicate (in Rep- 
\ low mnrpl>(ilo|;ical rank, the Carinaie type ilchthyorms) 
:h icelh wt in tuckets and shnwins a hi^'her de^eliipiiienl. 
hand, this early Carinatc type has venehne, whoic rom- 
lle biconcave form is equally e\ idencc of a rank unques- 
Ut tlw saddle-shaped vertcbrTt- of the cuntcmpomry- Ratite 

HHSU^^MWUn (^aaht^l ndsilion. kr' IV-iTnrn has hiwri 


the jaw was still variable. There is no reason to think that at ( 
any Reptile (with the exception of Pterodactyl es. which, as k 
been said, are certainly not in the line of Birds' ancestor) ha 
sternum. Hence it seenis almost impossible that the first B 
have had one ; that is to say, it must have been practically rf 
type. Professor Marsh has shown that there is good reason fa 
that the power of flight was gradually acquired by Birds, anc 
power would be associated the development of a keel to the st 
which the volant faculty so much depends. . . . Thus the Car 
would, from all we can see at present, appear to have been m 
the Ratite." After observing that embryological and distribuf 
afford support to this view, Professor Newton continues a 
" No doubt the difficulty presented by the biconcave vertebra: 
liest known representative of the Cannate type is a considerabl 
to the view just uken. But Professor Marsh has shown that ii 
cervical vertebra of Icktkyomis ' we catch nature in the aa \ 
of modifying one fonn of vertebra into another, for this singi 
in Ichtkyornis is in vertical section ' moderately convex, while tr 
it is strongly concave, thus presenting a.n^r approach to the ; 
articulation ; ' and he proceeds tagteinf out that this specialis 
occurs at the first bend of the nftl^nd, greatly facilitating n 
vertical plane, is 'mainly due originally to its predominance.' 
of the vertebnc would accordingly seem to be as much cone 
the mobility of the neck as is the fonn of the stemum with the 
flight. If, therefore, the development of the saddle shape be i 
tion of development, as well may be the outgrowth of a keel" '. 
sion, the Professor observes that the question must be rq^aid 
unsettled, although his own opinion is strongly in favour of i 
being the earlier type. 

On the other hand, Dr Gadow, in a communication of later 
eludes that the Ratitx were most probably descendants v& Bi 
formerly possessed the power of flight ; this view being said i 
ported by the structure of the wings, and the nature ofthefcatt 

In the following chapter a brief summary is given of 
divisions of Birds, with mention of those families known 
presented in a fossil state. It would, however, exceed the 
this work to give even the leading osteological characten 
families, since, owing to the great general similarity in the 
of all Carinate Birds, such characters could only be indicat 
introduction of a great mass of detail. 

It should also be observed thai the majority of writers 
three primary divisions of Birds as subclasses, and the : 
divisions as orders, with the proviso that such orders ar 
diflerent value from those of Reptiles. With the object ol 
this inequality the view of Professor Huxley, who has te 
primary divisions orders and the secondary ones suborders, 
followed in this work. 




Orders Saukur^c, Ratitx, and CarixaTjC. 

1. Sauaur^. — This exlinct order is represented only hy 
>trryx, arxl may be characterised by Uie metacarpob being 
Mr, and by the tutl being lonyer ihan the body, and not 
fiaunjt in a py^osiyle. 

M>i-V Akch-copiekvuiim!, — Anha^leryx, in(!luding birds of 
I the ^ize of the common Rook, is found in the tichographic 
BDites of Sotenhofen, near Pappenheim, in Bavniia, which nre 
C|vc9cntatn'e$ of the lower part of ihc El1gli^h Rimeridgc Cby. 
genus was first madeknon-n by the impression ofa single feather, 
tucfa Ibc late ProfesBOr H. von Meyer gave the name A. iUho- 
idea. Sutieequeitily the greater jKirt of a KkeEetoii, with im- 
tioas of the feathers of the wings and tail (fig. 1 1 1 o), was 
oned, vfhich Sir R. Owen named A, maanira ; while recently 
Kond skeleton has been found which some writers regard as 
eating a distinct species from the first. Of these two skeletons 
former is prescr^'ed in the British and the latter in the Berlin 
mm. Professor Carl Vagi, who first described thi; Berlin 
Einen, regarded Arehaopteryx as a Reptile : but there car be 
baitaiion in cUlssing it among the Birds. The jaws were fur- 
led with teeth ; the vertebKe were biconcave ; and there was a 
k>suficd sternum, of coi>^idcr.-iblc breadth, and probably pro- 
td with a carin.1. In the m.iniis the three metacarpals rcmamcd 
iDCt : and there were also three separate digits, each of which 
tcnnitutcd by a claw. In the pelm the three component 
cicnts exhibit the Reptilian character of retnatning distinct 
VghoQt life ; and it is thought probable that the ischia united 
I ventral symphysis. The distal portion of the fibula is placed 
ftont of the tibia ; and the metaursals were either separate or 
upetfectly united togcilier. The uil, again (fig. mo), 



differs from that of all other icnovn Birds in that it 
series of long vcrtcbrx:, gradually upering co the ei 
each of which carriesi a pair of feathers. From the 
impTcssions o( feathers in the region of the body, tt 
thought that only the wtn^s and tail had these appcr 
it is far moa- prolwiblc thai the feathers had fallen from 
as it \ay on the uld bea-shore, while tliiKC of the wings and i 




[y ' " M 


FIc. iiio.— ?aTtaribiiJidcion, HriiblinBraidMi(d'ih«taili«rt«(ihpi 

adhered to their attachments. In the figure of th« British Ml 
specimen of A. matrura (fig, riio) ihc head is not Hhown ; 
exists in an imperfect state of preservation in the &bb. T^l 
know more of the other Hirds of the Jumsiiir, the true leUtia 
of Anhaoputyx to exiitttng fonns cannot be determined. 

Orhbr II. Ratit.«. — 'ITie Ratiue, or Sliuthioua Birds, 
from the preceding order by ihe nnchylostii of the 
(when present) and the short tail, which may icnninaie b i 
pygoMyle. They are further characterised by the Kiemum 
1105, till) being devoid of a keel; by the long axes of 
adjacent portions of the scapula and coracoid being approxiB 
in the same line (fig. mi), or at least forinir^ an.i 
obtuse angle at their junction; by the wings being usdss 
flight ; and by the peculiar characters of the pelvis (figs, t loj 
1 1 11), which have been already mentioned. Tlicre are. 



tt CKatcological characters connected with the palatal aspect 

skull which cannot be noticed here ; and some other reatures 

been nientioncd in the preceding chapter, The iiia»i\'e Iwnes 

»lly filled with manxiw. In all forms the fibula remains 

dicunct from the tibia ; and the daul end of the latter frc- 

1*1 int.r—Ltli tall of tbc fftcimi rinlle met iiunum of Unftrtntii n/ptlit; from ibc 
laoMv «f Narlb Antnu. R(duui£ (, St^^HiU ; h, Huncrui e, CoiuukI;/, CUrkIc; 
n«a^ lAftcr Uanh.) 

(mly has no bridge over the extensor i(.>ncIon<t. In all existing 
Itas (he plumage prcKnts the remarkable peculiarity that the 
Irtlt of the feathers, instead of being connected with one another 
' booked barbutcs as is usually the case, are leniote and di»- 
nnected from one another, pre^niing some resemblance to 


order embraces ibe largest known members of the whole 


E^h> Uit.— L<A hitfaf l]w pdritot HiifrrmmU rrfali'ii fruni ilic Cicidimuiof Kotth 
Awwrica. Reouctd. Ltiunaain Ak. do;. (After Uarah.) 

In; anil from the scattered distribution of both its exi»tiiii){ and 
■bI rcpTesentatives is evidently an extremely ancient type. Its 
daboos to the Carinatic have been already alluded to in the prc- 
■diqg chapter. The order may lie divided into two scries, accord- 
iBg to the pnscocc or absctKc of teeth. 



Toothed Series. — In this extinct series teeth are proB 
the jaws. 

Suborder i. Odontolc*. — The type and only known i 
sentative of this suborder is Hesperomis, of the Cretaceous of I 
America. In this remarkable Bird (fig. 1 1 13) the jaws (fig. 1 
were provided with a series of sharp-pointed teeth, sunk in ai 
continuous groove ; but the anterior portion of the upper }n 

Kin- Illy— ^Su^Xov of Hnperfmiirttath: rrom the Craacmiu uf Nonh ABcrki 

About one-Knih natural >at. (After Hanh.) 

udentulous, and probably sheathed in a homy covering like th 
existing Birds. Various parts of the skeleton are representt 
figs, 1 1 1 1, 1 1 12, and II 17. 

in its whole skeletal organisation Hesperomis conforms strictly I 
cvisiinft Ratite type ; but there were four digits in the pes (all of i 
were directed forwards), and I'rofcssor Marsh believes that it » 
aquatic hiibits, and compares it to a swimming Ostrich. Accord' 
the description of the same authority, the tail consists of about t 
vertebnc, of which the last three or four are amalgamated to form 



there bcinj; at the >utnc ttiiM: clear indkation^ thiit the 
blc uf an up and down moveincnl in a veitiuil plnnc, thus 
ling iiioAcrvc aa a swiraminK- piddle or rudder. The vcr- 
ih« cervical and dorsal regions arc of the ordinarj' ornithic ly^e. 
wen powerfully consirmrlcd, and the feet were adapted to assist 
m r«pid motion (hruuKh the t^-aicr. 'Die known rcm.iins of ihc 
titi^rvntit rtgedis (fig. iiij) prove it to hiive bcvn of larger 
icn> than any of the »<|uatic members of the class u'lth which 
aci^uaimcd ai the present day. It appears to have^iood between 
d SIX feel high, and its inability to tly is fully compensated for 
nunicrous adaptations of its structure to a ivaicry life. Its lecth 
c it 10 hAVc been ratnivortius in its habits, and it probably lived 
ti-ihrs. A XMOiid ipctit-» of this genus occurs in the wtnc deuosits, 
1 IS knowTi as Hespfrirrmz imtupts ; but it wiLi orit;innUy rcjjarocd as 
daa*:iti^ ifi a distinct tfcnus, and lumcd Ltilomii. 

TooniLEis Series. — This scries, which includes the whole of the 
■anting forms, i« chnnicterised by the nbs«nce of teeth. 
SunnRitKR 3. ■■l-^rYORNmiB. — This suboidcT is rfprcscntcd by a 
Dg^ family, the jUpyomithida. of the Pleistocene of Madagascar. 
one lerK>wn genus, ^.pyornis, i.s characterised by the shortness 
die beak ; the smalt vingK ; the absence of a tibial bridge over 
extensor tendons ; and the )>Tcscncc of a hallux in the pes. 
tyjncal /£. maximitt appears to have atLiined dimensions 
ling those of the larKett »pecie» of Dtnomis {to be sboitl}* 
); and eggs have bcrn found in xs.socialion with the 
Dftcasunng fotirteen inches in diameter, and computed to be 
to capacity 10 three eggs of the 0^trich. At k-ast tno 
icr species of the same genus occur in the Madagascar 

SVBOILDF.R 3. APTEftVGES. — ^Thc members of the second suborder 
f this MTies arc confined to New Zealantl, and may all be included 
the family A^terypda, which is Jiow represented hy the Kiwiit 
Afttryx (fig. 1 1 14>. They arc distinguished from all other ex- 
ing DManben of the order by their extremely long and slender 
■k, which b adapted for probing the soft inaTsby ground which 
fre(|uent in xenrcb of worm.'« and other fond. Omitting montion 
peculiar cranial and ancral characters, it may )h: ul»crved that 
wing has a coin|uratively shun humerus, and not tnore than 
ungual phalai^cal. The tibia is furnished with a bony bridge 
the extensor tendons', and there is a hallux to the pes. The 
,lher% bavc no aftcrshafis. Tltc Kiwis are essentially nocturnal 
in iheir habits. Remains of the existing species of Aptcryx arc 
jfeanil fossil in the Recent and Pleistocene deposits of Nck- Zealand ; 
*hi)e some much larger hones from the same depnsiu have been 
tcsrnberl by the late Sir Julius von Haasi under the name of 
Hfgalaptrryx, whirb appcurs to have been a giant form closely 
CO tbc ousting genus. 


Suborder 4, Immakes. — This recently extinct groups 
Ust, is almost peculiar to New Zealand, and compmcs son 
largest known Birds. The beak (fig. 1115) is sbon ; the 
either very Hinall or totally wanting; the tibia has a d»ttli 
over the extensor tendons ; and in Kome instances (here I 
hallux in the pc-i. The characters of the skull and pehfb' 
nearest to those of the next siuhorder, and the featherx have) 
shafts. I'hia group iias k«en divided into the Dinomitkidt i 
Paiapterjgiday on account of the absence of the hallux in < 
former. Although this distinction has been doubted l>y 
wTitcrs, who consider tliat Dinornis had a hallujc, it appcan M 


rit|. \\\^—AKtryx mmf^itt. Hew EBdkad. 

a valid one. Apart, however, from this point, according to tlic 
Sir J. von Haait, the i'alapttry^da were provided witJi r 
menlary win){s, while in the Dinomithida: Lhoae appcnda^ci 1 
totally absenL Mr De VLs has described some bird-bono I 
the Pleistocene of Queensknd under the name of Diit»mii ^m 
iandia ; this Itein^ the only instance in which remains i>f ihitgn 
have been recorded elsewhere than in New Zealand. Sir ^ 1 
Haast proposes to divide the Palapttrygida into Piilaftrryx I 
£uiyapUryx^ and the Dinomithida itito Dinomit and MJt i NnA 

The first evidence of ihe existence »f this marvelloa* eroap wasafloc 
by a fratpncnt of the shaft of one of the bones of the leg Drought U 


Owen at a time when ihc cxiatcncc of s«ch huRP Birds wai 

basuspcctcd, nnd upon which oidcnce he founded ihc type 

Tlui the Mn.m, ns thnc Birds arc icnncd by the M.iories of 

^Ztalanrf, tuivc only lircn exicmtinated at a compaia lively recent 

i jmwr*! by ihe ociuirrnte of nearly mlirc ^tkclciuns with the skin 

trs Mill adhering: to them, x» iiifcll »■> hy fnijjTnent* of the ckk* 

ibcir oriinaal palc-gneen colour. The UrKCit tpcck's i» ff. 

of vhicb ihc loul height wa» about icn feet, the tibia 

a yard in length. Another species, D. fltp/iofttepus (fig. 

, ttn • fl» !■ wn if /*■'■■ I III I i/r/>l«->/V*>( : fmni ih* PltlMocin* oIKtw Zctluid. 
(kuily reiiufld. (Alter Ovio.) 

I), slihough not standing more than about six feet in height, was of 
■ more ina§sive construciion. the tuc-bonn almost rivallin)^ ihose of 
f«lepbani in si«r. The number of sjiedes desi-ribed i% very Inrife. 

JBORDKJi $. Megistanes. — The Emeus and Cassowaries are 

by certain stnicttirnl peculiarities in the base of the 

cranium ; by the motkrately long humerus ; the presence of 

one complete digit in the manus, which ■<: rnrnished ivith a 

; the absence of a ventral symphysis in the pube;s or ischia ; 


of the 

d by him 

iliOBCd Uttl 

ecu named 

u now by far 

.laeriscd by the 

(Im) long axes of 

■ fatmiDg at their 

! i» b; the upward 

>, 1 toa). In most 

'■me instances tbc^ 

. are modified into 

. HS obtaining anionic 

<1'U order, snii all such 

' more or less provisionAl. 

.'■(1 l)y MrP. I- SclattT 

M.lificAtions suggested by 

' ' 1. 

<>>ch are susceptible of beinfi 

- udraable to mention briefly 

Kii not yet been drtcrmincd. 

iti- mijorit}* should find a place 

'nx, of thf Upper Jurassic of 

■ .jly mcmioncfl, to Ik; coiisidcr- 

;' at all, In the Cretaceous of 

' u'u Craaiiavui, Laomis, PalxO' 

-II of which probably belong to 

■ 1 the oldest known bird-rcnjains 

,-ii<l. and have tjLvn named Kna- 

i*-lc that ihcy may indicate more 

'.iTiebra; have more or less flai- 

< Tdtanua, in which the fusion o f the 

,nie, 18 compared to ih,ii of i 

Lit of Jchtfiyprnit, and it is hi) 


the want of a bridge over the extensor tendons in the tibii 
the suppression of the hallux. The family Dromaida is 
sented by the existing Emeu {Dromaus) of Australia, and al 
fossil species in the Pleistocene of the same country. /Vm 
from the latter deposits, is an extinct genus referable to this i 
The characters of the pelvis of the existing genus are shown 
1107. The Casuariida, now characteristic of the Austia 
region, have not hitherto been definitely recorded in a fossil 
although it is possible that a phalangeal from the Indian Sii 
may be referable to this family. 

Suborder 6. Rhea. — The Rfuida, which alone coostitub 
group, and are confined to South America, differ from the 
stanes by the structure of the palate, as well as by the long* 
merus, the presence of three digits (of which two are clawed) i 
manus, by the ventral union of the ischia, and the absence 
aftershaft to the feathers. Remains of Rfua, which are refen 
the existing species, occur in the Pleistocene cave-deposits of 1 

Suborder 7. Struthiones. — The family Struthionida i 
sole representative of this suborder ; the only existing species 
the Ostrich {Struthw eame/us), which is now confined to Afric 
Arabia, although it formerly ranged into Persia, and probaU 
to Baluchistan and the north-west frontier of India. In ad 
to the characters of the palate, Struthio differs from RMta I 
union of the pubes in a ventral symphysis, and also by the su 
sion of the second digit of the pes, in consequence of whit 
distal end of the tarso-metatarsus has but two trochlex. This 
is represented in the Pliocene Siwaliks of India, and also i 
Lower Pliocene of the Isle of Samos, in the Turkish archip 
by remains referred to two species. These forms, which m 
specifically the same, appear closely allied to the existing O 
An egg, from Tertiary beds near Gallipoli — the ancient Chers 
— described under the name of Struthiolitkus, probably beloi 
the existing genus, and very likely to the species occurring atS 
These fossil forms point to the conclusion that the original ho 
the genus was probably in Asia. 

SuiiORDER 8. (Iastgrnithes. — The Gastornilkida, who: 
mains occur in the Lower Eocene of Europe, were large 
which may probably be classed with the Ratitse, and are appa 
entitled to distinct sul>ordinal rank. Their tibia agrees with tl 
the Aptoryges and Immanes in having a bony bridge overtl 
tensor tendons, but makes a curious approximation in shape ti 
of certain members of the Carinate suborder Anseres. The 
were somewhat larger than in the Ostrich ; and the cranium, 
is estimated to have been fifteen inches in length, had the al 
margins of the jaws serrated, as in the genus Odontoptery: 



CMiuins of Gast&mii have been wcortlcd from Mcudoii, 

from Rhcim^. and fronj Croydon, and have been referred 

The huge Dtatfyma, from the I-ower Eocene of 

DCficsu appear* 10 be cIcmcIv allied lu, if not identical with. 

Bird-bones from the Tertiary of South America, cle- 

y Dr Moreno under the name of Afexm/'riornisy and com- 

tlw AnMTies, prolulily indicate a ineinber of this group, 

^ are fully as large as the ccMTcsponding Itone^i of the 


I ma; be noticed an imperfect cranium froni the London 
bcribed by Sir K. Owen as. Dasfimis, and regarded by him 
^ng to a Katitc Bird. And ii may be also mentioned that 
ifect limh-bone from ihei-e deposits, which haK been named 
1^, may perhaps belong to tht; sutiie genus. 

■ III. CarinaTjR. — The third order, which is now by far 
i numerously represented, is gi-nerally characterised by the 
I of a median Iteel to the sternum, and by the long axes of 
Bent portions of the scapula and coracoid forming »t their 

tan acute or slightly obtuse angle, as well as by the upward 
\ of the ischium towards the ilium (fig. iioi). In most 
wings are adapted for flight, but in some instances they 
JDome atrophied, while in others they are modified into 

■ organs. 

\ is uill coniiiderable diversity of views obtaining among 
ktigists a5 to the clauification of this order, and all such 
I must consequently be regarded an more or less provisional, 
jresent work the claMification adopted by Mr P. I„ Scl;itcr 

rl in the main, although certain modificilions suggested by 
Newton have been tncorpomicd. 
ft noticing those fossil fonns which are susceptible of being 
|n definite groups it will lie advisable 10 mention briefly 
tetnains of which the affinity has not yet been determined, 
^ tt is probable that at leaAt the majority sliould fmd a place 
present order. As to Lacpttryx, of the Upper Jurassic of 
[zDcrtca, there appears, as already mentioned, to be consider- 
|lbt whether it is really Avian at all. In the Cretaceous of 
ic country we have Afaiornis, Graathvut, Laornis, Palafi- 
|tnd Telmatornii : many or all of which probably belong to 
k suborder. In England the oldest known bird-remains 
B the Cambridge (Jrccasand, and h,ivc been named Ena- 
.although it is quite probable that they may indicate more 
le genu)'. Some of these vcrtebrat have more or less flat- 
Jntra ; while the tarso-metaursus, in which the fusion of the 
lent elements is incomplete, is compared 10 that of the cxist- 
but and also to that of iihthyomis^ and it is highly prol>- 



able that these Birds ircrc allied lo ihc latter gmus. TV I 
from the Cretnceous of Kur[>|x; desicribed u Paioffrnis, OJN^I 
and Crctomis, belong to Oniilliosauriji. 

In the Tertiary we have Euptrrormi and Remianas (ma 
liOncT Eocene of Rheims; while tlic Upper Eocene (I^wctC 



I. OooMTORM-r- — The one suWdcr which is the 

representative of ihis series contains the family Jchlhy- 

from the North American Crct^iccous, of which Ickthy- 

only definitely known genus although it is highly prob- 

1 Ap^t^mii, and some of the other Cretaceous fonn* alrcady 

1, may alsu belong to this or an allied family. Tlie ireth 

nit (fig. tti7, d) arc comparatively large, and siv set in 

Itcls ; while the centra of the vertebrse are amiiliiccHlous. 

about twenty teeth in each jaw, which arc directed ob- 

Hwckwords. The rami of ihe mandible were but loosely 

and it IS probable that the jaws were not sheathed in horn. 

lU of the typical /. dis/>tir (f\g. 11 16) was about equal in 

Rcxdc -pigeon; and in nil essential features of its organisa- 

I nnM at lka_ oundibte of UMf—itft. illjihlly mbcscd ; 6, Ua. of Uti' 
mnb tu*ar^ tw ; e, /, Anlcrior »iiJ bilerat iu|xcu of ccrriijil *«r(«4jrii vf 
I utml VIM ; i, TAMh of Haftranit^ vmos rumiml tijc^ (Afici Mjifih'> 

genus conforms so exactly with the existing Carinalc lypc, 

appears every reason for including it in the same order, 

in following Professor Marsh's view of placing it, together 

\sf>ervrm$, in a separate order under the name of Odomor- 

ILBS8 SufCS. — In this series, which eompriises all existing 
no ueth are ever functionally developed, although germs 
the young of one group. 

>EX 2. CuVKTUKl.^'rhc Tinamous {Tinamtii, &c) whirh 

suborder, show more signs of .iftinity in the stniclure 

and skull to the KatitEC, than ia cxhibued by any 


Other group of this order. They are confined to the New H 
and arc represented in a fossil state by remains of existing 9 
of Crypttirus, Notkura, Ti'namus, and JtAymckatus, in the PldM 
cave-deposits of Brazil 

Suborder 3. Impennes. — The Penguins {Aftenodyks, te 
the Antarctic regions form a peculiarly interesting and weEtdd 
group of Birds, in which the wings are modified for swimming 
the component bones of the tarso-metatarsus are separated byi 
ties. It has recently been proposed that the Impennes shodd^ 
a primary group of equivalent rank with the Caiinatx^ uwkrl 
name of Eupodomithes. Unfortunately scarcely anything is bl 
of their palseontological history, the only fossil type being li 
eudypUs from the Tertiary of New Zealand. 

Suborder 4. Tobinares. — The Petrels, or Proallaraia, 
the only family of this group. The only known fossil repRM 
tives are members of the existing genus Puffinus (Sbeaimter), ri 
have been recorded from the Lower Miocene of Allier, in Fn 
and also from the Miocene of the United States. 

Suborder 5. pyfioPODES. — According to the opinion <rf I 
fessor Newton this and the two following groups should be i^ 
merely as .sections of a single suborder, but since no name bash 
proposed for this larger group the three divisions are retained 
the Pygopodes, the A/cida, or Auks, include the Great Auk fy 
impennis) of the Arctic regions, which now j^)pears to be totally 
tinct, but of which the remains are found abundantly in the ) 
and other superficial deposits of northern Europe. Remain 
ferred to the genus Uria (Guillemots) are fotmd in the Upper I 
cenc of Italy ; and Guillemots also occur in the Tertiary of 
United States, where they have been described iind«' the nam 
Catarracies. In the Colymbida^ which includes the Grebes : 
Divers, remains of the Red-throated Diver {^Colymius ^amlii\ 
found in the Pleistocene deposits of Mundesley, in NoTfialk;v 
the extinct Colymboides of the Ixiwer Miocene of Allier appev 
be an allied form. 

SuuoKiiER 6. Gavia:. — Of the Zani& (Gulls and Terns) a spe 
of Lams occurs in the Allier Miocene ; while Hydromis of the 
ter do|>osits may probably be referred to the same family, 
undetermined genus from the London Clay may perhaps be 
referable to the present group. 

SuBOKDKR 7. LiMicoi,.*;, — The Limicoix are somewhat a 
dantly represented in Tertiary deposits ; the subaquatic habil 
many of its members being probably conducive to the prescrvi 
of their remains. In the family Seolopadda the genus Nutik 
(Curlew) is recorded from the Middle Miocene of Gers, in Fn 
and the Pliocene of Italy ; Limosa (Godwit) occurs in the U 

I iu iPUAuu (.uiN.pi mmA\ m u 

locnic of WcMphalia; and a species of Himanlofm (Still) 
in the Allier Miocene. In the fAmily CharaJriido: (I'loven) 
erf the type geniu CfiaraJritii occur* in llie Upper Eocene 
tdo ; while the genus CamasctJut {with uhich DolUhoptirut 
probably ulcnivoJ) is known from the Lower Miocene beds 
tn, in France. 

LUER 8, AixcroRiDrA — Tbc Alcctoridcs form a somewhat 
d group, which is token by Mr Sctatet to include thv 
although l*rofcs!>or Ncwion regards the LttKr as more 
Died to the CiaviiE and Limicolx. The family Gruida, or 
}• rcprcicnted by the type genus Grus in thv Plei»tocenc of 
[India, and the United States, and also in tht Lower Pliocene 
fbeds of Ureece. and the Miocvnc of Allier. Allied extinct 

f! Paiaogmi of (he Eocene of Italy, and AMornis of thai of 
The OlidtJa uk represented by a. species of Ilu^tard 
the Allirr Miocene. 
|l>eR 9. FuucARLC. — This suborder comprises the Rails, 
{jlstcr-hens, etc; all of which are included in the single 
utUi^a, oikI arc of more or less aquatic habiU. Birds re- 
b the type gcnuN Kallus (Kail) o<cur in Uie Monlmarlre 
jthe Miocene of Alltcr and Gcrs, and the Italian IMIocene. 
i of Gaiiinu/a (Water-ht-n) are recorded from the Pleisto- 
Bs of Brazil and Queensland ; in both of which depOMls 
I with othcn referred to Porpkyrio (I'urplc IValer-hcn)— a 
bw widely distributed over the warmer regions of the globe, 
■let species of Coot {I-'Mtiea) has also been described from 
lensland Pleistocene. JVoiornii, which occurs in the i'leisto- 
[ New Zx'aland and was also found living sonic years ago, 
^ Rail allied to ihe Au.'UtT^iIian Trilxmyx ; while ^//(?r*r«, 
ia very brge form from the Aanit: dtpo»its totally incapable 
^^niore nearly related to the existing Otydr^mus of New 



(Pheasants, Turkeys, etc.), and TetraomiUe (Grouse). The 
together with the following group of Columbas were 
bracketed together under the name of Rafiores, and it i 
means certain that the departure from this arrangement is ) 
able one. The skull (fig. 1 1 03, p. 1 2 1 o) has peculiar palatal 
a sharp curved beak, and a recurved process to the an^ 
mandible. Many of the genera of Gallins (especially ibi 
are characterised hv the presence of one or more strong bo 
on the inner side of the tarso-m 
(fig. 1 1 18). The first two familit 
present unknown in a fossil condii 
the Fhasianida, however, the typif 
Phasianus occurs in the Allier a 
Miocene deposits, and also in th 
Pliocene of Pikermi ; FratueUiM 
colin) is represented by remains 
ing species in the Pleistocene of ! 
India (fig. 11 18); CtHWrwix (Quai 
Montmartre Eocene gypstmi; thi 
Falaortyx in both the latter depc 
the Is^re and Allier Miocene b 
soK:alled Palat^erdix of the ] 
Miocene being probably identtca 

Muliu. -.PoMtrior;/, Anterior ^ spCCicS of GalluS, SOmewhat Ul 

the existing Indian G. Sonneratiy is 
the Pikermi Pliocene. From the Miocene of the United 
Turkey {Meleagris antiqud) has been recorded, and is desc 
equal in size to the living species now characteristic of j 
In the TetraonidiB remains of the living Capercaillie (7>j 
gallus) occur in the Norfolk Forest-bed ; while an extinci 
of the same genus has been described from the Upper E 
Languedoc. Remains of the existing Willow Grouse ( 
albus) are found in the Pleistocene of Westphalia. 

Suborder i i. Columb^. — This group is taken to inc! 
existing Sand^rouse (Pteroclida), and the Pigeons (Cobtmbi 
the first-named family a species of the type genus Pterxles I 
described from the Allier Miocene. The Columbida are k: 
a species referred to Columba from the last-named deposits 
as by another provisionally referred to the same genus i 
Pleistocene of Rodriguez. Of especial interest is a tarso-mi 
from the Pleistocene of Queensland described by Mr De V 
the name of Progoura, and regarded as indicating a bird ; 
the Crowned- Pigeons {Got/ra) of New Guinea. Gain 
some signs of affinities to the Phastanida, and these reser 
are said to be more marked in Progoura, which is do 

Fig. \\ii.—FrancaliiaH ffniii- 
eeriaMHi, The left tarso-meta' 
taraiu ; from iht Pleistocene of 



e existing forms. Here also may b« placed Ihe now 
mil)- Didida, represented by the Dodo {Didus inrptm) of 
and ihe Solitaire {Pesopkaps soiilaha) of Kodrifju^^. 

: two stnguW birds, the I>odo formcrlj* inhabited Mauritius 
Dumbrn, but the last record of its octuncnce dates from the 
It w-A» a Urge and hcav>- bird [tig. 1 1 19), bi^grr than a swan, 
eiy unlike (be niKcons in ycncial appearance. The win^s were 
iiy uid compfeiely useless ai organs of flight The Ick* were 

slotu, the feet had four too each, and (he tail waa cxircmely 

I «f iki Dado (DiJMi imftmt), naarrd. (AA*r Owtc. 

irrying, like the wings, a tuft of soft plumes. The beak <un- 
\<k any of the Coiui^a except the liitic IMdunculut strigiros- 
'f arched tow.trds the end, and thtr upper jaw had a strongly- 
bpcx, not unlrkc Ihat oif a bird of pTwy. The froninl region of 
{was ureally elevated and lumld, from the CKces^ive development 
jtr cavitks betvfccn the two tables of the skull, and the actual 
|c was vcrj- imall in propottion to the siie of the cranium. 1q 
tpccts allied to the Dodo, and, like it, incapable of flight, wgu 
aire, of which the last recorded Hppcarancc was in the yxar 
fce Solitaire bad longer legs and neck ihan the Dodo, the bill 
ftiTonKly arched, its fonhcad flatter, and thecc was developed 



upnn tlic rndial tide of the mcucarpus an extraordinaire spheric 
like mass of bone, ahnui as Urge as a rauskvt-b^ll. and with a i ^, 
surface. This sinj^ulat callosity is much more developed ia] 
indiviclunls— supposed lo be males— Ihan in oihcrs, which ret 
sume to be females; it was duubile<i« covered durinte life \n , 
tniGxtunenc, and lecin^ tn have been used as an crfTenxtiT 
Bolli these Birds arc known lo us by nearly entire skeleiunt 
reccnily from the islands which they inhabited ; and of the 
have also a few reni.iins belonging to entire specimens once j_ 
our museums, which were unRiriunately allowed to (all into i 
A{>[ureiU ignorance of Iheir pricelcis value. 

Remnins of exiting species of several genera of Caf* 
found in the l*lci»tocene cave-deposits of Bradl. 

Suborder i j. Anskkes. — The Anscrcs, or Goose-like 
a well-defined subwdinal group, characteriwd by peculiar 
ill the palatal region of tfw? skull, and hy the perfectly 
the be&k being generally broad and spAtulati, an<l the anf 
mandible with a recurved pruccss (fig. iiao). All the 
members of this group arc referred to the family Amal 
is, however, Nplii up into several subfamilies. A peculiar 
is represented by the living Cena^stT, of Australia; allied 
is the much larger extinct Cnfmiomis, of the Pleistocene i 

FSi nM.— Skull «fSpui-wu>K«dOo«M</'ltai'T/ivrwfiuK{>rwMli^ K«4au<- 

Zcalniid, which was quite incapable of flight. In the sabfN 
Afiserime, remains of the Grey-!^ Goose {Anitr amnta) o 
in the European F]c;i.stac«nc I'hc Cfgrtina:, or Swans, aic fl 
senlcd by the CKtiiict Cji^ur Fa/contri, from the Pleistocene C 
deposits of Malta. In the Anatina (Ducks) an extinct hpMil 
Treeduclc {UeHdraty^a) is recorded from the Pleistocene 
Queeitftland. Keuiaitis of the Wild-duck iyAnas ^oscas) oota 
the Pleistocene of Kurope ; A. afav^ and A. tygni/ormis are ft 
in the Middle Miocene of Bavaria, the tatter species being d 
as Urge as a Swart ; A. cmingtrtsis, from the Upper Mioccn 
SwiUerland ; A. lignifila, from the Middle Miocene of Italy i 
A. Blatahardi, from the Allicr Miocene. Remains of the Sboi 



^ia eJyftatJt) have boen found in the Norfolk Foresl- 

the /^ligu/itur, the type genus J-ii/igu/a (I'ochard) is 

from ihc Upper Pliocene of the Val d'Amo, in Italy, 

(Whiic-cycd duck), from the I*)eisioecne of Queens- 

in ihe Mergina. it is prohable that Mergvs {Merganser) 

itcd in the Pliotvnc SiMnliks of India. The eitinct 

of the luli-in Miocene, nUo belongs to this faniily, 

its precise ixisitiun is unceriAin. 

>KR 13. Oi>ONToiTRRVcKs. — The Odontofttry^dir, tqire^ 
H>f Odfitt/ffiteryx, of the London Clay, appear to indicate a 
su1>ordinal group, whit'h miiy he pru\i:iionally placed here. 
sint^Lir bird the alveolar raari;ins of both jaws arc furnished 
\toa(h-like serration!! (fig. mi) wliich differ from true teeth 
■ctoally piins of the osseous substance of the jaw itself, 
3y aigree with those found in the Chelonian genera ffar- 
Baiagur. The)' arc of triangular or eompre»sed conical 


( «^^,-«^t— 'i-'*"^ 

- Stnll of l>dtnltfUrjx Miatitui, found ; rram ihc Undon Clark 

•n, and of two sizes. Upon the whole, Odcniopteryx would 
pear to be tnost nearly allied to the Anaiida, but the serration of 
Javs ii an entirely unique character, unknown in any existing 

SvBOKDFJt 14. PAlJtMEnR£. — Of this gTOUp, comprehending only 
k AmcricAn Screamers {C^twitt, &c), no fossil representatives are 

Bt;w>itDElt 15. Olx>KTWJl/D«s,R. — Tbc only family of this group 
the PkteHiaifttridit, or Flamingos, which arc exceedingly long- 
tbed waders, disiingui^cd hy a pec\iliar downward bend of the 
(fc, and presenting characters conncciins them on the one hand 
b the .'\n5crc5, and on the other with the Herodioncs. The 
Iting gmtn Phtemcopterut is found in the Allier Miorenc ; while 
the «ame bedsL, ns wt;]| ait in tlie equivnient deposits of Ihe 
venoe basin, and also in the somewhat higher strata of Stein- 
B, in Bavaria, oorurs the Remis Palirlodtii^ which, 
apparently allied to PhamcfpUrtis, presenLa aomc afEntty to 


the LimicolEe, and also shows one osteolc^cal feature 1 
occurring among the Py^podes in Potties (Grebes) and 
(Divers). Rlomis, from Uie Lower Miocene of Ronxon, 
to be also allied to the Flamingos; while Agtwpterus, 
Upper Eocene of Montmartre, may perhaps be also inc 
the present group. 

Suborder 16. Herodiones. — This suborder includes the j 
leida, or Spoonbills and Ibises; the Ciconiida, or Storks; 
Ardeida, or Herons ; all of which are waders. The /Vis 
represented in past epochs by an extinct species oi Ibii (/j 
from the Allier and Steinheim Miocene ; while the existing . 
Black-headed Ibis (/ melanocephala) has left its remains (fig. il 
p. 1217) in the Pleistocene cave-deposits of southern Indu. 
other existing species of this genus occurs in the cave-de 
Brazil. In the Ciconiida, an extinct species of the AfncaB 1 
Oriental genus Leptoptilus (Argala), which includes the 
Adjutant Stork of India, is found in the Pliocene Siwaliks of^ 
latter country, and another in the Middle Miocene of 
while an undetermined Ciconioid, from the Pikenni beds 
Attica, may possibly belong to the same genus. Part of a 
tarsus, from the I'ertiary of Argentina, indicates a bird douUe ' 
size of the Pampean Stork, and has been made the type of I 
genus Palaodconia. The Indian Siwaltks have also yielded 1 
of another giant Stork, of which the genus has not yet been 1 
mined. In the Pleistocene of Queensland there occurs an 
species of Xenorhymhus. In the Ardeida the type genus At 
(Heron) is represented in the Bavarian Miocene by a species (i 
similis) apparently closely allied to, but rather stouter than, dj 
common Heron (A. cirured); and remains of the same genus ill 
occur in the Miocene of Allier and Gers. The Night-Heid 
{NyctUorax) are known by an extinct species in the Pleistocem ■ 
posits of the Island of Rodriguez. Finally, certain remains M 
the London Clay may possibly indicate that this family dates frd 
that epoch. 

Suborder 17. Steganopodes. — In the Steganopodes ire b 
eluded a number of web-footed Birds, such as the Darters {PhHid 
Cormorants {Phalaerocoracida), Albatrosses and Frigate-Birds (A 
gatidie), and the Pelicans (Pe/icamdie), some of which are regadi 
as more or less closely allied to the Gavise, while it is su^eslf 
that there may also be a connection between this group and A 
Accipitres. The Plotida are only known in a fossil state by 
species of the one genus Plotus, from the Pleistocene of Quetn 
land. In the Phalacrocoraadte we find the t>'pe genus /*Ad 
crocorax {Graeulus or Cormoranus) in the Eocene of MontmaiV 
the Allier Miocene, the Pliocene of the United States, and pro 



tn the Indian Siwnliks ; remains of the <!xisting Corirorant 

9) being found in the Norfolk Forest-bed. vWj (Gannct) 

the Miocene beds of Ctriorado and of Roruon (Puy-en- 

while Ptlagomis, of the Allier Miocene, ts pro-isionally 

[in this family, tn the I^gatida rcnuins of a Diomedia, 

closely Allied to the Albatrosses of the Soulliem seas, 

described from beds at the top of the Suffolk Cray ; 

is considered probable that Arf^'lhrmt, of the London 

ites the existence of this faniily in the Lower Eocene. 

PeUeamda remains of true Pelicans {Ptirtamn) occur 

Miocene of AUier and Bavaria, as wet! as in the Indian 

iCR 18. AtciriTHES. — The Acdpitrcs, or Diurnal Birds 
pivjr, are characterised by their curved beak (fig. 1123, b), the 
encc of a circle of feathers round the eye, and the powerful 
pf of the foot (fig. 1 132, a), as well as by many osteological 
tHCB, utd especially the nearly straight line formed by the three 



[V%. iictL— 4. r«Mortlw Pcrtcrinc Fkkoij ; u, Head of Brannl. KoJuiel. 

trochlcjc of the tano- metatarsus (fig. 1123), and the ab- 
[of a bony bridge over the extensor tendons at the distal 
of the tibia. The probability of this group being related 
Sl^janopodes has been already noticed. The Cathartida, 
acrican Vultures, are represented by existing species of CaHt- 
D and Gy/^rtkus in the Pleistocene of the Brazilian caves. It 
also bc«i considered that this group is represented in Europe 
ijtkomis Tfuttarimus, of the London Clay; an opinion which, if 
ifinned, will be of considerable interest from a distril)utional 
H of i-iew. The peculiar SerfKntariida, ax Secretary Vultures, 
Africa, are known by a species of the one existing genus Ser- 
tarimt frcwn the Allicr Miocene. The Fakonida include all the 
■ining genera, which arc grouped in several subfamilies. Of 
le the VMlturina, or tnic Vultures, are represented in the Plcis- 
»e breccia of Sardinia by remains of the type genus Vutttir; 
le those of the existing Afro-Indian Neophron jvrctwpierus arc 
OU U. K 



recorded from the et^uivnlent cave-deposits of southern 
Ihe other groups w« havi; evidence of a species of cither . 

Circus {fig. 1123) from the 
deposits, Uie %uri: being gim 
to show the form of the distal cad] 
tarso-tD4--tat3isu<i so characteristiei 
suborder. Mthut is rcoonlcd 
Allier Miocene ; and faim fn 
Montinartie Eocene. Species 
to AquUa are oientioned botll 
Miocene of Allier and Gen, 
Sardinian Pleistocene ; while 
is recorded from Gers. Of 
genera Palaotuirax, from the 
Miocene, i& icgarded as being 
Agttifa: while Ptilaodmf, of the Montmarirc Eocene, bi 
as showing al^nity to the Utuutards {Butta), and the 
dion) ; TeraCHJ, from the Ronzon Miocene, being an 
known form. The largest knuwn member of this sot 
Harpagornis, from ihe Pleistocene of New Zealand, whS 
apparently allied to Cirtus. Finally, several existing v^tom 
Falwmda are recorded from the Plcixioccne of the Bod 

SuDORDEK 19. Stkiges. — The Striges, or Owls, were fonn 

Fig. I ri3.~(?)jf/'A'iu or C'riBj, 
«p. 1li*di>Ul)i»iri>rihcleft iirio- 
(n<uianusj ftvin tha uiifrior (a) 
Hid iKHWriar (u) aipccU; fram loe 
PlilMix*!!* cf MAaJrai 


B '^^'^^^ ^ A 

PIf. III4--1. FoM crilH Loncwnd Owl (OMa v^ifritY, m, Hc«I vTibc 1 

grouped with the Accipitres, but are now regarded u being pr 
■bly more nearly allied to tlie Parrots. Comparatively few (6 
forms are known ; but in the Atienida, or Eagle-OwU, wc htw 



;le-Owl {Suh ignavHt) m the Norfolk Forest-bed, and 
ng Indian B. toromaadui in ihc Pleistocene of Madras; 
genus is also recorded from the Atlicr Miocmc, and is 
ed in th« Eocene of the United States by B. Uptasteus, 
about iwo-thirds the si^e of ihc existing B. vir^niaHus 
otHting the same regions J'he existing Ceylon Fish- 
bifia ^ ccyfotumii) occurs in the I'lei^locenc of Madras, and 
KuropettTi Snowy Owl {/v'yetea seandiaea) in ihat of West- 
In t}ic Strigidfe, or true Owls, remains of an extinct species 
re havK been described from Ihc Pleistocene of llie Island 
rigucji 1 while bones from the Miocene of Allicr and Gers 
en refeTted to S/rix. 

iDER ao. PsiTTACl. — The Parrots, Cockatoos, and their 
hicb constitute this suborder, are now confined to the 
regions of the globe, and are remarkable for the presence 
ge-joim at the base of the strongly-curved cranial portion of 
k, whereby the upper jaw can be moved upon the cranium 
ns is shown in fig. 1135, a. All ihe genera are of scansorial 

.— RtcbtlutnlaiyMrt (f ibediulUA) And oT Ibr M* i»» Cd) of/'iiff-fm fttkt t ia. 
■i, ^VkKOMUni):', Seoond: (. T^iidi i/, rourthdlslt- (Aficr Uluitlianl.} 

U)d (he foot had a hallux (Jig. 1125, b). In the PstUaddtr^ 

Parrots, remains from the Altier Miocene have been referred 
typical .\fri(:an genus PsUtacus, but that term must W used 

ler senw: than the one in which it is employed by the students 
t Ornithology. LopkapsiHaats is an extinct genus from ihc 

cene of Rodriguez. Remains of the genus Ntsior, peculiar 
ZeaUnd, occur in the dcpoi>its of that country which yield 
pmV, and probably belong to existing sjjecies. The American 
rs axe represented by species of Ara in the Brazilian cave- 

ts. In the Palaomithidit, which includes the Lories and 
lets, an extinct species of the existing .\Xrican and Oriental 
J'aMomis occurs in the Pleistocene of Rodriguez, which has 

eldcd the extir>cl Ncnvpsittucui. The Stringopidti of New 

* AmKuieJ from Ktfufa, 


Zealand, and the Ctuatutda (Cockatoos) of Australia, have 
been recorded in a fossil state. 

Suborder 21. Picari*. — The Piearia are a somewhat 
geneous group of Birds, of which it will be unnecessaiy to 1 
all the families, since only a few are definitely in a fossil 00 
To the African Musophagida, or Plaintain-cutters, it is thoq 
extinct Neeromis, from the Miocene of Gers, may possibly 
The remarkable Leptosomatida^ of Madagascar, which am 
Coraciida with the Cuculida, are represented by a spedes 
type genus Leptosoma in the AUier Miocene. The latter < 
have also yielded a species of Tro^n, the type of the &m 
ganidcB. Limnatamis, of the same beds, is referred to the V 
or Hoopoes ; while it is considered that the Upper Eocene i 
nis may belong to the Buarotida, or Hombills, of the El 
and Oriental regions. The Aicedinida, or King-fishers, ar 
sented in the London Clay by Hal^omis ; while in the 
(Woodpeckers) we have the existing genus Piaa in the 
Miocene of Isfere and the Lower Miocene of Allier, and the 
Uintomis in the Eocene of Wyoming, Finally, the Cypu 
Swifts, are known to have existed since the Allier MiocerU 
we find a species of the type genus Cypselus closely allied to 

Suborder 22. Passeres. — Of the Passeres, the last ai 
highly organised group of the class, an enormous number ( 
genera and species are known ; but from the comparative 
size of the majority of species, and the difBculty of disdnj 
even genera by fragmentary bones, scarcely anything is ki 
their palaeontological history. To the Alaudida (Larks) h 
provisionally referred Protornis, from the Lower Eocene <rf 
in Switzerland; and Aiauda is recorded from the Upper 1 
of Italy. In the Corvida (Crows) the type genus Corvus h 
described from the Allier Miocene. In the Pleistocene of R( 
the extinct Necropsar is a Starling {Sturnida) closely allied 
pied and crested Fre^lopus of Reunion, which also appears 
recently become extinct Among the FringUlida (Finches 
and Passer are provisionally recorded from the Allier Miocer 
to this family may perhaps be referred the extinct genus Pal 
from the Upper Eocene of Colorado. Finally, we have ; 
sentative of the Zanii'da, or Shrikes, in a species of Lani 
the Allier Miocene ; while the Sittida (Nuthatches) date th< 
ence at least from the fossil Sitta of the Montmartre Eocc 
are also represented by a species in the Upper Pliocene < 
Lastly, the cave-deposits of Brazil have yielded remains of 
existing forms of Passerines, among which it will suffice to 1 
a species of Swallow {Hirundo). 




IKS <>V. B.)— '■ UebCT Ardurofittfyx." ' PaL AbhandL.' vol ii.. 
an. 3 (1884). 
kviES C\V.>— "On some Fowi! Biid-Remaim from the Siwalik 
Hills. 'Geological Maguine,' dec ii., vol. vii. (1880). 
rAKDS f A. MiLNK-;. — " R^herchcs .Anatomiques ei Pal^ontolo- 
riquc» pour scnir i ITiistoire drs Oiscauic fossilcs dc la France." 
,>»ri. (1867-77). 

7irTMR.R [A.),ind Newton (E.)— "The Extinct Uirds of Kodri- 
fues." 'Trantit of Venus Expedition.' 

(J, VON). — "On Harpugornii, an Extinct Genus of gigantic 
Raptorial Birds o( New Zealand." 'Trans. New Zealand Insti- 
tote.' 1874. 

** On MegatapUryx Hulvri, a new Gigantic Species of Aptmy- 
fian Bird.^ ' Twn»- Zool. S«c..' vol. xii. ( iM«). 
mXLtv \V. H.;— "On the Classification of Birdv" 'Proc Zool. 
Soc- 1867. 

lOiNs {V.)— "R6:herches sur les Oiscaux Fos&iles des Terrains 
Tertiaires des Environs de Reims," nans 1 and 3. Rhcims 
-rDEKKKR (R.)—" Siwalik Birds." ' Pabpontologia Indica' (Mem. 

* GeoL Surv, Ind.'l, ser. to, vol. ill., pan 4 (1884). 
Marsh <0. C.}— "Odoniomiihes : A MonogTaph of the Extinct 
Toothed Birds of North America." W^^hin^ton (1880J. 

" Discwenr of a Fossil Bird in the Junissic i)f Wyoming." 

' American Journal of Science,' ser. 3, vol. xxi. (1881). 
NkwioK (A.)—" Ornithology .• ■ Encyclopaxlia Hritannica,' 9th ed. 

- and (£.)—■ Osteology of the Solitaire or Uidine Bird of the 
island of Rodriguet" ' Phil. Trans." 1869. 

- (E.), and Clark (J. W.)— " On the Osteology of the Solitaire 
{PtMOpkapt jniilaria, Cmel)." ' Transit of Venus Kx|>«^iti«n.' 

. (A.) and Parkkr (W. K.)—" Birds." ' Encydopiedia Hriian- 

nica,' 9th ed. (187;). 
, OWEJI (H.}—" Ar^fiai<fUryx macn/ra." ' PhiL Trans,' 1S63. 

. "Memoironihe Dodo." 1S66. 

, "Memoir on the Extinct Wingless Birds of New Zealand." 

,^^ "On Argt'J/armi IiHieip*nMis,Owi!n,n lar^'e Bird of Flight, from 

ibe Eocene Clay of Sheppey." 'Quart Joum. Gcol. Soc.,' vol. 

xxxvi (1&80X 
, "On the Skull of s Dentigerous Bird from the London Clay of 

Sheppej-" ' Quart Joum. Geol. Soc,,' vol. xxix, (1S73). 

- " On ninomit," ■ Tnins. Zool. Soc.' i839-«5. 

- " Otieolog>- of the Dodo." ' Trans. ZooL Soc' 1867. 
[fORTIS (A.) — " Comribuiioni alia Omiiolocia ttaliana," part 2. 

' Mem. Ac. R, Torino,' »er. a, voL xxxviii. (1887). 
PSeelev (H. G.)— "On the British Fossil Cretaceous Birds." 'Quart. 

Joum. GcoL Soc.,' vol. xxxii, (1876), 
SrSTCKLAXO (H. E,) and Melvii.le (A. G.)— "The Dodo and iu 
Kiodfcd." Loodoo (i&|6)- 


26. Vis (C. W. de).—" A Glimpse of the Post-tertiary A vi&nna of Q 

land." ' Proc. Linn. Soc. N. South Wales,' voL iii. (1888). 

27. VOGT (C.) — " L' ArchaofUryx macrura — Un IntermMiaic et 

Oiscaux et les Reptiles." ' Rev. Sci. France et rEtranp 
ix. (1879). Translated in the ' Ibis ' for 18801 

28. WiNGE(0.)— "Fugle fra Knoglehaler i Brasilien" (Birds fr< 

Brazilian Bone-Caves). 'Museo Lundii' (1887). 
39. Woodward (H.)— "On Flightless Birds." 'Proc Geologists 
ciation,' vol ix. (1886). 





Mammalia, or highest class of the Vertebrata, are character- 
by having some part of the integument provided with hairs at 
s period of life, and by the young being nourished for a longer 
\onex time by the milk, or special secretion of the mammary 
ds. As characters available in the case of fossils, it may be 
Tved that the cranium articulates with the atlas vertebra by two 
pttal condyles, mainly formed by the exoccipitals ; while each 
15 of the mandible consists of only a single piece, which proxi- 
y articulates directly with the squamosal element of the cranium 
out the intervention of a quadrate ; and there is no movable 
; between the proximal and distal rows of the tarsus. Like the 
opsida, Mammals possess during development an amnion and 
itois, and are totally devoid of gills. They differ from Reptiles 
agree with Birds in having a four-chambered heart, warm blood, 
a complete double circulation. They are peculiar in that the 
corpuscles of the blood are not nucleated and usually circular; 
le lungs being freely suspended in the thoracic cavity, which is 
rated from the abdomen by a muscular partition termed the 
kragm ; in the presence of only the left aortic arch ; and in the 
xrtion of the transverse commissure {corpus callosum) connect- 
the two cerebral hemispheres. Feathers, moreover, are never 
mt, and there is no syrinx or lower vocal organ, although 
mplete larynx is always developed in the upper portion of the 
\ea, or respiratory tube 

will be unnecessary in this work to make any further mention 
le soft parts, but a few remarks must be made concerning the 
mentary and dental systems, and the endoskeleton ; although 
itudent must refer to other works for fuller information on these 
^ts. With regard to the tegumentary system, it will suffice to 


observe that imbricated homy scales occur in the epidennit c 
the family Manida among the Edentata ; and flat honif d 
with their edges in apposition, in the tails of the Beaver, Rd 
certain Insectivores and Marsupials. The Anuadillos and G 
donts develop, however, a series of bony scutes aitknlatJDi 
one another in the true dermis, which are covered by hon 
dermal shields ; the whole structure being thus [H^dsely confi 
to that obtaining in the Crocodilia. Smaller separate bonyi 
also occur in the dermis of Mylodon. The homs of the Rnmi 
and Rhinoceroses are entirely epidermal structures ; the I 
being hollow sheaths enveloping bony cotcs, while the latti 
solid throughout. 

The dental system, as being of extreme importance for the 
mination of the extinct forms, must be noticed somewhat more 
Calcified teeth are developed in the great majority of Mamnu] 
in the true Whales they occur only in the embryo, in Ormtkerkj 
they disappear in the adult, while in Echidmty Mtam^ and it 
cophaga, no traces of them have as yet been detected. Id tb< 
of Omithorhynchus and Jihytina the function of teeth is diid 
by horny plates, or comuies on the palate. In all other foimi 
ever, true teeth, which are developed only in the iweinaxilla, d 
and dentary bones, are present ; and are usually composed 
three elements, dentine, enamel, and cement, although occasii 
as in the existing Edentates, the enamel is absent The d< 
or ivory, forms the chief constituent of most teeth. This is t 
either completely or partially, in the majority of cases by a tl 
vesting layer of the hard flint-like enamel, which is readily 
guished from the dentine by its bluish-white and translucent a 
ance, while the outermost coat of cement, when present, is (^ 
opaque white, or buff, colour. The cement is frequently foum 
as a thin coating at the roots of the teeth ; but it is very 1 
developed in the crowns of the hinder teeth of many Ungi 
In the teeth of the great majority of Mammals (as in fig. 1 1 1 
crown, or exposed portion, is sharply defined by a constr 
known as the tieek from the root, or embedded portion j but in 
teeth to be immediately noticed, which grow continuously, tt 
no such distinction between the crown and the root. In no 
mals are the teeth anchylosed to the bones of the jaw ; * am 
are invariably implanted in distinct alveoli, or sockets, whic 
however, very imperfect in certain Cetacea. In all young an 
while the teeth are still growing, the inferior extremity of tht 
or roots, is widely open ; but in the majority of instances thii 
ture becomes completely closed in the adult (fig. 1 1 36). In c 

' Except, perbap*, the inciion oT the Shmn. 



fever, as in th« anterior (nitting-tecLh of the Rodents and 
of the Elephant, the root remains pcrmanundy open be- 
nd the tooth consequently continues to grow throughout the 
t Uie animaL In such circumstances the tct'lh are said to have 
Iftcnt pulps. I'bc anterior tcclh (fig. 1126, f, i), with some 
exceptions, are of simple structure and have hut a swingle root ; 
he htnticj ones (MJ.,/m, m) very geiierally have more or Icsa 
■lex crown». which may be iupported by from two to four roots ; 
I division of the roots being unknown outside the MaminaJian 
L In many forms the summits or sides of the crowns of the 
In icclh nrny b< interpeneiratwi by d<cp rc-cnttring foldi of 
, which maybe filled up with cement ; thc:>c fold:^ being cspe- 


. inii.— Tank ol ihari^ uitt/ihaiotniftwotihrn ChimpaiuM. /, Inckon; 
' ' I ; fm, PranuUira : m, Tnj« noUrt. (Afl«r Owtn. ) 

developed in many Rodents and Ungulates. From this 
it will naturally result that when a horizontal section of the 
fn of such a toolh is made by ihc wearing of the up[)cr against 
I lover seriesL, an extremely complex pattern will appear, as will 
(Jfeca in the figures of the chueli-teeth of the above-mentioned 
Mps which are given below. Much more rarely, as in the Horse, 
BC may be an infolding of tlic enamel in the summits uf the 
fenn of the anterior teeth. 

Wkb the exception of Ihc above-mentioned edentulous forms, jn 
lodsdng Mammals one definite set of teeth, which is almost 
9mjk txmstant in nunilicr, is developed, and this set when it ap- 
Hs Dsaally persists throughout the remaining portion of the life 
Rs owner. In a large number of species this is ihe only set ever 
Kloped ; and sudi species, or groups of species, are conse<iuenily 
I to be i^onophyodeni. In the greater majority of Mammals, 
PCTcr, the development of occasionally only one, but usually of 
number of the anterior teeth of this permanent set is 



retarded, and their function filled for a time hy an earlier i 
of so-called milk-teeth ; such Mammals being accordinglj 
Diphyodont. As development proceeds the permanent teeth ■ 
Mammals come up beneath the milk-teeth, and thus replace 
in a vertical direction ; but there are instances where certain of I 
milk-teeth have no such permanent successors, while in otha 1 
the anterior teeth which come into use with the permanent 1 
tion have no milk predecessors. Those Mammals with a 

Fig. iii7-—0uler lateral aipect of the left d«n(iiion uf the Pig {Sta trrwlt), widi Ac« 
lamina of bane removed, in order lo exhibit the rooti of the teeth, i, Incuon; r, CaaiM;^ 

Premolara ; hTi True molan. 

phyodont dentition present the least specialised develoiMnent, im 
the milk-series appears from the latest researches to be an addtti 
grafted on to the permanent one; and there is accordingly no hoo 
1(^ between this definite single replacement and the irregular cc 
tinuous change which takes place in many Reptiles. In some Ha 
mals, like the Dolphins {Dtlpkinida), all the teeth are so much afi 
that they cannot be divided into groups, and the dentition is tb 
described as Homaodont ; but in the majority the permanent ted 
either from their position or their mode of succession, can be sq 
rated into four distinct groups (as in figs. 11 26, 1 127), and the d( 
tition is then termed Heterodont. In Eutherian Diphyodont Mai 
mals the total number of teeth of the permanent series does il 
normally exceed 44; and in forms like Sus (fig. 1127), or the < 
tinct Anoplotherium, where this full complement is present, the fi: 
three upper teeth (/ i-i' 3) on either side, which are situated in i 
premaxills, are termed indsors ; the last three {m i-mj), which) 
distinguished by having no milk-predecessors, true molars ; the ft 
submolariform teeth {/« i-pm i^ in advance of the latter, of whi 
the last three have such deciduous predecessors, prtmolan; a 



ftubconic^l toolh (c) situated between the premolars and 
( the tanitu ; the latter tooth being the first of those 

lie maxilla. The same terms aic applied to the corre- 
mer teeth, although it iriU be unnecessary here to indicate 
serial correspondence is worked out. For the sake uf 

ich a deDtiiion may be expressed by the nutiiencal 

_ - 3 — 3 _ » — t _ 4—4 ,. 3~i . ^ 

-J. ; C. ; Pm. j M, = 44; but 

3 — 3 '— ' 4—4 3—3 

teeth of opposite sides of the jaws a]wa>'s coneipond, such 
may be further Bimplified into — 

I, ^; C.-; Fm. ^ ; M. ^ = 22x2 = 44. 
31 4 3 

ridual cecih of each group are enumerated from befote 

and by such a formula as the followinj; — vijt., 

, I. 3, C , Pnt. t. Pm. a, Pm. 3, Pm. 4, M. r, M. a, Af . 3^ 

^- 3* C., Pm. I, Pm. 1, Pm. 3, Pm. 4, M. i, M. 2, M. 3, 

tdividuil tooth can be ftpcci-illy noted. Thus, for example, 

indicate the first upper premolar, and m. 3 the third, or 

true molar. It will frequently, moreover, be convenient 

of the incisors xnd the canine collectively as the cutting-, 

premoUrs and true molars as the chrfk-tecth. It is very 

the caae that when the true molars arc reduced to less 

« it is the hinder tooth, or teeth, that disappear ; in the 

n it is, however, frequently the anterior teeth that arc w-mt- 

tnough this is by no means invariably the case, and there are 

ees kriown where the second and fourth disappear, while the 

ild third remain. In the figure of the lower dentition of the 

given on page 1247, the two premolars arc usually 

as /m. 3 and pm. 4, but ic is not certain that such is 

case Again, it has been suggested that the two incisors 

that species, in common with other Frim.iies, may be the 

of the typical EuLhcrian aeries of three, although other 

;ard ihena as the second and third. It nuy aho he oU- 

t in some groups, as the Carnivora, the ^specialised forms 

lose the molani and retain tlie full number of anterior teeth ; 

Others, like the Ungulates, the reverse conditioii obtains. 

^•dentition may be expressed by a similar formula with 

Ibc Icttci M. to the symbols. The typical milk-series 

be written as M.i. ?, M. c-, M.m. 5 ; the three milk- 
3 ■ 3 

ding to the last three premolars of the permanent 

In a few Ungulates, however, such as Tapirui, and some- 
'noares and PalaalAerium, four milk-mobrs are developed. 


Among the existing Metatheria the number <d true 

generally ~, while the premolars are veiy frequently 

number, and there may be five upper incisors. In one < 
and several Mesozoic members of that subclass, the nuiober 
molars exceeds four ; but, with the possible exception of i 
these extinct types, there is no known instance of a 
Mammal normally having more than four premolars ; and 
is never more than a single canine tooth on either side si 

Since in Ungulates it is sometimes difficult to <1iirt^ngiiiih 
from premolars, it may be well to mention how the divisioa bd 
the two series can always be determined. Since the first loodi 
true molar series always comes into use before the last miltiM 
shed, it is obvious that in the adult the first true molar will 
be more worn than the last premolar. Thus, in the thtee 
Hipparion represented in fig. 1233, the tooth on the left 
the figure being more worn than the one in the middle 11 
shown to be the first of the true molar series ; the other t«D 
consequently premolars. 

The different types of cheek-teeth will be mentioned undd 
head of the various orders and families, but a few genoal obi 
tions may be recorded here. Professor Osbtmi considers thit' 
primitive Mammals had simple conical teeth with undivided fil 
and that the crowns of the teeth of the upper and tower jam 
tually interlocked. Teeth neariy approaching to this type ocxfl 
the Triassic Dromathfrium (fig. 1 140), while those of the Dolfl 
are looked upon as a reversion to this type. Another simple % 
according to the views of the same author, is that found ia Trdl 
don and Priatodon (fig. 1147), where the upper and lower M 
alike consist of three cusps in a line ; the upper teeth biting oa 
outer side of the lower. A third common, and apparendyi 
generalised, type of tooth is that known as the trituSertular. 1 
consists in the upper teeth of ont inner and two outer ciufi^ 
ranged in a triangle ; while in the lower jaw the reverse anangtn 
obtains, so that there is one cusp on the outer and two cuspson 
inner side of the crown. An example of this type of structm 
its simplest form occurs in the Mesozoic genus ^altuother 
Modifications of this type occur in the lower teeth of many M: 
pials {e.g., fig. 1 145), and also in the lower camassial teeth of 
Camivora, of which mention is made in the sequel The trie 
cular tjrpe of tooth is regarded by the American Palseontologi! 
one which has given rise to a large number of the more con 
modifications ; and it is extremely common among the genen 
Mammals of the Lower Eocene. 


1 351 

lammaU, sa in the Sauropsida, the whole of the primi- 
inau» cranmni is replaced by i-xtensive ossiRcations, 
f ethmoidal region ; and these bones, vrith the exccp- 
e of the mandible, fiyoid arch, and internal auditory 
» similarly articulated togeiher at their edges by suture, 
I on there is generally a tcndcnf:y 10 the oblitemcioii of 
I, this being most marked in Mam's. In all cases the 
i has ceased 10 exist as a disltncl ossificatioii. It has 
k observed that the hinder pan of the cranium aniculates 
Bt, or atlas, venebra by two exoccipital condyles, and 
ich of the two rami of the mandible is composed of a 
articulating at its pruximal or hinder extrcmily with the 
if the cranium. Owing to the comjiWif incorporation of 
isal and parietal in the walls of the brain-ease there is 
{irds) rK> superior temporal arcade ; but an inferior, or 
■rcade is nearly a]wa>-s prct^ent ; and, as in Crocodiles 
tot in Birds), forms the lower border of the orbit. This 
tporal arcade, or arch, differs, however, from that of moat 
in that its jugal element articulator directly with the 
in vhkh respect it accords with that of the I'icynodont 
md should be known as a squamoso-maxiltary arcade. 
to the view of Professor Huxlej-, the Sauropsidan quad- 
een taken up into the inner ear to form the m.illeus. 
ntiy, howewf, other ■n-riters h.nve taken a different view, 
ling to Dr Batir. the reprtrscnlativi; of the quadrate is to 
n the zygomatic process of the squamosal, with which the 
nUtcs ; the quadratoju](al being also represented at the 
f these two bones, Dr Gadow, however, disputes this 
finds the representative of the quadrate in the tympanic 
letennination according with the view here taken as to 
Og7 of the tygomatic arcade, which appears to have no 
(pi eientenL 

Rrben, as in the Primates and many Ungulates, the orbit 
1 posteriorly by a bony postorhilal, the asrending pro- 
e jugal articulates directly with the frDnlal, without the 
<n of the po«torbital or postfronlal, which forms such a 
U feature in the Reptilian skulL 

cases the premaxilUe, maxilla, and palatines develop in- 
tal plates which meet in the modinn line below the nasal 
ad thus completely separate the Utter from the cavity of 
, Except, however, in the Anteaiers {Afyr'nfrt}^A/tga), in 
of Armadillo (Tatusio), in certain Cetaceans, and in one 
i Hyttnodon, this flooring of the nasal passage does not 
ckwards to include the pterygoids, as it does in tnadcm 
U. The palatines are always placed behind the maxillae. 


Finally,, the sclerotic of the eyeball never develops a rii^i 
plates like that so frequently found in the Sauropsida. 

A characteristic feature of the larger bones of Mammal^ i 
one by no means peculiar to the class, consists in their < 
from several distinct centres. In the case of the Icmg 
shaft is formed by one centre of ossification, while two i 
ments termed epiphyses form the extremities ; in the aduh die^ 
of these being welded together into a solid mass. The loagl 
are also tubular, and their vacuity is filled with the fatty nunv 

The vertebrae always have well-developed articular procesa 
their arches ; the ends of the centra are generally flattened, I 
the cervical region of certain Ungulata. they may be opisthoco 
Terminal epiphyses, so generally wanting in the Sauropsida 
nearly always present The number of vertebr% varies p 
owing to the great difference in the length of the tail in dil 
species ; but in the majority of Mammals the number of pcec 
vertebrae does not vary very far from thirty, although in I 
and Cholcepus their number reaches forty. In spite of the 
difference in the length of the neck in different Mammal 
number of cervical vertebrae in existing forms is, with three D 
exceptions, seven. These exceptions are Manatus amtrali 
Cholapus Hoffmanni, in which the number is reduced to sii 
Bradypus tridaetyius, in which it is increased to nine. A 
ing, however, to Professor W, K. Parker there may occasii 
be eight cervicals in the Pangolin {Manis). The first, or 
vertebra always has two articular cups for the occipital con 
and, except in certain Cetacea, the second, or axis, has a wi 
fined odontoid process. Usually the cervical vertebrae are 
free ; but they are anchylosed together in some of the Cetaco 
in the Armadillos. The dorsal vertebrae are usually well de 
from the lumbar, although this is not invariably the case ; an 
number of dorso-Iumbars in any one given group is usually 
constant, and among the Ungulata afTords assistance in classifio 
A distinct sacral region is present in all Mammals except tb 
tacea, where the iliac bones are absent. The number d" a 
vertebne varies from three (certain Primates) to forty-six (Jfi 
Chevron-bones are present in the caudal region of many long-t 
forms. The sternum is always present, although varying greai 
form. It usually consists of a presternum (fig. 1 1 28, /) and 
posterior xiphistemum (x), between which are a varying numb 
segments (m) constituting the meiostemum. The segments ol 
mesostemum (fig. 1128, a) may be anchylosed together; u 
the Balanida, among the Cetacea, only the presternum is pre 
The connection of the ribs with the sternum is generally by ( 
lage ; but in the Armadillos the costal cartilages ossify, and ue 



r^Mat ri^t. lo the anterior doi^al region the capitulu 

ribs onicuhlcs with the vertetirx in a pic at the junction 

while (he tubercular head joins the transverse process 

;d of these two vcnebne ; but in the posterior dorsal 

mro heads generally coalesce l)r Raur regards the 

ral Kiticulation of the capicula in the anterior region a<t a 

Tftl of the iniercentrai stcachmeni of the ribs of primitive 

d to Thcriodont Anomondontia, wliich has been totally 

other Reptiles ; and the mode of costal articulation is 


I, Surnma wd ruhl catuU onihcn of Mui: B, Surnum ani! IeA coiXtl larilbja 

dly very near to that of those Anomodonts which have 
Dtercentra. Uncinate procesnes are never present on the 

Mammals the pectoral Umb is well developed. The pec- 
me is, however, usually simpler than in lower forms ; the 
persisting as a distinct bone only in the Monotremata, 
)wever, it anchyloses to the scapulx .\n iiiterdavicle is 
>nly in the last-named order; where it articulates with the 
after the Reiwilian manner. In the Euthcria the clavicles 
in their complete development in those grou[>;, such as the 
n, many Rodentia. the Chiroptcra, and the Primates, 
e the pectoral limb for Sight, Isurrowing, or prehension. 
Mctus frequently has an emcpicondylar foramen, like that of 
nodont Kcptiles. The radius and ulna retain their original 


pre- and postaxial position in the Cetacea ; but in most i 
Rials they are crossed at their distal ends, so that the 
become reversed, and in the " prone " or normal positioa > 
the radial, or preaxial border of the hand, becomes intern 
Primates, however, these bones admit of motion upon on 
and when the hand is " supine " (that is, with its palm d 
wards or upwards), the bones of the fore-arm occupy thi 
primitive position. In the majority of those Mamiiials w 
are adapted solely for walking, the ulna is more or less re 
the radius, especially at its proximal end, is much enlarg 
it articulates with the whole of the anterior sur^ure of the 
and thus comes in front of the ulna, instead of at its s 
.carpus essentially corresponds with that of the type repi 
fig. 829 (p. 907) ; the radiale,' intermedium, and ulnare be 
the scaphoid, lunar, and cuneiform ; the trapezium, traf 
magnum representing the ist, 2d, and 3d carpalia, and thi 
the 4th and 5th of that series. A centrale is present in 
of embryos of pentedactylate forms ; but in the adult 
fuses with the scaphoid, although it remains distinct i 
the Primates. In some groups others of these elec 
also coalesce, and one or more may be absent The p 
presumed representative of the seventh digit, is genera] 
veloped ; while in pentadactylate types there may be an ( 
representing the prepollex. The metacarpals and digits n 
in number (Proboscidea and Primates), or may be reduc 
or even to one functional member. Among the Ungu 
the metacarpus is reduced to a single functional element 
Horse, such element is frequently termed the cannon-itm 
tomy, however, this term is more usually restricted to th 
dium of those Artiodactyla which consists of the coale 
and fourth metapodials. Except in the Cetacea there 
more than three phalangeals to each digit, but by sui^ 
anchylosis this number is occasionally reduced ; and the 
(pollex) has but two phalangeals. The pelvic girdle is 
developed in all Mammals except the Cetacea and Sireni 
the aduit the three elements coalesce to form an infumii 
The pubis and ischium of the same side always unite to e 
obturator foramen ; and the two pubes meet in a ventral • 
which is, however, not completely united in certain Insed 
a large number of instances the ischia meet in a ventral 1 

' Since the preceding chapters were in type, Dr Baur bu expmsec 
that the bone termed radUle in fig. 829 is reallr ■ tecood cenirale ; 1 
radiale is represented by a minuie bone generally known m the ndii 
The Mammalian scaphoid is accordingly also regarded a« a second c 
the same communication, Dr Baur exprejses his disbelief in the exist 
nants of a prepollex and of a seventh digit in Mammals and oihei V< 



itly etongatnl in Miiie Un^uLiCcs {fig. 1 1 3& di's), bui in 
es and soint: other (wmt. there is no such union. In (he 
kU and Monupiolia trpipubic, or marm/ia/, hotKH arc 

the anteriur border of ihc pubic symphysis (fig. 1 131J). 

may have a third trocKanlcr (fig. taib) for the atlAch- 
ic of the gluteal muscles. The tibia and fibuU ari- never 

their di^ataJ extremity, but lie in their primitive parallel 
the tibia, or prcaxial bone, being internal in the usual 

ilion, and the fibula external. The latter bone may be 
ruditnentar)', and completely aiichy!o»cd ai one or both 


->^ % , 




^Kt.—Tht UA die «f ttw ptiiuaf ihe Eland (OrtMi taiuui\. OncttTtb nuanil tiie. 
li, IkIuiwb; >■ SrBphriii «r dbi ■, EjiphyiUan >)iini4iy>U of licbluin and vubi>; 
tt, ffbtvr%lar t 

ties to the tibtJi. The patella is present in all except some 
la. If the pes (fig. 1139} be cntnpared with the typicil 
mentioned on p. 907, it will he Tound Ihai the libiiUre i» 
ited by the calcancum, which may also contain an clement 
londing lo the jHsifunii of (he inanu.t : the a»tni^alus has 
iually r^ardcd as the coalesced tibiale and ijitemiedium, 
ht by Or Baur to correspond solely to the latter : 
I-, and cctocuneifomi reprcBent the ist, ad, and .^d 
] while the 4th and 5th tnntatia have coalesced to form the 
The cenliale persists as the navicular, which may unite 
cuboid.* Other modifirations occur an.ilogous to iho^e 
Orpos; but in no instance, as already obser\-ed, is the joint 
the leg and the pes fontred on the line Ixitwcen the proxi- 
Bd disul rows of the tarsus. The meUlorfals and phnlan- 

Ittdekbin liu poinied out to the wnici thfti in Cryfifprttia ihcre may lie 


) otatnle, which luually <oal««xi with ibc ectocauct. 
> IL 

a A 





geals repeat the characters of the homologous bones of th 
although they deviate in a lesser degree from a conun 
In the Cetacea and Sirenia the pes is entirely absent ; althi 
proximal portions of the pelvic limb may be detected in an 
form in many genera. 

We may now talce a brief glance at the general distril 
time of the Mammalia, in the course of which we shall tun 
stall some of the information given in the sequel 

If the theory of evolution be the true eiqilanadfKi of t 
of nature we should expect to find that the earliest repres 

of the class would be smal 
more or less closely related 
another, not numerous in genei 
and allied to those orders whi 
comprise the most generalise 
sentatives of the class. Our kr 
of these earlier types is indeed 
ingly imperfect, but so far as i 
appears to accord fairly well 
foregoing conditions ; all the 
forms being of small size, an 
ently more or less closely allie 
existing Marsupials, and probi 
to the Monotremes and peri 
Insectivores. The earliest evi 
the occurrence of Mammals ye 
is in the upper portion of the 
system, which forms the bast 
great Mesozoic period. Of 
known genera from these dep 
larger type, termed Dromalfut 
hibits some curious approximations in the structure of its 
Reptiles and Amphibians, and it is possible that we may 
this form an ancestral type of the Prototheria. In th 
Jurassic, immediately above the Lias, remains of small ! 
become more common, and in the Upper Jurassic they ai 
abundant, although they comprise comparatively few genei 
These Jurassic Mammals are readily divisible into two grouj 
guished by the characters of their teeth. Of the first groi 
ample of the lower jaw is shown in fig. 1141, p. i»74, wht 
be seen that the teeth are very numerous, the hinder one 
several sharp cusps on the crowns. Although there has be 
discussion as to the affinities of these small Mammals, it is 
that they are realty Marsupials of the suborder Polypro 
with a relationship to the Australian KnttaXa {Myrmteotiu: 

Fig. 1119. — nortal a«pcct or ihe 
right pcK or Mul. Retluced. Ca, 
Cucancum (fibulirc 4- T piiiroim) ; 
Atf A,<itraeatii-^ (imcrmcdium) ; c. 
Navicular (centralc); Cti, 1-3, Ento^. 
meM>-, and cctocunciform (iftt, ad, 3d 
lar^alLa): Cub, Cuboid (^tn, 5th tar- 
salta): i-v, MctatarsaJ!!. (After Wie- 



aatl see in the setiuel, is the only existing hetcrodont 
I witH more ihan four cruc molars. In the second group 
tition is of a very dificrcnt type ; the lower jaw having a 
jkur of curved and chiscMikc incisors, separated by an 
from the chee): -leeih, which are characieriised by the 
£ of one ur more lun^tudinal grooves. The serial position 
i group, whK:h has been named Multituberculau, is still 
o doubt, but not improbably it may indicate an extinct order 

irou^hout the greater part of the Cretaceous period our know- 
It t>f Mammalian Mh is a blank, doubtless owing to the ctTcum- 
IX that the grisitcT iwrtion of the Cretaceous beds, such as the 
ft, t» of jHirely marine origin ; hut evidence has recently been 
joned of Uie existenec of a Mammalian fauna in the topmost 
jlKtous of North America allied lo [hat of the JuniNsic. 
flTah the dawn of the Tertiary period, which may be rej^rded as 
fifst commencement of the present order of nature, we meet, 
iMwr, with an abundant Mamntnlinn faun:], containing repre- 
Utivcs of nearly all the cxistmg orders, but aNci including several 

Kinal tv-pcs now totally passed away, some of which are of 
e intercut to the zoologut as connecting together more 
Hea completely groups which are now M-idely separated. In 
' Eocene wc are indebted lo the palscontologiM.s of the New 
irM for mo6t of our knowledge of these primitive connecting 
ts ; but we can only afford space to notice very briefly some of 
' more iiitereuing group». 

Of CarDivDTOUs types there ia a group of Eocene genera known 

the Creodonta, remarkable, among other features, fur their 

tcralibcd denblion, which has resemblances lo thai of the Poly- 

Kodonl Nfarsupialia and Insectivora, and also to that of the 

Carnivort, of which these forms were probably the anccs- 

We thus have indications how the Camivora of the present 

ly have been gradually evolved from a Marsupial type by 

of forms more or le*» nearly relatrd to the Insectivora of 

Ent epoch. In another direction the Kocene has afTorded 

of a transition from the Insectivorous type towards that 

l^cmuroid Primates, and we may thus readily conceive how 

higher members of the latter order may likewise trace back 

or^in lo the same primitive stock, 

the present tlay no orders of Mammals appear more sharply 
ined from one another than the Carnivores and tiie Ungulates. 
the Eocene, however, we meet with a group of primitive Un- 
otes, known ».% the Condylarthra, presenting such remarkable 
emblances to the primitive Camivore^t, that wc are led to the 
tclusion that the Ungulates are probably another liranch derived 


from this same prolilic stock. From this Condylarthrous sabot 
we have abundant evidence that the two existing suborders tt 
Artiodactyla and Perissodactyla, now so sharply distinguished^ h 
both taken their origin ; and perhaps the Hyracoidea may 
trace their derivation from this group. Ano^er extinct Eon 
suborder of Ungulates, comprising the Coryphodons <^ Europe 
.\merica, and the huge and uncouth Dinocerata of the Un 
States, tends to show a connection between the Perissodactyk 
Proboscidea, which till recently were referred to distinct orden. 
All living Ungulates, as we shall subsequently mention, we 
tinguished by the total absence of clavicles, but in the remaikil 
Typotherium, of the Pleistocene of South America, these bonei 
retained. In its dentition, moreover, that genus shows fotn 
now peculiar to the Rodents ; and by its help, together with 
evidence afforded by an Eocene group known as the Till^ 
we can dimly see how the Rodents may have been connected 
the ancestors of both the Ungulates and the Carnivores. 

Having seen from these brief notices how intimate z^ipeaa 

have been the relationship between the chief terrestrial onten 

Mammals in the Lower Eocene, we may glance at the 

afforded by the Mammals of the Eocene as to the comieditfj 

between some of the families of these orders. Among the Oit 

nivora, no two families are better distinguished than the Dogi, dC 

Canida, with their triangular upper molars and digiiigrade ftf^ 

and the Bears, or Ursida, in which the upper molars are rhoB- 

hoidfti and the feet are plantigrade. In the Eocene, however, K 

have Ampkicyon, with the teeth of a Dog and the limbs of a Bar; 

and in the later Tertiary the Bear-like Hyanarctus, of which Ae 

dentition retains many Oog-like features. So perfect, indeed, it 

the transition between Dogs and Bears, that it seems convenient ti 

include both groups in a single family. Again, in another directing 

the Dogs of the Eocene seem to pass imperceptibly into theCrnn 

{Viverridie) through Cynodktis ; while it is almost impossible » 

distinguish the Civets from the Hyaenas on the one hand, andlbe 

Cats {Felidie) on the other. Still more remarkable is the appinal 

connection in the Eocene of the Civets with the Weasels {M<u- 

lelidie), since these two families are widely sundered at the pro- 

ent day. 

Turning to the Ungulates, at the present day the Artiodanybtt 
suborder can be readily divided into four sections — vit, the Pip, 
or Suina ; the Camels, or Tylopoda ; the Chevrotains, or Traguliiu; 
and the Cattle and Deer, or Pecora. When, however, we go bid 
to the early Tertiary, we find a complete transition from the Suittt 
to the Pecora ; while it is almost impossible to distinguish Deer 
from Chevrotains ; and the early Camels exhibit signs of clo>t 



ion with ihc other families. Similarly, at the 5amc {iciind 

of the Old Wcwld appear to be inscnarahle from the 

of the New; while ihc Giraffes were proI>abIy as closely 

with the Antelopes on the one hand, and the Deer on 

Again, in ihc Pi:ri^%ud^ctylale subordtrr Tapir like funiia 

to have pas.sed into Rhinoceroses on the one hand, and into 

on the other. 

ly other equally striking instances could be cited of the 

connection of the Eocene MammaU with one another ; but 

has been stated to show that the present sharply defined 

of the orders and families into which vi-c find it con- 

to divide the r.las!i 1.%, so to &pcak, but a feature of 

[llic TiOirer Eocene all the genera of MammaU appear to be 
hot in the Upper divwion of the same period, which is 
Jy termed the IjOwct Oligoccne, we meet with a few existing 
surh as DidiJphys^ /i/tiitMeros, i^iverra, Musle/a (VVea«el), 
cihaiis Cant's. In the succeeding period, or Miocene, existing 
Ijerome more common : thus in the Lower and Mitldk- divi- 
iis of that period wc meet with Otters {Lutra). Rhinweros, Tapirs 
m), and (}ibl>ons [NyloSatti). The middle division of the 
le is, indeed, noteworthy for the appearance of Anthropoid 
I and the ProboKcirleo, or Klephants and Mastodons. Deer of 
genera are abundant ; but ihese were either devoid of antlers, 
these appenda|{e« were presenl, they were small and simple. 
teeth of lite Ruminant Ungulates {Deer and Cuttle) were, 
low-crowned or braehydoni (5g. isij), and the Rhino- 
had in most cases not yet developed horns, 
the commencement of the i'liocene period the Mammalian 
assumes a much more modem appe.ira.nce. Thus we have 
[lines (ffysirix), Ii)'3!nas. targe 'ligcr-likc C!ats {Jvlis), numer- 
Antelopcs, Giraffes, Deer (Cervus), and Horse-like antmah 
■urn) ; while in India true Elephants {F.Uphas) had made 
appearance. 'Iltere was stilL, however, a lai^o number of 
ct genera. By this time many of the Tlccr h,^d acquired 
Neated antlers ; many of the Ruminants had tall-crowned or 
W teeth ; the Rhinoceroses had horns ; and the I'igs had 
large tusk* in the males. In India during some t>onion 
PHoccnc, not only Elephants, but true Horses (E^uusX 
tami, WoK-es, Bears (f/nw*), and Oxen {lion) had already 
ide their appearance ; but in Europe these genera arc unknown 
fore the top of the Pliocene, when we first meet with remains of 
few- cxiMmg Hpccicit, such a-s the .\frican Hippopotamus and the 
iped Hywna. 
to the succeeding Pleistocene period, which may in reality be 


regarded merely as the commencement of the epoch in wfaiiA^ 
are now living, the greater number of the Mammals of 
belong to existing genera, and a considerable proportion of 
living species, 'llie fauna of Europe in the early part of thii] 
included, however, a large number of Mammals belonging to | 
or species now confined to the wanner regions of the globes i 
as Rhinoceroses, Hippopotami, Elephants, Lions, Hysnu, 
and it was not till after the great cold of the glacial period I 
these generic types were finally swept away from the 
area. In many other parts of the World the Pleistocene 
was equally prolific in large forms of Mammalian life, morecrl 
closely allied to those now inhabiting the same areas, of whidi ' 
have remarkable instances in the extinct Edentates of South . 
and the Marsupials of Australia, many of which vastly exceeded i 
size their living relatives. Science has, indeed, yet to account i 
factorily for the disappearance of this exuberant life, and the i 
quently impoverished fauna among which we now dwell. 

In regard to the Mammals of the Eocene, we have alreadyi 
marked that they are frequently of a more generalised type 
those now existing, and in many groups a well-marked 
specialisation can be traced as we approach the existing 
Thus many of the Eocene Mammals possess the full Eut 
complement of forty-four teeth, which in the Ungulates were mnt^ 
uniform in size, less conspicuously differentiated into groups, nl' 
more approximated to one another than in recent forms. In the 
older Ungulates, moreover, the crowns of the cheek-teeth were re- 
latively short ; and we can trace a gradual increase in the height of 
the crowns as we advance in time, this increase affording a greater 
capacity to withstand wear, and thus indicating a greater length df 
life in the individual. In other instances we may observe a gradnd 
reduction in the lateral digits of the typical pentedactylate limb^ 
accom]>anied by a progressive elongation and strengthening of ooe 
or more of the remaining digits. Again, it has been shown thit 
there has been a gradual increase in the relative size and the com- 
plexity of the brain as we approach the present day. Thus the 
Eocene Mammals, as a rule, had very small brains, in which the 
hemispheres left the cerebellum nearly uncovered, and were them- 
selves nearly smooth, or but slightly convoluted ; while in the higher 
forms the hemispheres spread backwards over the cerebellum, and 
are often marked by most complex convolutions. In some Eocene 
Ungulates the brain was so small that it could pass through the 
neural canal of the lumbar vertebrse. 

tinally, it should be observed that we may trace a gradual e^x)lu- 
tion of local faunas. In the Eocene of any given region we find 
the Mammals differing widely in generic types from those now in- 


that arcA. yet as we aM:end in tbi: gcolugic^tl s<:alc vrc trace 
appronimaiion to the existing fauna; ;ijk1 in the Flcisto- 
find the cfaaxacteristtc features of Jiuch fauna distinctly 
oui, although many of the gcni;ric typrs, e&pecia)]^ those 
forms of large corporeal bulk, may Ik diflcrciit. Ex- 
of ibesc features are nfToTded by the Mamupials of ihe 
of AuiiUalta, oiid the numerous Edentates of the same 
in South America ; although it in practically ceilain thai in 
case the fauna has always been Marsupial, and therefore 
ror less closely related to the present one. Thm many of the 
Mammalian fiuna)^ now rharacieristic of particular ro^ons 
I itut thu:s circumscHlKc) till a late period is, however, shown by 
urrcnre of Baboons, Hippopotami, (iiraffcs, and /Urican 
• oi Antelopes in the Pliocene and I'lcistoccnc of India and 
of Europe ; and equally by the Mammals of the Pleiiitocene 
being in many rases specifically identical with those of 
. while the number of common species h now very few indeed, 
iin, many forms like the Rhinoceroses and Horses, which are 
Mi exclusively Old World types, formerly wandered over the plains 
America, anil thus point to a more unifonii distribution of types 
now exisu. There are, huwever, itidications of certain groups 
Mammah having always been restricted to one hemisphere, 
we ha*-e no evidence of the existence of Apes, Hyrenas, or 
. at any epoch in the New World ; and neither arc there any 
of the Dinoccrau or Titanoshtriidtr in the Old World, 
was considered probable *oine years ago thai the Maunnalia 
directly descended from some primitive Amphibian types, and 
|wt they stood altogether apart from the Reptiles. The striking 
JHemblance of the pectoral girdle of the oviparous Monotremes to 
Itat of many Reptiles, and more especially the Anomodonts, to- 
pether with the remarkable approxmiaiion to n low Muniinalian 
9pe presented by the skeleton of the latter, renders it, however, 
Mac probable that Mammals are a divct(;i:nt branch of the sanie 
itphibian stock which gave origin to the last-mentioned group, 
Iced they he not the direct descendants of the earlier forms 
It group. I)r Baur, who considers that Mammals were dc- 
from true Reptiles allied to the primitive Khynchoccphalia 
Ud SaorapterygLi, has, propof;cd thnt the^e early hyixtthelical forms 
Ultmld be tenncd Sauroniammalia. Professor Mivart has, indeed, 
that Mammals may have had a dual otij;in ; and that 
Be the Monotremcs may have l>een derived at a comparatively 
date from the Anomodonts or kindred types, the Marsupials 
Bay have ori^nalcd at an earlier epoch from a totally distinct and 
pvinps Amphibian stock. M.iny objections have, however, Iwen 
tins! this view ; and Mr Poulton cxpicascs his opinion that 


" whether the Monotremes are the descendants of the i 
Mammalia or not, it is quite certain that the hi^er Mammi 
at one time have passed through a condition such as now « 
the Monotremes, in nearly all parts of their organisation ; an 
powerful arguments can be brought against the assumption i 
same stage has been reached independently, and at widely se 
periods, in the course of evolution." ' 

In the following chapters are given the leading palseoot 
characters of each order of the class, with the range in time 
different groups, and the names of the more important genen 
number of genera is, however, so great that only a very br 
general sketch of their characters and afiinities can be given ; 
attention being drawn, where it may seem necessary, to those 
types which are of more than ordinary interest from an e\'oli 
point of view. 

' The opinion has been recently exprened by some Continental writen 
Cetacea are the roost archaic type of Mamoials, and that they haTe bee 
derived from the Ichlhyopterygian Reptiles. There is, however, so ■ 
dence against this view that it may be contideied a* pncticftlly dispcDie 



CLASS MAMMALIA— amtinued. 

Orders Monotrehata and Marsupialia. 

LASS I. Prototheria. — This subclass, now represented by 
:wo genera, may be characterised as follows. The brain has 
fc anterior commissure, and a very small corpus callosum ; * 

the auditory ossicles are simple, and the stapes is rod-like 
nelliform). The coracoid is a distinct, although small, bone, 
'losing in the adult to the scapula, and articulating with the 
im ; while there is a separate precoracoid (epicoracoid), which 
not articulate with the scapula, and also a large T-shaped inter- 
:Ie ; the form and relations of these bones being very like those 
e corresponding parts of the skeleton in the Anomodont Rep- 
The pelvis has epipubic bones, and the ilia are inclined to 
icral axis after the Batrachian fashion ; thus resembling those 
e Pariasaurian Anomodonts. The urinogenital and excretory 
IS open into a common outlet, or cloaca ; and the former are 
similar to those of the Sauropsida ; the mammary glands are 
avided with nipples, and the reproduction is oviparous ; the 

being meroblastic like those of Birds. 

nay here be observed that the small bone in the pectoral girdle of 
Honotremes placed in advance of the coracoid, which is usually 
xi the epicoracoid, appears to correspond with the precoracoid of 
jiomodont Reptiles * (fig. 978 bis, p. 1054), although it docs not ex- 
upwards to articulate with the acromial process of the scapula, as 
inomodont precoracoid articulates with the process of the scapula 
ified by Sir R. Owen with the acromion.^ Further, the scapula of 
lonotremes differs from that of all other Mammals, and resembles 

he structure connecting the two hemispheres of the brain. 

tiis appears to be the view taken by Professor Cope. 

his is the original view. In deKribing the scnpiila of Plalyfiodasaurus, 

. Owen confused the process situated above a "i fig. gyS 6is with that 

-d a, and tenned the fonner the acromion. 



that of Reptiles in that the acromion is situated on the anterior v 
axial border, which in the higher Mammals has become twisted 
to the dorsal surface to fonn the spine of the scapula. The 
some of the Anomodont Reptiles appears to indicate how this 
the preaxial axial to fonn a ridge on the dorsal suz&ce has taken 

At the close of the precetling chapter reference has been 
the relationship of the Prototheria, or rather of its existing 
sentatives the Monotremata, to Reptiles and Amphibia, lit 
ture of the pectoral and pelvic girdles presents, indeed, so 
striking resemblances to the same parts of the skeleton in 
Labyrinthodont Amphibia and Anomodont Reptiles, as to inA 
as already mentioned, a distinct genetic connection betweet 
three groups ; the characters of the humerus aj^arently indkl 

Fie. iijo.— A. Lateral view of skull of fcitiVaa; M,Do. at OrmilM»rtr^AMa: ci 

manJible of do. ; u, Siernnl riuion of do. (, Clavicle; (, InterclBviclcijt, Pw 

sternum ; r. Ribs ; ir, Sienial ril» ; i>, laicrmcdule Kb* ; ■■, Connllcl. 
Flower; ihe others after Giebel.) 

that the nearest relationship is with the Anomodonts. Tfce 
ciduous teeth of Ornithorhynchus indicate, moreover, witbout 
doubt that the ancestors of the Monotremes were provided 
persistent teeth, which were probably monophyodont. H 
over, a distant resemblance between these deciduous teeUi 
the cheek-teeth of the extinct group, mentioned below under 
name of Multituberculata, suggests that these forms may themsi 
be Prototheria. If this should prove to be the case, it would 
be ap[urent that that branch of the subclass could not have 
the ancestral stock of the Marsupials ; and we shall accord 
have to look for another group or order of Prototheria, wi 
dentition akin to that of the Pol)-protodont Marsupials. 



quite within the bouivls of probability (hat (he fiiniily 
i$da mentioned under the head of ih« latter group 
to be Prolotherians. And in any case it is quite 
that the vko exiting genera of Monotremcs can in no 
looked upon as actual ancestral t>-pes. 

MoNOTKiiMATA. — ;\s already mentioned, both the 

, genera of Protothcrians arc included in this order, which 

' pccnrisianally characterised by the proiluction nf the cranium 

or less elongated rostrum supporting a horny beak ; by 

cc of lecth in the fully adult animal ; the smoothness of 

; and ihe absence of ar auditory bulla. The humerus 

'expanded, and lia:i an L-ntepicondylar foramen ; its whole 

approximating to the corresponding bone of the Anomodont 

(fig- 983). 
ILV ORNrmoRHYNCHiDX. — In this family the evrcbral hemi- 
arc smooth ; the extremity of the muzzle is produced into a 






Fi|-' > ■ ) I .— Om/Ufnt/miM f*r»Jt. 

. Autlralij- RbtiKvd, 

losd bcak-like expansion (fig. 1131); tcclh arc present in the 
img. and are succeeded by horny platen or comules (fig. 1 130, b, 
inii the skin is covered with hair. The feet arc webbed. This 



family is represented solely by the genus OmitfiorhynekMS (fig. 
of which a single species inhabits the Australian rivers, in the 
of which its burrows are constructed According to the d 
tions of Mr O. Thomas, it appears that there are usually m 
on either side of the upper and three in the lower jaw, whidi 
till the animal is somewhat more than one-third grown. Tbea 
have at first stnall but distinct roots, and present a distant : 
blance to the true molars of some of the Multitub«cubti 
MieroUstes ; one of their longitudinal walls, or ridges, catr 
number of minute cusps. It appears that the comules grow 
neath and around these teeth, which are gradually worn ava 
finally shed like the milk-molars of other Mammals ; the hott 
the comules being the remnants of the original alveoli. 

Family EcHiDNiD-e. — The second family is characterised 
convoluted cerebral hemispheres ; the production of the mun 
a long tube-like beak (fig. 1150, a); the slendemess of the 
dible ; the total absence of teeth ; and the presence of stout 
mingled with the fur. Further, the feet are very strong, and at 
for digging, and the centre of the acetabulum is imperfectly o 
There are two living species found in Australia and New C 
both of which, at least for paUeontological purposes, may be in 
in the genus .£cAf'i/ffti, although the hige £. Bnnjmii, of New C 
is frequently separated under the name of J'roecAidMa. Ri 
of a large species considerably exceeding the latter in size bav 
obtained from the Pleistocene of New South Wales. 

Group Multituberculata. — In this place it will be com 
to notice a group of very imperfectly known Mesoroic anc 
Tertiary Mammals which were formerly regarded as Dipn 
Marsupials allied to Tkylaojleo, but which differ in several n 
from that group, and may perhaps eventually prove to be mi 
of the subclass Prototheria, Although these peculiar for 
semble the Diprotodonts in having a single pair of lower i 
like those of Rodents, while many of them also appronn 
certain members of the same group in having a secant and g 
fourth premolar ; yet they differ in that it is the second in p 
the first upper incisor which becomes enlarged and c^po 
the incisor of the lower jaw ; while when the fourth lower pr 
is secant the summit of its crown is extremely convex inst 
more or less concave. The true molars, as already men' 
appear to approximate in general structure to the deciduous 
teeth of Ornilhorkynehus, and are quite unlike those of any 
Marsupials. Till, however, the structure of the pectoral gi 
known the serial position of these forms cannot be definiteh 
mined. And it may be observed, that the humerus of th 
which may be referable to Triiylodon is unlike that of the 

BO that if tlwse fornw be Prototheria. they probnbly indi- 
tstioct specialised order of that subclass. 

ition lu the features mentioned above, this group is chanic- 
jy the true molars (fig, 1 136} carrjirg longitudinal rows of 
•i, separated by one or more grooves, and also by the ab> 
r a pit or perforatioD in the m.'i»9eteri<; form of the itiandible. 
uv Pi.A*iiAULACiD.e. — In ihe Plngiauitiddxr the premolars, 
•s in number from one lo four in the mandible (figs, iija- 




iti Owcr iri<* of ikc richi rBinin rf itw nundibl* of PUfianlur mijtpr ; Tour 
H ((■•. I,. FtHftk IsBer pmnoUr of P. BnAtai ; fivt aiul a hBiriimniialunliiM 
VvfaKk of DHMttbin. lAheiOmi.) 

are always of a secant nature, and are usu&lly mjirked by a 
f oblique Literal grooves (fig. 1 1 32) ; whi!e the true molars 
1 and HKluccd to two in number. In the type genus J*/a- 
of the Purbcclt (Upper Jurassic) of Dorsetshire, there may 
four (lig. 1 132), or three lower premolars ; Professor Cope 
tg the Utter va- 
2A indicating a 
genus, for which 
propoiied the 
^ioprion. The 
ceth are unknown. 

>^« <fifr "33). 
Upper Jurassic 

th America, is a 
allied but per- 
, specialised type, 
r lower prenio- 
rhich arc much 
the figured sped- 

In the upper )aw the anterior premolars ore like those of 
1. The figure shows the relatively low position of the con- 
the mandible in this family : — a feature shared with the 
Irene Tf»yiaa>{io, among the Marsupials. From the Lowest 
both of france and North ."Kmerica, we have the more 
Itsed genus JW*9*A»jr/'itJ'/<J.c (fig. 1 134), in which only the fourth 
iUr remains in the mandible. The Puerco Koccnc of North 
also yielded Plilodus (fig. 1 133), characterised by the 

Flit- tij)'^)iJBn atpsEI cjr the rifht r*mu4 of thf inmn. 
dibia of c rnMrnAn nmUiu; fram iht Uppcf junutic of 
Konh ArMrica. Uiiinr Afun, naiirrul iiiio ; luwi). faur 
limtn naiurat uH o, Incuori h, Candyli; c, L'aronond 
tatxHL CAflcr Manb.) 



presence of a minute third lower premolar. Uotomus of th 
Eocene differs from all the others by its snaooth fourth [ 
Afeniscoessus, from the uppermost or Laramie Cretaceous of tt 

Fig. iiu.— InntT vlewof the Irflramiuof the mandibUof JViM^i&V>'"'>'< 

Lowtr Eocene of Rheimt. The line indicate* the Imetue. (After LeontBi 

States, is a somewhat larger form which may probably be in 
this family. It was first made known by an upper true n 
secant premolars from the same deposits subsequently i 

under the name of Halodon, 
with other teeth figured under t1 
of Trifiriodon, DiprodiotL, and 
odon, are probably all referable 
iscikssus. Another form from i 
mte Cretaceous which may be ] 
^^^ ally known as Cimoliomys, appe 

^^^^Bl b closely allied to the Eocene Pti, 

^^^^Hm^ may have affinity with Plagiam 

^^^^^^B^ teeth described as Cime/odon ai 

^^^^^^^^ mys, as well as one of those r€ 

Haiodon appear inseparable f 

form. Other generic names h 
applied to teeth of a more or 1e 
allied type from the same depo 
Certain bones from the Lai 
scribed as Camptamus probabl 
to this group, and are note* 
showing a distinct coracoid a 
clavicle ; thus, if rightly referred and determined, clearly i 
the Prototherian affinities of the Multituberculata. 

From the Tertiaries of Patagonia Dr Am^hino has i 
certain remains under the generic names of AbderiUs, Afn 
Palaotheutts, which are referred to this family, and are a 
nearly related to the European genera. The genus Microb 

Fii. 1135.— The lefi rarom of the 
manoibie of Ptitttiut mfdiinttti 
frorn iKe outer (a), inner (/•), and 
oral (<0 aspects : from the Puerco 
Eocene of New Mexico. (After 



describ«<I rrom the «nine deposits, and is regarded as 
^ & distincl bmily. 

jr PoLYM*STODOKTiDJL — Tlie h-pe genus Pofymaslodon 
I the Lowest Eocene of North America, and haa one pre- 
d two true molars. The prcmnkr is tubercular, and (he 
lie molars have three longitudinal ridges, and are elongated 

jflr TarrviijDONTiix*. — The genus TritylodoH was fiftt de- 
tom a nearly entire cranium found in ttte K^ruu system of 
frica, in a horiiton which i« probably of Lower Mcsozoic 
[ a tooth (fig, 1136) pToiously 
t from the Upper Trias of 
|r, and described under the 
pied name of Tn'gfypius, 

bclor^ to the same genus. 
)ia dental formula » I. 2, C. o, 
Jif. 4. The innermost upper 
B large and scaiprifomi, while 
er one \i very minute. The 
rue molars (fig. 1136) carry 
Itgitudinal ridges, and fiave the 
linDeler of the crown directed 
icly. The anterior portion of 
flum is remarkable for its (treat 
nd bitmtness. The j\fncan 
UKUcatcs an animal about the 
i Rabbit. From the same de- 
\ the Cape has been obtained 
fhowing the impression of a 
ilimb apparently referable to a 
[amnul, whteh has been described under the name of Tfitr- 
}. If, as is proliabty the case, this specimen be really 
llian, there is, however, no reason why it should not belong to 
(m. According to Professor Bardclcbcn this Hmb has two 
il in the carpus (as in some InsL-ctivora), and a distinct pre- 

the alleged fusion of the scaphoid with the lunar being in* 
To the present family may likewise be referred the very 
tdy known SferttrgnatAus, of the I-owcr Jurassic of Slones- 
ilc upper cheek-teeth of which rlo»ly resemble those of Tri- 
• Hnally, CAirvx, from the Upper Juranaic of North America, 
BTtato characters connecting the pre^etit with the next family, 
h it is provisionally placed by Professor Osbom. 
jLV BOLOUOKTiD,*- — Nearly allied to the preceding family, 

1 the upper true molars antcro-posteriorly instead of trans- 
ited, and with only two longitudinal ridges, is the 


TrifM ci SEFivlHibr]!' 'I'bd lHrt> tantraJ 
fictifci an of ihc ruturi) tirr : |b« 
olh«T« atilarcHl itirt* rliria*. if, Crt>wn 

i«n> 1d'tt«J 4iirfAc«4 ; /, •■. Ani+rii^r find 
pOHltnor 4urfat«f. Thv poi^tlon of fi 
It St rithl iLii£l«« t4^ thit of <he fntilMt 



genus Bolodoit, of the Dorsetshire Purbeck. In this genn 
1137) there were apparently three upper premolars and ftm 
molars ; with probably three incisors, of which the first ii 
minute. Although of much smaller size, the cranium of thi 
species presents a striking resemblance to that of the Aincan j 


Fie. 1137.— The righl maxilla of Beledtu trmaiJtm: Tram ihc DoncMbn hrtad 
Maxilla; /"i.r, Premaxilla: ^, Ponunen ; ■.Tnicoolan; fm, Pnmobn; (', lodnn. 
enlarged. (After Gaborn.) 

odon. Allodon, from the Upper Jurassic of Korth America 
very closely allied, if not identical form, in which there wer 
tainly three upper incisors, of which the second (fig. 1137) 
larged, and apparently corresponds to the large inner ind 
Tritylodon. The genus Microlestes, from the Upper Trias <A 
temberg and England, is only known 1 
tached molars (fig. 1138) closely resei 
the true molars of Plagiaulax. This 
is referred by Professor Osbom to thi 
giaulacida : but till it is proved to 
trenchant premolars it seems prefecal 
place it provisionally in the present f 
Figure a in the accompanying woodcut 
a side view of the type tooth, whidi ha 
rows of tubercles separated by a tongiti 
groove ; the imperfect crown of another 
shown in b, and provisionally referred to the same genus, is re 
able for the resemblance presented by its two remaining tub 
to those of the teeth of Ornithorkynchus, The name Hypai 
Hopiis has been applied to a tooth of AticroUstes, from the Ei 

Subclass II. Metatheria. — The members of this subclass 
with the Prototheria in the structure of the brain and the [ve 

rig. ii^S.-n. Lalcial view 
of a tooth of Mkrvtestri 
unliquui; ^.Parlofthecrown 
of a tooth on a «ri]| more en- 
larffed scale ; from the Upper 
Tria« of WOrtemberB. (Alter 
Lyet] and Falconer.) 


bic bones ' (fig. 1 139, «), but differ in the more specialised 

of the aufUtory ouicles ; in the reduction of the conicuid 

prooess of the scapula, and its 

ntlAtion with the sternum ; in the 

of an inteiclavicle ; nnd in the pres- 

nipplea to the mainmai^' gland-s. 

atter characters they agree with the 

■b. fTvin which tbcy arc distinguialicd 

I proence of an imperfect cloaca, 

I the chanctera of the urinogenital 

I which are in «onu; reaperti^ inler- 

||K between those of the Eutheria 

{koUNheria. The young, although 

Bed rit-iparously, arc bom in nn cx- 

gty imperfect condition, and are 

.pourtshed in uUro by an allantoic 

la ; but at an early period are trans> 

[to the nipplea of the mother, to 

tthey adhere for a long time. The 

f themselves are nearly always con- 

y in a fold of skin forming the so- 


arc in>-ariably present, and are 

into tfie usual four groups. With the exception of tlie 

HjriJte, the number of inci&ors in the upper jaw uf all c»ist- 

exceeds that in the mandible* The true molars are very 

iy four in number on cither hide of each jaw; and in all 

fonni the number of premolars does not exceed three, 

p) four (which may be taken aa the i)'pical number) are 

Hin icveral Mesozoic genera. The most remarkable feaiure 

jlhc dentition is, however, that it i» only the last premolar (the 

tof the typical series) among the whole number of teeth that 

■s a milk predecessor; such predecessor generally resembling 

K molars in stniCTure, Some genera, like rhaicch'tnys, show, 

r, no signs of even this single replacement ; and it is pretty 

we have here the first coramcnccment of a replacing scries 

which in the early Eutheria must have gnidu,-illy extended 

tly, until it normally embraced all the teeth in advance of 

LwKh the very general exception of J*m. !, although in certain 

Odactyla even that tooth liad a milk prcdecassor. 

pther very characteristic, although not universal, feature in this 

IfB is the inileciian of tiic angle of the mandible (Fig. ) 1 52) ^ 


OuAtd ia Tiyiotiuus. 

t MBic fnliuc oocurt waong the Anomixlont Itepltles in the GaUiaaridtt. 3 B 

V\z- M_>^— V«nr»l view of lli« 
iyin^>«ui and ir(\ ht\( itf Ih* 
|kIm-i< vF It KufigutM. ■«, Epi- 
publc bonc:!. KqIikbI. 


and in many genera the palatal region of the cmiiuiB oc 
unossified vacuities of considerable size. 

Order II. Marsufiaua. — The whole of the known Men 
are included in the single order Marsupialia, and it is not at p 
necessary to give ordinal characters as distinct from those < 
subclass. At the present day this order is restricted to Ai 
and the Australian region ; the greater number of forms ooc 
in the latter area, while only the Diddphyida are found in the fi 
In earlier epochs, however, this order was much more wide! 
tributed, and it apparently contains some of the earliest 1 
representatives of the entire class. 

Marsupials form in some respects the intermediate stage be 
the Prototheria and Eutheria, and it is probable that there « 
eventually found a complete transition from the Pol3rprotodoa 
order of this group to unknown Prototherian Mammals wit 
same general type of dentition — possibly more or less closely 
to the undermentioned Triassic family Dromatheriida. 

Suborder i. Polyprotodontia. — In existing Polyprat 
Marsupials, which occur both in America and the AustraUan r 
there are never fewer than three lower and four upper incisoo 
there may be five upper and four lower teeth of this series. ' 
teeth are small and subequal, and are followed by a larger c 
(fig. 1 148). Normally there are three premolars, correspondi 
the first, third, and fourth of the typiod series, but the fourt! 
be absent, Dasyurui and Sarcophilus {^%. 1148). The numl 
true molars is generally four, but in Mymutobius these teet 

increased lo - ; and their crowns are nearly always characteris 

carrying a number of small, sharply pointed cusps. In no s{ 
is there a fourth premolar of the elongated secant form charactf 
of many Diprotodonts, Several of the Mesozoic forms included i 
group agree very closely with the recent ones, but many of them 
four premolars, and in some there are as many as seven true tn 
Family Dromatheriid*. — This family, typically represent! 
the genus Dromatherium (fig. 1 140), of the reputed Trias rf 1 

America, is provisionally p 
here, since it may be relati 
some of the members of the 
family, although ProfessorO 
makes it the type of a di: 

V\g. ii40.-lnner view of ihc left nmut of „,J„, ,l„ t> . j -_ 

Ihe mandit'le of Drvma/iirium rxhtilr, .- OrQCr the rrOtOOOnta — 

Enlom' )''''" "' """' *^""' **"" ^^^" suggests that it ma; 

Reptilian. If not Marsu| 
these early Mammals may prove to be representatives of a grw 
Prototheria from which the Polyprotodont Marsupials have 01 



The chcck-Iccth differ from those of all other Marsupials in 
the Eangs imperfectly divided, and thus approximate to those 
Anomodont Reptiles. There arc seven true niolurs, tlie in> 
UK spaced, there is long diastema behind the canine, and the 
; premoUn are of a \xty simple structure. The crowns of the 
taKilars coDsist of one main cusp, with small accessuTy cusps ; 
lil would appear that the teeth of the up|x:r and lower jaws 
[ly interlocked- This type of tooth is regarded by Professor 
as the most archaic yet known. Mkroconcdon, which 
in the same depo&its as Dromatherium, '\% an allied but 
IILV .\iiPHtTHERliD.C — This Nfcsoioic family is provisionally 
to include a number of small and imperfectly known forms, 
of which are re^rdcd by almost all writers an undoubtedly 
, although others have been referred by Professor Osbom, 
some hoiiationr to the Insectivora, under the name of In^cc- 
Pnmitivn ; those which are retained by that writer in the 
order being termed Pnxlidclphia. Although it is rjuilc 
thai with fuller information the two sections into which 
family t« divided may be raided to the tank of separate families, 
^the e\ideiK;e broujjht fomard by the writer mentioned above in 
of referring some of the genera to ihc Inacctivora appears to 
*lti^uflieieni ; no members of that order having more than three 
Iparr incisors or more than the normal three molars of the other 
ital MammaU.' Coniiderable confusion has arisen in regard 
itfic dentition of many of the mcmlwrs included in this family 
to the circumstance tlut in the mandible only one side of 
I teeth u generally- seen, so that several genera have been made 
the evidence of remains of a few closely allied species. The 
rirt^ up of this confusion is mainly due to the careful obscrva- 
tJons of Professor Osborri. In this family all the genera are char- 
klerised by a channel on the inner side of the mandibular rami 
bKiWD as the Myhhyeid gmove (fig. 1 14;) — a feature oncuning in 
Duy recent Polyprotodonts. The number of lower incisors was 
pobably always four (as in the modem Diddphyida), and the 
k«er true m<:^ars are frequently in excess of that number, as in 
MfrtMcMus alone among existing heterodont Mammals. These 
taie molars may either consist of three or more cuspK ,irr.nii^d in a 
kicjlc line, or they may be differentiated into a trituliercular blade 
lollowed by a posterior heel or talon. The premolars are very 
Seocrally four in number, but they may be reduced to three, or 
pofaaps two. Not unfrequently, as in the existing Peramtlida, the 
toot of the canine may be grooved, 

' (oilgine fran hi* lalol memoir on the lulijeci, ii if probuMc that Profewai 
'toami would now conndmbly mcxltfy hi> views its 10 thnc divikionft. 


As » pro\nsional nuasure this family may be divided in 
groups or subfaTnilics. of which the first may be cvciitiuUy i 
the ranlt of a diwinct family. The fir*i subCunily, or fk 
theriina, is referred by Professor Osborn to his ProdkJdp 
is included in the Triainodontida (^infra). The lovrer inic 
consist typically of three main cusps arranged in a line, 


Fix. Ii4].~'li>ii' I - I III III, hi I nil frill nunililili nf r*ii>Wi«rt«f<i» 

from Ihc Siaiwtlirlil suiit i tiv:t niUDn.1 lin. lie oulluM Ggiarc li taianl ■>■ 
ihouU |jea(ou.tili uikImi. (Afiei Oircn.) 

with some accessory cusps; and it would appear that tbc 
niolnrs were of iimilar siruciure. The lower incisors arc 
from one another hy intervals, The typical genus Phc 
(fig. 1 1 4 ' )i f'''^™ t^'*-' Lower J urassic slate of Stuncsfield, in 
shire, has the condyle of the mandible placed very low 
the lower dental formula is / 4, C. 1, Pm^-\-M. 7, the 
separated by a diastema from the first premolar, and the tnaei 
hive a well-marked dngulum on the inner side. AmphiUsiit\ 

Pis< 114a. — Rcvtnvd InDci vie* uT iKc liTl nniinor cli< nundibli a(i.. 
ffom th* ^inne^tlil Stale. THk* nuri^raf tXm. Tba n«i«tailoo ^ Ibt i 
JKIunil -, and On eoodyli ii pUnd loo hi|[h. (AJUi 0<na.} 

1143), from the same deposits, has a more numerous scncs 
chcck'Tceih. and a higher mandibular condyle. The exact doOil 
formula is not known, although it may hAvc been tbc same at m 
the undermentioned Amblotktrium. The mandible from the ime 
beds shown in fig. 1143 ^^^ been niade tht type of the jjeo* 
AmphiiyUii: and according to Professor Osbom has teeiti of '^ 



l^ type as those of the preceding genera. If this be 
appear to countenance the view that the present group 
be separated from the Amphithfriidtr. hut owing to the 
condition of the spcci- 
It camion is necessary ^^^ 
iking positively as to the 
: of the teeth. 
tlic second subfamily, or 
MtJuriimr, the lower true 
arc diifcrcntiatcU into a 

aaCerioT blade and Sion«fKU^is.~ EiaMsed. (.\i^ct'Ow*n.] 
Ipmcrior becl or taloo. Four 

tonolars are present in all those genera of which the entire lower 
BKttkin is luiown; and tlie mandibular condyle is high. The 
jpCT molan are unlike the lower. Many of [hese genera are 
(trred by Professor Osbom to his Insectivora Primitivo. 
Brfore proceeding further some explanation is nL-ccN<%ary aK to 
t ttniciure of this type of lower molar. Following the nomcn- 

Pic. ixi'- Ou(» nAntci nf ihe richt mat* at 
ihe nuBdlUe of ^mMilyima Ovtml; fmin iIk 

^IM. — OUHC Mptci vT lh« Uft riBjln of (hll inin.1il.lii nf lyrjulltltt rpni-r ; rrum tbe 
■xuk of Nordi Anwna. Three lima nalunil •>». «. Caiiiiic : c, Curonaid jicvecM i 
. TW JDCMn sad 6r>( rnmolai on aWiM- (AImi Manh.> 

lt«re adopted by Professor Flower the three cusps in the anterior 
itf of such a "iritubercular" tooth may he collectively spoken of 
I the hiadt (fig. 1145, 1, b, «•); while the hinder part (rf) may be 
imcd the takn. In tbe blade the cusp a is termed the anterior, 


Hf^^Dpp«r, atit*r, u^ iantr *irw« iT a Irfi Towrr irup tniiU.' of 774//»ru. d, Anterier 
r Uu< (/M^atwaiO. t. IVMatior Ciup of do. i^rrlitfniit): t. Inncf cuip uf ilu. i/iitt' 

he large one A the posterior, and the small one <*, which h the 
BDennost, the inner cusp. According to Professor Osbom these 
hnte cusps correspond to the three cusps of Pn'acodon (fig. 1147), 
fid in both cases be applies to ihem the names of para-^ prutO'^ 


and metaconid ; calling the talon of the present type the kff^ 
Corresponding terms ending in cone are applied to the upper t 
of the same types. 

It will be obvious that when a jaw is embedded in matm 
the inner surface exposed we shall only see the cusps a andi 
the talon d; while when the outer surface is visible ooljtiie 
cusp b will be observed. An example of the latter occonegi 
shown in fig. 1146; and this circumstance has been the fti 
source of error in regard to a number of the Mesozoic types • 

In the type genus Amfhitherium of the Stonesfield SUM 
Lower Jurassic, the exact dental formula is unknown, but it 
perhaps have been the same as in the next genos.^ In A 
iherium ( = Peraspalax, Phascokstes) of the Purbedc, or I 
Jurassic, of Dorsetshire, the lower dental formula is /. 4, 

Fig. 1 146. — Outer view of the left nintuof (he mandible of AmbUtlitrium frmdb\ 

Jpper TuraAsic of North Americjt^ Thr« ' . . * , 

^ Condyle; .^ Muidibk. (After Marab.) 

Upper Jurassic of Nonh Ameriot. Thm tinn natorml liic ■, Caniiw; c, Connoii 

hf Condyle; ,""' 

Pm. 4, M. (7-8), or the same as in the American Jurassic 
kstes (fig. 1144). Several species are known, in some of 
there were seven, and in others eight lower molan. The 
Styiodon has been founded upon the outer side of mandil 
Amblotherium, in which only the large posterior cusp (protc 
of the blade of the molars is visible, as in the lower jaw sb 
the accompanying woodcut, which has been made the type 
genus Stylacodotty of the Upper Jurassic of North America. 
latter has eight lower molars, while the English form ha; 
seven. Achyrodon^ of the English Purbeck, is closely all 
Ambloiherium^ but differs in the form of the cusps of the i 
Peramus {Leptocladus) is a third Purbeck genus, with rel 
stouter lower molars, in which Professor Osbom gives the 
dental formula as / 3, C. i, Pm. 6, M. 3. The North An 
DryoUstes has relatively shorter lower molars (fig. 1144), the 
cut clearly showing the three cusps of the blade and the tJ 
these teeth. Aslhenodon and Laodon are other North An 

' ProressoT Osbom regards it &s /. ?, C. I, Pm, 5, M. 6. 



oncterUed by the iunall size of ihe lalon of Ihe lower 
while the names DitnxynodoH {Difi/wyHffdan), Doa>doH^ 
ttcdtm 1 have been applied to more or less closely allied 
XD the same depuait:(. Finally, Trofcs^or Osburn hns 
t name Kurttnion lo certain upper jaws from the Engliiih 
wfaicb are probably rcrcmbic to one or other of the aliovc- 
d genenu From the Laramie Cretaceous of North America 
Marsh has described the remains of allied types. One of 
been termed DidtipHops {Didfhfhodon or CinmlenUs), while 
I has been referred to the Jurassic gtin\i> DryolesUi, and 
las been made ihc type of the genu* Fuiwmys. The re- 
e in the structure of the lower molnri^ of ihe AmphitfuHina 
'the corresponding teeth oi Dmyurns and the Didt/phyida, 
ihat the latter have originated from an allied stoclc. 
r Spalacotheriid*. — The genus Spa/aw/Aen'um, with 
TaUsln is identical, has a dentition which may probably 


Esed by the formula /. -, C. ^ 'Pm. ^, M. \. 

3 f 4 6 

ITS consist of a single column carrying three cuspii, and 
•sponding to the blade of the tooth of the Amphitheriintr. 
iction in the number of the lower incisors diKtin^uishes 
ly from the last; but it is difhcull lo say whether (lie 
if the lalon in the lower true niolais is or is not a more 
d feature. The molars, which are of the typical trituber- 
r, approximate very closely in their plan of stmcture to 
the genus Chrysochlfirit among the Insectivora, but this 
C taken as iiKlicativ'e llial the present genus should be 

that order, since a precisely analogous resemblance cx- 
«i the molars of Tupaia in the Inscttivora and Ptrameles 
irsnpials. Menaeodjfn^ from the Upper Jurassic of North 
is an allied genua. 

r Trjcokodontid-k. — The last family of MesoKoic Mnm- 
have lo consider is represented by the English Purbeck 
VifCHffdon (Triaeanthodon), and the allied or identical 

{fig. ti47) of the Upper Jurassic of North America. 

Osbom includes in tJiis faintly the Phauolalheriitia 
tentioncd ; but the reduction in the number of the in- 

1 the general fatiti of the teeth seems to indicate con- 
difference — although the two groups arc probably more 
early related. The mandibular condyle is placed still 
n in PhoitoUiMerium, and a|)parently, indeed, than in 

knonn Mammal. The upper cheek-leeth resemble ihe 
'be dental formula in the mandible is / 3, C. i, Pm. 4, 
The inciKors were approximated, ihe cheek-teeth have 
' Prcoccujiicd, Mc page loi;. 


an inner cingulum, and the molars consist of three subeqiial i 
or cusps of a trenchant fonn, arranged longitudinally, lixtei 
apparently either three or four molars in different indiiidiiab < 
single species. The fourth premolar was preceded by a wlkA 
resembling the true molars. It was considered by Sir R. 
that the lower true molars of this genus corresponded to tbocl 
the existing Thylacinvs, in which there is a bilobed bbde udj 
talon. According, however, to Professor Osbom, this inter 
is incorrect ; and that observer considers that the three cusps of t 

Fig._ii47.— Inner view of the tcrt rarnui of tbc muidiblc of /VufwAx jferaur ; fraa AtQ 
Turauic of Nonb Ajncrico. Three tirno natuTAj liie. c, Cofvooiil ; f, Hykj^roU fnop 
Symphysis. The Anterior teeth ue wnnting. (Af^er Mwh.] 

Triconodont type represent the three main cusps of the tootb 

Phascohtherium (^g. 1 141), and also correspond to the three 16 

of the blade of the tritubercular molar of the Amphitfurium t 

(Bg. 1 145); and he accordingly terms these three cusps the /« 

proio-^ and metaconid, and considers that the talon Qiypocomi^ 


Family Dasyurid.*;, — With this family we come to the considi 

tion of the existing Marsupials. The Dasyurida is an exclusii 

Australian family, comprising the largest known members of 

suborder, and is divided into the subfamilies Dan-nrina : 

4 t 

Myrmecohiina. These are always / -, C - ; but the numbe 

cheek-teeth varies, although there are never more than three ] 
molars. In the pes the hallux is usually either rudimental 
absent, but the other four digits are well developed and subequaL 

the Dasyurinee the number of cheek-teeth does not exceed {, 

upper true molars have triangular crowns, and those of the lo 
molars are differentiated into an anterior blade and a posterior ta 
(fig. 1 145), like the lower carnassial tooth of many of the place 
Carnivora of the present day. The mylohyoid groove maj 
present both in this and the next subfamily. The Tasmar 
Wolf is the sole living representative of the genus Thyiaciimi 



cheek-tcdh number Pm. -, M. -, &nd Uic hutncnui has 

3 4. 
but in Ifae Pleistocene of Australia there occurs the 
iWy larger 71 ifit/irus. Sarmphi/us, of whirh the dcnlition 
in fig. 1 148, is also conned at the present day to 'L'a»- 



|ti,a.~I^ri bt(*il *lev of Uw rfentilion ^r Sarafklin unimttM. Recant, I'urni 
ban ; c, C^bibc ; /m, PnaisUn ; m, MoUn- ni lootli niul«J /■• i i > lulljr 

'urnanla. i. 

but a larger ^lecies inhabited the mainland of Australia in 
Pfcistoccnc. Indications of specialisation arc shown Ly the 
ction of the premolars to two, and also hy the loss of the fors- 
\a ihc humerus. The most generalised genua of tlie subfamily 
tms, compnsing sevenil species of ^mnller Ktje tlian the pre- 
In the lower molars (fij;. 1145) the blade has three cusps 
in a triangle, and thus differs from those of Thyhidntn, in 

the inner cusp ts wanting. There is a mylohyoid groove in 
CauncKblc, and the humcnw has a foramen. Species of Dasyurus 
iihe present day range over the whole of the Australian continent, 
neuf them dating from the Pleistocene. The subfamily Afyrmteo- 




biina is represented solely by the genus Myrmtcobius. lo I 

markable animal (fig. 1149) the lower incisors are sepantedl 

one another, the cheek-teeth number ~, of which the fint 

either jaw are premolars ; while the molars have quadiai^piliri 

cuspidate crowns, and are not well differentiated from the 
Although some writers doubt the connection, it is probable 1 
Afyrmecobius is the direct descendant of forms closely 
AmphiUsUs^ of the Lower Jurassic of England ; the relations of I 
two being perhaps somewhat similar to that existing between 1 
living Sphemdon of New Zealand, and the more specialised 
odapedon of the English Trias. 

Family Pbramelida. — In this family, comprisiiig the 

of Australia and New Guinea, the dental formub is I. --_ , C 

Pm. -, M. -, and the pes has two of the digits reduced and 1 

3 4 
nected by int^piment, as in the Macropodida among the 
donts. Occasionally the canines have grooved or double 
Remains of existing species of PerameUs and Ptragaie occur in 1 
Pleistocene of New South Wales. 

Family Didelphvid^ — In all the niembers of this lamilyl 

dental formula is /. -, C. -, Pm. -, M. -. The incisors are 

4 I 3 4 

small and sharp; the canines are large; the premolars compresMd^ 
and the true molars constructed on the general plan of those 1 

Fig. 115a — Left lateral view of ibc denlilion oC DidtlfMjit Ammrr; Sooib Umiriii t. Iv 
ciiois; r, Canine; fm, Prenialan : m. True molin. lie mili nuritcd >« j •booM ta/B«i 
the leeth beiweeti ihcK and/jH i being /m i. 

Dasyurus ; the lower ones having a blade and talon, with a distinct 
inner cusp to the former. Each foot is furnished with five com- 
plete digits i and the humerus has an entepicondybr fonunen. .\l 
the present day the Opossums, as the members of this femily m 
commonly called, are confined to the New World, where they m 
represented by the genera Didelphys (fig. 1150) and ChinmeAi; 



being known only by a single species. Remains of cxiist- 

of both genera occur very cummonly in. the Pleistocene 

1 of the Bmilian caves ; and to Ihe ij-pe genus may also he 

a large number of species from the I-owcr Miocene and 

of liurope, which by some writers are referred lo 

genera under ihe names of Peratherium and Amphipera' 

S^liuodon, of the Eocene of Hordwell, i^ probitbly a 

It <*a& in DiJtlphis Cuvicri, of tbe Paris g>-psuin, tbut 


l.^Did»(fSvff)^t»i: humllM While-river Miocen* (rftolDnido. Tn-ice naiur»l 
I litfenor ^i blew *>e*i «f tkiill ; t. 


J, 3u|ierl«r uid laivnl viev* of rigtii mwi- 

demon&tratcd the existence of matsupial bones by a careful 
[ng of the matrix. Other extinct forms from the Miocene of 
^America (fig. 1151) may in all probabiiily be referred lo the 

KitDeR 3. DiPROTODOKTiA. — If we exclude the Multituber- 
ft which have been already menliojied, tliiD :suborder will be 
Bed to the Australian region, where it has been known since 
Pleistocene; and with this limitation it may prob;ibIy be re- 
, as an offset from the more generalised Polyprotodoniia. In 
there is only a single pair of lower incisors, but in the upper 
are usually three pairs of such teeth, although they are 
10 one in the Wombais. 
lower indMnt, and the first, 
inennosl, pair of upper in- 
s are always of large size and 
ted for cutting. The canines 
^uently absent, and when 
t arc of relatively smal I size. 
3wns of the true molars 
tSer tubcrculatc or have 
nrte ridges ; and as a general 
are not more than two 

. Very frequently the last premolar has a long and narrow 
, with a concave superior border adapted solely for cutting. 

Fif. .ijiv— Voticflor r<iilii-"f vi«» nl ih* 

1 382 


Family Phascolomyid^ — This family is now rep 
by the genus Phas(o!omys, or Wombats ; in which tlie dcnti) I 

is /. -, C. -, Pw. -, Af. -, and all ihc tcclh grow ftxwn 

putps. The tnie molars h.ive curred crowns cron»i5iing of i 
equal lo))es, while the premolar has only a single lobe^ ii>d ! 
preceded by a milk-toolh. The mandible (figs. iis>> 
characterised by having a pit and perforation in the 
fossa. The fore and hind limbs arc of equal length, [he I 
being of great <itrcngth in accordance with the fos&orial 
the genus ; and the humcni)i has a foramen. There are kn* 
in the manus, all of which are proindud with long curved ctiKi,! 
are of subcqual size ; but in the pes ihc hallux is iinpcifcd,! 
the three middle digits are of nearly equal size^ and poitlf 1 
in a common integument. Three existir^ species of Woabni 
known, which are divided into two groups according tu lbc< 



actcrs of the skull and teeth ; none of thetn being of large fiic. 
the Pleistocene of Australia we mcci with remains not only of 1 
existing species, but also with several extinct tj-pes, one of 
was of considerably larger dimensions. In the wimc deposit* 
also found remains of the extinct genus Phaaolonus (with 
the ao-called Sctpamodon appears to be identical), characterised 
certain peculiarities in the incisors. The one known tipedes ' 
ably attained the dimensions of a Tapir, although of consiii 
stouter build. 

Famii-v Ncitothkriiu.*:. — Thi.s family is represented onlybfl^ 
singlt; delinitcly known genus from the Australian Pleistocene 1 
scribed as NotoHurinm.^ This includes one (or possibly mort) iMtf ' 

) It tiiui rncntly l>ct« !iu|,i;<^l«il thai llio fi|iur<'<1 sl^ull Atr* oM l>cipn{ t«/B 
tktrium, Mil thai il abovld Iw tcrmeil Zygtm^umn \M thu \i«w U ncAi 



hd appcan. to lutvc becti allied in many rcspecU lo the 
although presenting several of the dental characters of the 

Dy. The dental formula is / ■*, C. -, Pm. - , J/: - ; and 

thai, at leaM normally, there was no deciduous milk- 

The check-teeth are rooted ; the crowns of the true molars 

two simple iranavenie ridf^es. The cranium (fig. 1154) 

a *«ry singular contour, the nasals being transversely ck- 

: and the mandible diflcrs rrom that of the Phascalomyidm 

u— LiA bMnI *i*w tA lb* dcull -if StUtki'ittM ,\fittktlH: from ihc Pltijio«ne 
oT AsfinlU. Om-uitb naiunl mt. (Attu 0<nn.) 

absence of a pit or perforation in the masseteric fossa (fig. 

(). The limb-bones apjiear, however, to have resembled thotte 

\it latter family; the humerus having a distal foramen, and 

cvidenily adapted for fojsorial habits, although it is difficult 

that an animal of tuch coinpanitively large bulk could 

|ved in bunuws. 

ILV iJiPROTOuOKTlD*. — Thc gcnus Diprotadcn, of the Aus- 
Plelstocene, is the wlc rcprebcntativc of this extinct family, 
type species (lig. 1155) is the largest known mciuhcr of 
vder; its bulk being fully equal to that of a large HMnoeeros. 
dental furniub b the bsmc as that oi NoMhenum ; and Uie 
tore of the cheek-tcclh of thc two genera is also very similar, 
g^ the lower true molan of I>iprotfJ<?i hat'e no median 
Eliiul bridge. In the incisors of this genuiv the first pair are 

I \ff MiSdcnt ciUencc. A tnuU Notalheiold fmm Queensland hu re* 




scalprirorai, and grow from fK-rsistcm pulp* The (ate ■ 
limlK are of approximately equal length, and ac 
walking ; the humerus has no foramen ) and it is prat 
covering of the toes approximated to the nature of 
mandible, although marc convex below, is not unlike 
tMerium. Professor Huxley has named a second 

IHf. iru.— Left knnl vicv of itic iluU al Vifrfta^m Amli-mlii : f>aa cka I 
of Awlmlu. klufh mliKed. (AfurOwn.) 

evidence of premolars, but it is not certain that this deterai 
is corrccL 

Famii-v PHAi^N(iKKiD.c — Thii taxaWy incltidea the 
Fliatangers and the Koala [PAascolantus), as well M • resu 


extinct genuH. All these animals have /. ^, and an upper ■ 
quently also n minute lower canine ; while the premolars on 

2 3 I ' 

from to , aiid the true mohrs from - tu -. The stnid 

1 3 i 4 

the cheek-teeth is subject to great variation in the different g 
Xhii fourth premolar being either secant or lubcrcaiar. Tha 
pit or perforation in tlie ma.<»(.-tcric fossa* of the mandible- 
limbs arc of nearly equal length ; the manus has 6ve subeq' 
but the second and third digits nf the pes are very slend 
partially united by integument ; and the hallux is always op() 
In many of the true Phalangers the founh premolar is groon 
the dentition closely resemh!e<i that of the existing geous 
prymnoden among the Mucropodidtt. Of living genera the o 


D occur in a fossil state is Puud/ykim, of which the remains 
species are found in the Pleistocene cave-deposits of 
Wale*. The most interesting member of the family is, 
►the large Pleistocene TAy/aco/to (fig. 1156), which foms 
^of a distinct subfamily. 'Ilic dentition may be rcpre'senlcd 

lula / -, C. -, J*m. 5, Af. -. The true molars and 
I o' 3' 2 

loUns were small, and more or less functionless, while 

premolar is enormously developed, and has a long sharp 

s, so thai, in union with its fellow of the opposing jaw. 

cfAiuirella. Uix-flfib nalural tit«. 

a cutting instrument of extraordinary power. In originally 
this remarkable animal from fragments of jaws cuniaining 
MDth premolar, Sir Richard Owen came to the conclusion that 
Iructurc of this tooth indicated a carnLvorous animal adapted 
cy upon the huge Diprotodons and Xotothcrcs; but the dis- 
-y of the complete skull has shown that the animal was more 
\j allied to the existing Phalangers, and that it could not have 
lascd the destructive habits atliibuicd to it hy iu dcscriher, 
1^ it is quite probable that its diet may have included the 
ler numntalK, birds, and ^:g&- It was at one time considered 
the Multituberculnta were allied to this genu.^. 
ILV MACKoroDiD^ — The last, and in many reapects the 
:iaUsed fiimily of the Diprolodomia includ&s the Kan- 
Waltdiies, and Kangaroo-rats. The dentition is represented 

Ibrmula /. 

O 3 4 

The incisors are 

Jy secant, those of (he mandible being frequently movable 
. one another. 1'he premolars may have cither triangular or 
crowns, and in the latter case (5g. 115S) arc frequently 
icd ; ihe third premolar is always, and the fourth in some cases, 




deciduous, the latter being invariably preceded by a milk-toa 
true molars have either four tubercles or two transverse rid 
frequently an anterior talon. The mandible has a deep pit 
foration in the masseteric fossa. The pectoral limb is ain 
or less markedly shorter than the pelvic The manus is ] 
with five subequal digits; but in the pes (fig. 1 157) the: 
generally absent, the second and thin! digit 
very minute and enclosed in a common inti 
(syrtdacfy/ous), while the fourth is greatly enlai;i 
forms ^e main base of support llie ma; 
forms progress by making enormous leaps b 
of their powerful hind-limbs ; but a few, like 
/agus of New Guinea, are arboreal This fao 
be divided into three subfamilies. In the 
Hypsiprymtu^nHna^ there is a distinct hall 
the dentition closely resembles that of the 
gerida; the fourth premolar being small, t 
grooved, and directed inwardly at its anterioi 
In the small existing Hypsiprymnodon there is 
between the lower incisor and the fourth p 
but in a large extinct form from the Pleisl 
New South Wales, described under the nam 
elh, there is a minute tooth behind the lowe 
corresponding to the tooth in the Pfia/aHgtn 
monly reckoned as the representative of th< 
This genus, therefore, forms an important link 
the last-named and the present family. In th 
subfamily, or Potoroina, the hallux is abs) 
first upper incisor is narrower and longer than 
the others (fig. 1158); there is always an u] 
ine ; the fourth upper premolar is elongated and secant, am 
with the grooves strongly marked ; while the true molars a 
culate, with the fourth smaller than the third. TTie Kang 
are divided into the genera Potorous {Hypsiprymnui^ I 
.and jEpyprymnus ; the latter being represented in the Pli 
of New South Wales by remains of the existing species. 

The third subfamily, or Macropina, is distinguished froir 
by the following characters : The cutting-edges of the uppe: 
form nearly a straight line (fig. 11 59); the upper canine i 
either absent or very small ; and the fourtli premolar, whict 
shorter or longer than the flrst true molar, has either an ini 
ridge or lobe. In the existing genus Macropus (Kangai 
fourth upper premolar has a sharp cutting outer edge, and 
ridge or tubercle ; and this tooth in both jaws may be eithi 
or shorter than the first true molar. The two rami of the 1 

Fig, 1 157. — 
I>aT3al Mapect oi 
Ihe rigbi pes of 

tutii. Reduced. 
(Afler Flower.) 



anchyloscd together, and the hind-liml)s are much longer 

fiont ones, A large number of fossil species occur in the 

lEan Pleistocene, among which may be mentioned several 

»e forms like M. brtkus, vrhich have the fourth premolar 

Chan the first true molar, nnd are allied to the small existing 

M. ualabattts (which also occurs in the Pleistocene). 


.njl.— DwlUi«a4l/MtfiMU. /, ladun; r, Cuiiic;/H, tVtmcW; h, HoIu^ 

, again, we have in the same deposits remains of the existing 
l-Kangaroo, M. \PtlrogaU) ptnUiUalus, and also of the larger 
<tstMS, and some allied extinct forms. Another group, in 
the fourth premolar is very small and soon falls out, is tepre- 
by the existing M. gij^antem, .itid the Urgcr extinct M. titan 
M. firm^s. Stfunurui is an entirely extinct genus character- 
by the prcsetwe of a distinct inner lobe to the fourth upper 
ir, and is represented by a Mngle species of considerable 
In Protcptodon^ again, which is likewise extinct, the fourth 


Ptg. ii3» — Miitnfm Oemmtti, L.ueial vkw o( tkull ; riam AlutnUa. RoJimmL 

fiper premolar is like that of Sthenunts, hut the rami of the main- 
Bbie become artchylosed together in the adult : more than one 
pccics arc known. Finally, the extinct PahnhesUsy comprising 
be largest kitown member of the family, is distinguished from the 
ist-named genua by the longer mandibular hymphysis, and the 
bsence of an anterior talon in tlie upper true molars. The length 



of the skull of the one known species is estimated at as m 
sixteen inches. 

Of Uncertain Ordinal PosiHcn. — Here may be conveniently i 
some minute Mammals, mostly known by teeth or fiagnit 
jaws, from the Lowest Eocene of Rheims, of which it is diffi> 
say whether they are Marsupial or Placental, although it is 
probable that some of them belong to the former div'ision. i 
these may be mentioned Trieuspodon, with teeth resembling 
of Spalacotherium ; the allied Orthasfidotherium and Pleun 
tkerium ; and Fro^niciis, in which the true molars approxia 
those of Amblotherium. 



CLASS MAMMALIA— continued. 

Orders Edektata, Cetacea, and Sirenia. 

DBCLASS II, EuTHERiA. — The whole of the remaining orders of 
timmalia are grouped together in a single subclass,^ which is 
hnacterised by the foetus being nourished in utero by means of the 
■■tenial blood passing through an allantoic placenta. This sub- 
hss is sometimes termed the Placentalia, but more generally the 
Eutheria. Throughout this subclass the urinogenital organs are 
mvided with an external aperture quite distinct from that of the 
fimentary tube ; the corpus callosum of the brain is well developed ; 
here is never any marked inflection of the angle of the mandible ; 
ud distinct epipubic bones are absent in the pelvis. With the ex- 
ception of the three orders forming the subject of the present chap- 
to, the dental formula can always be reduced to some modification 
ofthat given on page 1249, At the present day the various orders 
fcive become so well differentiated as to render their definition com- 
puatively easy ; but fossil forms indicate such a close connection 
between the majority of them, that such definition becomes fre- 
<tDently a matter of extreme difiliculty, if not an absolute impossi- 
biUty ; and it is to be remembered that it is entirely due to our non- 
icquaintance with forms which must have once existed that renders 
nen these imperfect definitions practicable. 

There is at present no conclusive evidence of the existence of any 
iwniber of this subclass previous to the Eocene. 

Order Hi. Edentata. — The Edentata are widely different from 
ill oiher existing Mammals, although there are indications of affinity 
to certain extinct forms mentioned in the sequel under the heading 
of the Tillodontia. Almost the only common character presented 
oythe various existing members of this order is that the teeth, when 

li liM, indeed, been proposed lo form a separate subclass — PaTatheria — for 
'* Kuptioa of the Edeniaies. 


present, are devoid of enamel, are never developed at the adi 
ties of the jaws in the situation of the incisors of other Mam 
and are always homceodont and grow from persistent pulps ; i 
with the exception of one genus of the Dasypodida, they are 
wise monophyodont. It has, however, been recently observe 
Dr Ameghino that enamel was present in the teeth of certain \ 
American fossil forms ; while in the genus DiadomuSy from 
deposits, a pair of canine-like teeth occur in the symphysis c 
mandible. In many of the genera the teeth are simply cyliiu 
but they may be transversely ridged, and occasionally they have : 
complex internal structure. Not unfrequently the maxillary 
sends down a large descending process in the zygomatic arc 
II 63 bis); and certain members of the order are remarkable as 
the only known Mammals which develop a bony exoskeleton, 
cer\-ical vertebrae are short and wide, with nearly flat terminal 
to their centra. 

The distribution of the Edentates is very restricted. In E 
neither at the present day nor in past times is there any kno 
presentative of the order ; the so-called Macrotfurium bein| 
known to be identical with the Ungulate genus Chalicotherivm, 
Ancylotherium is also allied to the latter. In tropical Asia 11 
the Pangolins or Manida ; and in Africa the Orycteropodida, 
America is, however, the headquarters of the order, which is 
represented at the present day by the Anteaters, the Sloths, ar 
Armadillos, and in past epochs by the huge Ground-Sloths ai 
Glyptodonts. The gigantic size of these fossil forms as coir 
with their existing allies of the same area is paralleled by t 
stance of the fossil Diprotodont Marsupials of Australia. 

It is evident that the Edentates are widely separated frc 
other existing Eutherians ; and Professor W, K. Parker, in v 
the tendency to a variation in the number of cervical vertebr 
other features, has suggested a separate origin from a Prototl 
stock. Professor Cope, however, looks upon the order as all 
the Tillodonts, and the occurrence of enamel in the teeth of 
fossil forms may support this view. 

Family Orvcteropodid-«. — The Ant-Bears {Orycteropn 
Africa are characterised by the body being covered merely ' 
few hairs ; and by the numerous teeth, which are of a remai 
complex structure, owing to the presence of a number of v 
pulp-canals. In the fore-limb the pollex is absent, but the 
limb has five digits. The femur has a third trochanter, an 
terminal digits are provided with moderate-sized claws, suital 
digging the burrows in which these creatures dwell. At the p 
day Orycieropus is mainly characteristic of the Ethiopian i 
although one of the two species ranges into Egypt. Till vi 



naihing was known of tli« patxon to logical history of the 
ff but l>f FoTsj'th-Nfajor has recorded a species from the Ix>wer 
of the isle of Samos, in the Turkish .\rchipeIago, duttii- 
from the existing species by ihe largw size of the lateral 



-- '^. 

'* -- 


tht- x^6c—t»anlvUvorthtikllaa((^rJKarv^t4^fra^^^. Afttci. Reduced. 

The occurrence of this species seems to point to the 
elusion that A^a was ihe original home of the faniiEy. 
Famii-V >Iasii)*. — The Pangolins (.Viii/y) of Inrfia and Africa 
re distinguished fmm all other Mammals by iht: body being covcitrd 
riorly with .i coal of iinbricited, homy, epidermal scalt-s. 
arc absent ; the limbs arc short and furnished with Five digits, 
icb the terminal claws aie long, cunTd, and biAd at the extrc- 
The humerus has an cnic|>icondylar foramen, but there is 
I third femoral irothantcr, and clavicles are wanting. The large 
l/imj giganka of Western .Africa is found in a fosMi state in llie 
nnstocene cave-deposits of Southern India; while in the Ixjwer 
niocene of the isJe of Samos we have a species three times llie siie 
if the btier. which has been made the type of the geims Palaomams. 
K pbatar^eal from the Indian Siwaliks de.icrihed as Manis appearti 
to belong to Chaluothtrium. 

Family PAsvnot>iDA — The Armadillos (fig. 1161) of South 
America are ch-iraclcriscd by the presenLC of a bony dorsa! carapace, 
composed of a scries of dermal sctites, of which a certain numlwr 
are always arranged in movable bands, while the others may he 
Articnlated together into solid scapular and pelvic bucklers, as in fig. 
1161. I'hc fiontal region of the skull also has a hu>rklcr ; while 
the tail t^ defended by rings or tuliercles of bone. In the cxittttng 
Ktnen the teeth are simply conical \ and in Tatusia nil except the 
brt have milk predecessors. Many of the cervical verlebroe are 
Jnch>-iosed together; and the stout humerus has an enlepicondytar 
fonunen, and the femur a third trochanter. The fore-feet are ]iro- 
Tidcd with very strong curved claws ; and, like the MamJ^, the 
ocisting forms are of burrowing habits. The I'leistocenc cave- 
of Brazil yield remains of soDic existing and some extinct 



iipccics of the genera Da^-fius, Talusia, Tolypetttes (fig. \\h\\ 
Xenurus. In the Pleistocene of Aigciitina we uiect «uh 
extinct type known a!> Eutatus, which is chancteriscd by ibc' 
of ihe cirapace consisting of mo\'ab1e bands, which ore ih[rty4 
in number. The Tertiaiics of the same region have oIm pciT 
much Larger fonn known as Dasypothtrhim, which appears loi 
ncct the living forms vrith the next gvnu^ ITicre were ei^hi 
teeth, of which the scL'ond appears to have been entiiged. 
bomc spccioii of MyiodoH. 'Ihe most reiu^luble genus it, I 

rifi. ittt.- 

(AfMr Murfa.) 


ChlamyJothfrtHm, of the South American Pleistocene, in 
teeth appioximale in structure to those of the next famtlf. T 
carapace has several movable bands ; and the largest spedei »» « 
sidcrcd to h»ve equalled the bulk of a RhirMxeros. The onsd 
AmiacIiUos are therefore dw-arfs by the side of these huge al!i» 
an earlier epoch. 

Familv Cuvptodontida. — In this extinct American fatmlrt 
body was covered by a carapace as in the Armadillos ; tnt 'i 
carapace (fig. 1163} has no movable band^, so that the ailtt 
could not roll itself up ; and mhcc the forefeet have short tbi 
toes, it is evident that the habits of this group were rwt fouoti 
The carapace usually has iu component scutes united by saui 
but in one genus they were separate; the scutes are, morw* 
usually omamenled with a sculpture, which varies in the diflctt 

tityftHdn rrtAmtUmt; Tmni die Plcitiocfnc of 

lh« bul u enclosed in a complete bony sheath. 
larr ~ tn number, and have two deep grooves on either side 

Ing thcin into three nearly 

lolws (fig. 1163); the 

: name bcinjj derived from 

grooving or fluting. The 
ial pan o( the cranium is char- 
|cri«cd !>y iw ewreme shottness 
[, 1 1 63 fit's) ; and there is a 
Ig descending m-axillary pro- 
is in the z>gomatic arc}i. Nearly the whole of the vertebral 
Ittmn is anchylo««d into a long tube, but there Is a complex 
im at the baiie of the neck. Thia 
Btily is mainly charactefislic of 
nnh America ; but species of 
fyftodim rsngcd as far as Mexico 
td Texas into Nonh Amciicn. 
Rat confusion has arisen in re- 
KCt to the classification of the 
l]rpiodonti>, owing to the diffi- 
llty of referring isolalcfl caudal 
leath!! to their proper carapaces. 
he fonns with a solid carapace 
ay, bowev-er, be ananged as fol- 
m. In /fc/l^Aervt the scutes 
r the carapace are sculptured, and 
Flea comparatively thin, the peripheral scries being flat ; while the 
tuda) sheath has several mavable rings, and terminates in a long 


Fia. ttiitft.—lutt Uicril vt(w of the 
ukullor Cly^tttlfm ; htim ihc Plciitonnt 
cf South Amcilc*. One-Unlh lUilunl 
UMt (After Jlurmciitn). 



subcylindrical tube (fig. 1164), ornamented with a number of! 
disks, surrounded by a series of much smaller ones. It ii 1 
sidered probable that the caudal tube represented in fig. i^ 
l>elongs to this genus. The humeras has an entepicondylar : 
and there are four complete digits to each foot. An allied I 
from the infra-Pampean of Patagonia, has received the 
Pahfhoplopkorus ; while the terminal tube of a caudal sheath \ 
Uruguay has been made the type of the genus EUutk 
The latter specimen is characterised by its loose attachment tol 
enclosed vertebrae, and by the great number of perforations 
bristles, so that the tail of the living animal must have 
a huge bottle-brush. The genus Panoekthus is characterised bj I 

Fig. 1164. — The incomplete lerminBl tube of ibc caudal ^lutb o( Htfl^ktna; frmtle 
Pleijttocene of South America- One-thinl oaiurai vae. 

excessive thickness of its carapace, the scutes of which are tuberct ' 
lated, and hy a caudal sheath composed proximally of seven! 
movable rings, but terminating in a long compressed tube onu- 
mented with tubercles, of which some were of very latge dimensions, 
and marked with a radiate sculpture. In JSuryums the caud:! ' 
sheath is of somewhat similar type, but the scutes of the carapace 
arc simply rugose. Dadicurus, again, also has the scutes of the 
carapace rugose, but the terminal tube of the caudal sheath ii 
enlarged into a flattened club-like expansion, covered with coine 
tubercles, interspersed with a few larger rough disks having a radiatt 
srulpturo ; these disks having probably been surmounted, is in 
Ptiiioihthtts, with horny epidermal spines. The type species at- 
tained a length of about 1 2 feet. Finally, in Gfyptodon {fig, iifii), 
with which Sckisiopkurum is probably identical, the scutes of the 
carapace had a rosette-like sculpture, the peripheral ones heing 
raised into conical jirominences, and the caudal sheath, at lea* 
in several species, was entirely composed of a series of raortWe 
rinys, ornamented with large conical tubercles. 'ITie humerus «a* 
devoid of an cntepicondylar foramen ; and while there were fi« 
complete digits in the manus, those of the pes were reduced to fbtir. 
ThortKdphorus differs from all the foregoing in having the scutes c' 
the carapace separated from one another, and thereby approximates 



of the MesatheriidiT, in which there were a number of small 
I embedded in the dermis of the dorsal region, A similar coti- 
revailb in Cariodermtiy of the Ixjup-l-'orlc beds of Tcxait. 
It,* MvRM»xopHA(;in.n. ^This family comprises llic true 
of Soutli Anieticn, re[;reseiite<i by the f^cncia Myfmeoh 
Titmandutt, and Cydoturui: but appears 10 be unknown in a 
state. The javrs arc entirely destitute of teeth ; the body is 
Iwd with httir : the tail n long ; there are either four or five digits 
be pes ; and the third digit of the mantis Is the longest. 
^ll.v Mt'iATMlKiio*. — The mcnibers of This family are en- 
^kxtinct, and arc conAncd to the New World. They comprise 
Htber of \vrt lai{[e fonnii adapted solely for walking on the 
nd, and showing in their skeletal organi:ia(ion characters inter- 
liate between the preceding and the following families. Thus, 
le their vertebrae and limbs are constructed like those of the 
vfhdtpJ*t^ their crania and dentition resemble those of the 

^ — ATccKfAman amttriiamM i Troni ilu Pltitnotasof Sovih Auii^ca. 
Mucli Kdvwd. 

Udut, One species of the genus Sctiiitolhenum approxi- 
ie|^ howerer, in cranial chanclers to the foiincr family : and it 
Kbable that th« whole three families have originated from a 

^ *toclc. The number of the teeth is usually ~ ; and the firsl 

I second teeth may be cither in apposition or separated by an 
The femur has no third trochanter; and the under sur- 
tbe odontoid process of tht> axis vertetira presents a peculiar 
surface for antculation with the alia*. The type genus 
'gaiherium ' is found in the Pleistocene of both South and North 

TUt nsme itwitld properly be AftgaUikrriitm, but in aniiquiiy rcndcrt li 
mkM HCicd. 



America ; tbe typical M. amerUamim (fig. 1 1 65) or the fontierl 
being fully equal in bulk lo tlic largest spcdcs of Khiaoccnb 
teeth (fig. 1166) conKUt of square prj«tn», wearing into 
ridgca through Uic lue^nce of two %'crtical plate* of hanl 
inierolated between KOftef dentine and cement ; ihey ate do 

Structure, and arc all in conlaci. The feet arc ptx>vidcd itilh : 
ful and huge claws, the third digit in each Foot being the 
and the humerus lias no fommcn. 

There arc indications that the 3i»at was prolonged, and xom, > 
flexible ; And the tongue uns probably prehensile. From the clai 
of the molar teeth it is certain thai the Mcifnlhcrc was purely ticrh*' 
in il5 habits ; and from the enonnous >iec iind wcii;bi of ibc Im^i 
equally certain that it rould not have imii.itecl \\% mixlrm allien I 
Sloths, in ihc feat of climblnj,'. back dowTiwards, amonj;*! the wwfc 
is clear, therefore, th.11 it sought its sustenance upon ihc Krnsmlil 
wax originally supposed lo have lived upon tnnis : but by > ma 
piece of deductive reasoning. Sir R. Owtn showed (hat tin- 
Ground-iloth liv<?d upon ihc foliare of trees, like lhec^i»tiaK^■ 
bui with tliii dlTTcn-DLr. that instead of cliinbtni; am()n;;vi the br. 
it actually uprooted the tice bodily. In this lour iU for, r, \ht inir. 
upon its huge haunches and mighty tail, as on a tripod, and ttoi p 
in^ ihr trunk with its powerful arms, either vrrenchcd tiupfayllMi 
or broke 11 short ofTnbovc the ground. NfancUous as >hi» tnaytfa^J 
can be shown that o-er>- detail in the skeleton ot the Mc-jruihereao 
with the supposition that ii obtained its food in this way. 

A smaller but allied form from the Hcistocenc of Sot;-'- '■ 
has been named Onunnt/ius, but since tht<> term is px:- 
a genus of Palfcozoic Fi-shcs, it should Ik changed. 
Scf/iJoiAerium (fig. T167), which may lie taken to m 
imyx, and likewise occurs in the South American Flci-: 

prises a number of tpecies, and hxis chamciers in su: 

intermediate between the preceding and the following gencnk 
tccih in the upper jaw have an irrejjulaily oval section, while 
of the mandible arc usually subtriongubr ; tbe whole of the scnei I 
are in contact, and their crowns do not wear into ridges. Tlic ' 

prises another group of large Ground -Sloths, which has b«n 
up l>y sonic writers into ihc genera Ltsto^n, Pseud^iestpdom, G\ 
tJt^rium, Sec, according to certain not vcrf iinporunt diifa' 



!C and mode of arningenicm of ihc leclli. In the upper 
'h are usually i.uh triangular or oval in trnriKverse i^ection, 
-al species there is unly n short interv-al between the first 

rond tooth in each jaw, and the former is worn horizontally ; 
otbcr species thert: is a considcrahlc: interval bclwccn the 
and liie first i& worn ohlt<|uely, as in certain Sloths. The 
has no cntcpicondyloi foiainen ; and in the shortness of 
skull and the characters of the teeth this genus approaches 

to the modem Sloths than any other member of the family. 
beKt known specie!^ is the South AireTicin Mylodon roburlui 
■ 169^, which wa» smaller than Afrgat/uriufn amenMnum, its 

Kh being altoui 1 1 feet ; hui Af. iirmaius (the type of the so 
^ £^steJon\ from the same country, is considerably Ur^cr. The 
species it M. Harlani, from the Pleistocene uf " iiig-bone 
in Kentucky, North America ; while the Patagonian M. 
is a very aberrant form, regarded tiy some writers as 
ly distinct, and named Giypolhfrium. There arc numcr- 
dermal scutes, which do not aiticuUic with one another. 
fyx^ from the I'lcistocene of North .\mencn, is an allied 
acicriscd by the long interval between the large first and 
Her second tooth, and also by the presence of an entefn- 
forainen to the humerus. The type species is M. Jfffc' 
frooi Kentucky and Tennessee ; while M. aihtnsii, from the 
of Cuba, has I»ecn separated by some wrilcns under 
nunc of 3fegaloehnus {Af/omi'r/i/ius). Another furin hitherto 
m as Cttiodffn, but which may be named .Ve/ArotAen'uw, on 
Ntnl of the preoccupation of the foitner term, occurs in the Pleis- 
cave-deposits of Brazil, and agrees with Mti;aionyx in the 
ctnrc of its limbs, but ha-s teeth of the type of those of Aftga- 

MM*, alihoi^h their number is reduced to - ; the type species 

Inderably exceeded in size the largest Ant-eater of the present 

the Mammals from the Lower Pliocene and Miocene of North 
tBJca which have received the names Afarapus and MftrolhtriuHi^ 

1 arc rc^i^rded by Professor Marsh as forming the type of a 
linct family of Edcnlatc>i — thi- Moroftodidit — probably belong to 

Ungulate family ChalUotheriidir. 

pAWI LV BxADYPODiDjr. — This family is entirely confined to South 
lenca. aiid now comprises two genera of comparatively small 
nala which are of exclusively arboreal hnhiU- The body is 

lied with coarse hair j the teeth ore - in number in each jaw, 

are of sut^cylindrical form, with a central axis of soft dentine, 


surrounded by a coat of a harder kind of the same substance, 
fore-limbs are enoimously elongated ; and both the manns u 
are furnished with long, curved claws ; the number of digiti 
exceeding three in each foot. The skull (fig. 1 1 70) is short, 
descending maxillary process to the zygoma ; and the tail B 
mentary. In Bradypus (fig. 1 1 70) the first tooth is equal in 
the second, and is worn horizontally ; while the digits are n 
to two. In Cholcepus, however, the first tooth is consider^ 
than the second, from which it is separated by a much 
interval than in Bradypus, and wears obliquely ; while the 
three digits to each foot The only known fossil form is fri 

Fig. 117a — Skull of Anu/r/"' f^"i^- Rcomt. Sooth Anwrka. Rcdwa^ 

Pleistocene of Argentina, and has been named Nothrapus / 
it appears to have been about twice the bulk oi Bratfypus did, 
and has the first lower tooth separated by a very long intern 
the second, although it is of smaller size; the cranium, 
dentition, and feet are unknown. 

Order IV. Cetacea. — The Cetacea form, perhaps, th( 
readily defined and sharply differentiated order in the whoU 
Their contour is fish-like, the body being fusiform, and passi 
perceptibly into the head without any distinct external nee 
posteriorly gradually tapering to the extremity of the tail, wl 
furnished with a pair of horizontally-expanded "flukes," fan 
dense fibrous tissue covered with skin (fig. 11 78), The h 
frequently very large, and may be as much as one-third th 
length of the animal. The pectoral limbs are reduced to 
paddle-like, organs ; and there are no external traces of pelvic 
The skin is smooth and without hair ; although bristles 
present in the neighbourhood of the mouth, more especi 
young individuals. Frequently there is a median dorsal i 
1 1 78), which however has no bony supports. Both the eye 3 
external auditory aperture are small ; the former having no nic 
membrane, and the latter no pinna. The nostrils open by a 


1 301 


tlAc aperture usually near the vertex or the !>kutl. The bones 
^iBEuall]' of a ipongy nature, and contain a large amount or oil. 
cervKal region of the vertebral column in always vc-ry short; 
itbe Kven component elements may be p.'iitially ur completely 
together, while the odontoid process o( the axis when present 
rt and Ijlunt. and may lie enlircly want- 
None of the vertebne unite to form a 
The lumbar and caudal vertcbia: 
and numvrous, and from the ah- 
of <y|;apaph)-sial antculalion<i .illow of 
amount of motion in the hinder part e 
body ; the presence of chevron- bones 
it»hc« the eaudaU from the lumbarN. "^ 
terminal epiph)-»es of the ^-crtebne do 
I imtte with the centra till the animal is 
adult. The cranium presents peculiar 
■la which it will not be necessary to 
aibe here ; although it may be mentioned 
I the uKiull}' Mnall nasaU are £ener.illy 
Lght up near to the vertex, and that there 
I more or less elongated rostrum in ad- 
Bc of the eAtemal tiares, formed by the 
tasxiUa, ma»lla, mesethmoid, and vomer. 
vidcs are absent 1 the scapula and hu- 
are -well developed and freely niov- 
npoa one arvother, but the anterior r\ Witt 

of the limb admit of only a very 
amount of movement. There arc 
■Dy five d^ts in the manuii (fxg. 1171), 
I iheiie nuf be reduced to four ; the 
jbngcalt are unique amoi^ Mammal-i in 
ceding the number of three to a single 
it, and a\m in being furni.ihcd with epi- 
his. The pelvis us represented hy a pair ih, '"ftwa^Snib'^T'lkf 
Utifbrm bones, which are regarded as tl^f-^^.^i'jf^.'^.Z] 
t icchia : and there are occasionally small «■ L'lniw.CuTiii-cb. iwwr 

., - , puccd on tht .oiph^Kl 9t 

Dus at cartilagea representing the rutuie):>-i-w;.MruciirnHu: 

bones of the proximal part of the till^i,.,'''"''"*"'- **^'" 

btnb. Teelh arc u.sually present, hut 

irtty variable in number and size. The dentition in existing 

is bofnueodont and monophyodoni, but it was heterodoni 

extinct y£Uf^ii>donfida of the Kocene. 

Cctacca are not known vfith certainty before the Eocene. 

e most aliundani in the later Tertiary periods. "Wxq denti- 

tbe Eocctte Ziushdoniiitf iiidieates tliat the order has been 



probably derived from Mammals with a heterodont dentiti 
Professor Flower comes to the conclusion that it is most 
that their ancestors were allied to the Ungulata; while 
absence of Cetacean remains in the Cretaceous the same 
thinks that the earlier members of the order were ir 
of freshwater. With the exception of the PlatanisHda : 
Delphinida, all existing Cetaceans are of marine or 

In their increased number of phalangeals (hyperphalar 
Cetacea resemble the Reptilian hhthyosaurida and Pieswtia 
from this circumstance, coupled with their simple type of tc 
been argued that they represent the most archaic type of Hai 
even that they are directly descended from the Ichthyo&auis. 
mentioned there are, however, many practically decisive ob 
these views. 

For the determination of fossil Cetaceans the solid tyw. 
petrosal bones of the internal ear, and less frequently t 
rostrum, are of especial importance, since these parts are 
well preserved. 

Suborder i. Mystacoceti. — In this suborder, commo 
as the Whalebone ^^l^ales, functional teeth are no'er p 

though germs may be 
in the gums; " baleen' 
bone is always attach 
palate ; the t>-mpanic 
117a) is anchylosed t 
otic and involuted u] 
the nasal passages art 
by the nasals, the lai 
small and distinct 
jugal ; and the rami o 
dible are laterally ci 
do not meet in a sj-m 
Family BAijENti 
only family of this 
may be divided into 
nine and Balitnopt 
tions. In the first of 1 
is often no dorsal fin, the tympanic (fig. 1172) has a ch. 
flattened and angulated shape, and some or all of the ci 
icbrre are at least usually fused together. Remains of 
Bahena (including Baleenolus and Balartula) occur abu 
the Pliocene, and especially in the Enghsh and Belgian Ci 
of these fossil species {B. affinis) is closely allied to the 
Whale (fig. 1 1 73); while B. primigenia is more nearly 

Fig' 1173. — Inner vievr of the right tympanic 
"f the (frcenland Whale (fdA^fd tttystatttttt). 
One-ihird natural lize. {After Giay.) 



icit of (he Southern Seas. Pala&alui, which has been 
' Hi deiichbcT a^ of Mvmjwic ajjC, may be proviaioii^Iy 
. to ihU section ; ihe tj-pe was probably obwtned from ihe 
In ibe Saianapurine seclion a dorsal fin is very gcne- 
L (whence the nante " Kinners " apjilicd to many of these 

^->l>aft IU«ml*itwariWilnllitf iht >■■ 

Ii (Aa/MM mjttmtmt). 

the cervical venebnc are free, and thicker than in the 
section ; and the tympanic {fig. 1 1 74) is longer, more in- 
id more rounded (tian in the latter. The existing genus 
TO. (in which may be included the fossil liurtinopsis) is 
Uy represented in the Pliocene Crags of both England and 
-, Bakeno^era also occurs commonly m Uk- same deixxsits ; 
pto being apporcnily nearly alUcd to the existing B. siUaid', 

II}*— IwMr*Ww«f ihtrix'" 

i-tul( luiiml iliK. (AAcT Uriy.) 

RKsal. On«< 

tnins a length of 80 feet ; while H. etfiur^iMitia conies nearer 
Hng B. roitrata, which is seldom more than 30 feet tn length, 
"imm (including Cttolkeriopkants, PUfhxetm. and Pftsioctt- 
characti-riscd by ihe narrowing of the nnterior extremity of 
panic, and is likewise found in the Crags and other Pliocene 

U. 3D 



beds ; while Herpetocetus from the same deposits is an allied | 
with an egg-shaped tympanic, and a talon to the mandibular cot 
which re^ls that of Physeter. The names Ampkicetus, lA 
Isocetus, Hettro^tus, and Mesocetus, have been applied to Cea 
from the Belgian Crag, most or all which may apparently 1 
eluded in Cetotherium. Lastly, a vertebra from the Upper E 
of Hampshire has been referred to Balanoptera, but this reft 
requires confirmation. 

Suborder a. Arch^oceti. — This suborder is confined 
Eocene and Lower Miocene, and n^ay be characterised by th 
nasals, and the presence of teeth differentiated into groups i 

Family ZeuglodontidjE, — In the one genus of the only 

family the dental formula is /. \ C. -, Pm, + A/: 5. Tl 

3 I 5 

ting-teeth are simple and pointed ; but the cheek-teeth (fig. 

Pig. II7S. — Ztugbdm eilfidii : frtim [he Middle Eocene 
America- A. Left laier^ Aspect of cnniiun. mudi reduced; %, 
molar loolh less reduced. (After GandrY.) 

have two distinct roots, and compressed, \ 
crowns with denticulated cutting-edges. T 
nium is elongated and depressed ; the braii 
is small ; the temporal fossae and the sagilt 
are large ; the cranial rostrum is long, and 
sides largely composed of the premaxillae ; the nasals are loi 
and narrow ; and the external nares are placed more anterior 
in living Cetaceans. All the cervical vertebrse are free, whU 
of the lumbar region are unusually elongated ; but the nature 
limbs is not known. In their dentition, as well as in the du 
of the skull — especially the long nasals and the forward posi 
the nares — the Zeuglodonts depart less markedly from the j 
plan of Mammalian structure than any existing members 
order ; and it is remarkable that the Mystacoceti show a ne: 
semblance in cranial structure to these fossils than is made 
Odontoceti, The one genus Zeuglodon is known from the 
Tertiaries of Egypt, England, and North America ; the rem 

Z. ettmiis being extrn ordinarily abundant in parts of the 
intnr, where they have been weathered out of a deposit of 
ne age. 

ki»rK 3. OiX)?fTOCETr- — All living Cetaceans not included in 
fbcoceti Iidong to this suborder, which is characterised by 
knee ofcatcilicd teeth aflcr birth ; the runcltonal ones being 
f ntiinerot]<;. but Kometimes reduced to a sinf^le pair (ooca- 
kanting). Baleen, or whalebone, is inv-ariably abRcnt; the 
(5 more or less unsyromecricnl ; the lu^aK are reduced to 
tty nodules which do not roof over the narial passages; the 
ll is either united to the jugal. or of very large size; and 
fibular rarni are nearly straight, and meet in a median sym- 
The tympanic is not anchylosed to the iicriotic, and has 
(otnptetely involuted stracturc found in the Myslacoceti. 
y PHYSF-TKRitLK. — In this family there arc no functional 
the upper >aw ; and the anterior facet of the periolic for 
pn with the tympanic is smooth (fig. 1 176). while the pos- 
pponic surface of the former bone is broad, with a distinct 
Mge. In recent genera some or all of the cervical vertebrae 
I together. This family is dividwl into the two subfamilies 
Uf and ^jpkiiitit. In the fonner, which comprises the 
tachalot, or Sperm V^'hate (Pfiytfter), and the Short-nosed 
f{Cr>ftii), the niandibular teeth are numerous and implanted 
[ groo\*e partly diwded by imperfect sepia. Remains of the 
Speim >Vhale {P. macrocepkalm) arc found in the English 
tdf artd also in the Pleistocene of South .\merica ; the large 
Ire tw enamel at their summit. Allied to this genus arc 
rtrni the English and Relgian Crags, and Phytttodtm from 
etie of .^ustialia ; while Pkxieteruia is a genus founded on 
rom the former deposits, which does not exceed some so 
;ngth. A number of Pliocene and Miuccnc forms appa- 
Ilin! to the Cachalot, 
1 the crowns of the 
ped with enamel, have 
scribed as Ba/<rncJ'f», 
IS, Hofhattu, Phyfodpn, 
\mdft ; ffifhtthif and 
I comprise comparatively 
ecics from the English 

f Crags and the French 
'hile /Jphioides is from 
Xtioccnc of B.iltringcn 
mherg. In the Ziphiine subfamily, comprising the existing 
sed {ffyjxrwdcn) and Beaked -Whales {FJphiut and ,\feso' 
the mar»dibular teeth, vilh the exception of one or 

ltmetr«itrit : fntm ih« SuHolk CinX' 


occasionally two pairs, are rudimentary. Ifyperoodon is repRifl 
in the Crags of England and Belgium. These beds also yidd 
extinct genus Chonesiphius, as well as several species bdongit 
the living genus Mesoplodon, which is distinguished from C 
ziphius by the complete ossificatioR In the adult of all the da 
comprising the cranial rostrum. The Crag species of Mts^ 
are mainly knowp by these solid rostra, which in the living s 
are compKJsed of dense ivory-like bone, and are the most solid 
found in the whole of the Vertebrata. The periotics (fig. \ i] 
more rarely found, but are equally characteristic. 

Family PlatanistidjE, — This family, which is now rei»e 
by Platanista of the Ganges, and Inia and Pontqporia of lb 
rivers of South America, is characterised by the large nun 
simple teeth in the elongated jaws, and by the length of th 
dibular symphysis exceeding half that of the entire mandible. 
of the cervical vertebrae are anchylosed together. Remain 
to Pontoporia were found by Bravard in the Pleistocene of 
America, but these are referred by some authorities to a 1 
genus under the name of Pontistis or Palaoponioporia. The 
genera Champiodelphis and Schisodelphis, of the European PI 
are usually referred to this family, but the latter approximates 

Family Squalodontid*. — The extinct Squalodonts W) 
merly classed with the ZeuglodonHda, but the characters 
cranium are essentially Delphinoid. although the teeth are dil 

^>K' <i77- — 'Vb'ns\t>tiKrmMi%i:XSqiatadtm; from tb« Miocow of Europe 

ated into groups as in the former. The type genus Squalodon 
from the Middle Miocene to the Pleistocene of Europe, and 
found in the Tertiaries of North America, New ZeaUnd, an 

tralia. The teeth may be arranged as /, - , C. - , Pm. *, 

the premolars are simple, but the true molars (fig. 1177 
double roots, and crowns very like those of Zm^adon^ al 



Sed by the dcnliculations being more developed on the 
' than on the anterior border. The .so-called Rkizoprion \% 
ra species of this genus. 

lUr DELrHisiDA. — In this, the la&t family of the Cetacea, 

arc simple, nnd usually numerous in both jaws j and the 

of the mandibutnr symphyu's may be very siiull, and ni.'vor 

I one-third of tbat of the entire ramus. 'Wi\i pcriotics (fig. 

which are (mjuentty found in a fosiiil condition, are 

' distingutshed from those of the PhystteriJit, by having the 

&cet which aniculatcs Kith the tym]>anic marked by 3 

- of grooves, as wdl as by the turrowncss of the posterior 

their tympanic aspect. 'I"his family comprises all the Ceia- 

Commonly known as Porpoises, Grampuses, Kilters, and Dol- 

RemAins of the Narwhal i^Motiodim nn>in>ceros) are foimd 

Norfolk Foresi-bed and the Plcistocent; of .Maska. The 

Iklphinapltrut^ now represented by the "\Vhite While," 

in ibc Lower Pliocene of Tuscany, as well as in the Plioct-nc 

iocene of other pans of Europe. An extinct species of 

■Whale" (Onw) is found in both the Italian and English 

The existing Globicephalus 
Of '* Black-fish," has left its re- 
in the superficial deposits of 
I,, while the extinct G. Hnadem 
in the Suffolk Crags. A left 
of the latter species is repre- 
id in fig. 1179, in order 10 show 
iicstturcs characttrristic of this family. 
|rn»ttng Pstudorta rrttsridrns was 
illy described from a subfossil 
found in the fens of Lincotn- 
Of the smaller Dolphins it is probable that the genera 
ps, iMgenorfiVfuhus, and Dt/phinus (as now restricted) are all 
!)ted in ihc Pliocene of Kurope. Eurhintxitlphis. is a long- 
enus from the Pliocene of Belgium and Italy, which is 

GMtietfkaiut watfi-tmt : frvm the 
Corallini Cis>a of Suffclll. 



probably nearly related to the existing Siene. Delphinoid icsB 
from the Miocene of North America have been described bj 
Tessors Leidy and Cope under the names of PrisctftUiphiinu, H 
spkys, Zarkachis, LopJioeehts, £habdosteus, Ixaeantkus, An/fi/m 

E1(. iiBo.— Skdeton of ihe Muiuee iMmmmlia mmtinfu). Gnadjr nddcid. 

and Orycterocehis ; but further information is required as to t 
affinity of these forms and their right to generic distinctioiL 

Order V. Sirenia. — The Sirenia, now represented only by 1 
Manatees and Dugongs, agree with the Cetacea in their a(UpUti 
for a purely aquatic Ufe, and accordingly present a strong gene 
resemblance in their external contour to the members of that onl 
The head is, however, of normal relative size ; the tail has a b 

Fig. iiSi.— Donal view or the righi peciiical limb i^ ifiuutim aialrmlh. UnctinitaEi^ 
I, StapuU; k, Humcnu; r, Radiui; v, Uln>i /, Polka. 

zontal membranous expansion ; the pectoral limbs, although modi£ 
into paddles (fig. iiSi), retain the normal number of [^alangea 
all external traces of the pelvic limbs are wanting ; and the surii 
of the body is either naked or covered with sparse bristles. T 
lips are fleshy ; the nostrils placed near the extremity of the mitf 



io) -, ihc eyes minulc ; and the small car has no external 
The bones of the skeleton are extremely dense and .solid; 
vcncbfae mav be reduced to »ix, but are hcvlt anchy* 
; and the axis has a well-developed odontoid process, 
hjive no cpipliVM^; and none of them unite to form^e ch«Ton-lK)ncs are present. Clavicles are want- 
no recent furnis ^ow any trace of the femur, although a 
nidintent of this bone is found in the extinct Nalitfurium. 
ro bones of the forcaim in the existing genera (fig. 1181) are 
al in siec, and fmiuently ancliylosed together. In recent 
|be sltuJI (fi^ ti8i) is rcmarkal>le for the upward direciion 

inanal aperture, and for the absence or nidimental condition 
fOBSals. The teeth can only lie differentiated in the living 
I into tncisors and chcclc-Ietth, which are separated from one 
:r by a long interval ; but in one fo&sil genus there is a more 
dental series ; while in Rhytina these organs were totally 
. In the existing genera the dentition is monophyodont, 
c-molars occur in Halitktrium. This order is known from 
\t upwards, but appears to have been stca^dily dying out 
resent period. The fossil forms, although decidedly more 
llised than their living represenuiives, do not at present afford 
^ indication as to the origin of the order. 
CILV Halicorid^ — In this family the one existing genu^ 
wr (Dugongs) has the deflected preinaxillae (fig. 1 i8j, a) fiir- 
I with a pair of large tusk-like incisors; and there are also 
airs of functionlcss germs of cutting-teeth in the similarly dc- 
1 mandibular symphysis. Five or six cheek-teeth are dfvel- 

Ei each jaw, which aic usually cylindrical, and all grow from 
nt pulps, and are not coated with enamel. The Dugongs 
il the ousts of the Red Sea and Ihc Indian Ocean. Pruhaii- 
hORi the Pliocene of France, k regarded as nearly allied to ihe 
% genus. The only otlier form which has been refcned to 
amily is known by a molar from the Tertiary of California, 
t has been described by I'rofcs»oi Mar»Ji under the nami: of 


i>es»toiylHs. Funhcr information is, howerer, required 
reference can be definitely accepted. 

Family Hai.ithkriid^ — The fossil forms consiituting theJ 
tkeriidtr are charaaerbed by the presence of upper incison; 
cheek-teeth being coated with enamel, like Ihose of the .Win 
and by the retention, in at least some cases, of a milk 

Fig. til}-— CrindinBiarfuvof irK d(nuliiiiul«andlutncbi IwmrnaUn e( tfatltcMil 
/mil* ; mm it>* Huropiaa Mbocm. 

In the type gt-nus /fa/itherivm (which may \k taken to indf 
Haliamsia, Pugmeodon, and Ftlsimfthcrium) there is a pair o( It 
like upper incisors, and either five or »ix cheek-teeth in each f 
small na&aU arc present in at least some of the specks ; the | 
maxillx and mandibular symphysis are bent dotmwards; there i 
descending plate at the angle of the mandible ; and a small toiJ 
ossification represents the femur. The molars (6g. ii8j)haK 
pattern on their giinding suifaoe resembling that of Hi^tpt^ 



■St.— Skdctcpn or flalUMtrimm Stih—ii tnrn ihc Lwnr UiuoM of Ha 

This genus ranges from the I^wer Miocene (Middle Ot%ao( 
to the l^wcr Pliocene of Europe ; and remains oi ff. SA 
(fig. 1 1 &4) are especially abundant in the loiwer Miocote fl 
of ccrlnin distrias of Hessen - Dartnsudt. Promstemin^ 6 
Tertiary siraia in the West Indies, is a generalijied form appiici 

' In ihii ligure ihe deflectioa of the pmaxillw n oaiiHd, 


131 r 

Hafiiherinm, but with a fuller dcntilion, and without the 
ileBection of the extremities of the jaws. The dentition 

\ -, Pm.-^M. —^ ; the uppci incisors are not lusk-llke: 
I 7 or 8' '*^ ' 

cheek'ieeth have simple transverec ridg«s somewhat like 

the UngoUte (genera Dinotkeriuin and Lhtrifidon. Kg- 
is de6nitclj known by a o)»t of the brain-cavity from the 
if Egypt, but tftth from the same deposits described under 

of Manahis may perhaps belong to il ; its afFiniti« cannot 
fopcrly ilciemiined. Probably allied to Hn/itftenum is 
riuut from the Flioccnc of Itclgium ; while Diophtfarittm 

Miocene of South (Carolina is regarded as intermediate 
Halkert and Halilktrium. 

t RhiiTINID.*. — The Rkytinidce are known only by the 
ytina ffgas, or Steller's SeaCow, which was fornierly very 

on Behring and Copper I*.l;inds in the North Pacific, 
nmonly supposed to have been exterminated about 1768, 

a few individuals may have lingered on to a confticlerably 
■- The Rh)tina attained a length of from ao to 35 feet, 
laracteriscd by the entire absence of teeth, their function 
»p(ied by homy plates (cornulcs) on the palate. 'I'hc head 
nail in proportion to the length of the body ; and has the 
R and mandibular s^-mphysis moderately deflected. The 

naked, and covered with a rugged cpidemiis resembling 

of a tree. Nearly entire skeletons have been obtained 

peat of Behring Island. 

V Man.\T)d*,. — In AfaHtitus, the only known representative 
bily, the prcmaxilla: and mandibular symphysis (flg. 1 1 82, n) 
fcflected, and there are no functional incisors. The cervical 
»rc reduced to six ; the cheek-lccth, which are coated with 
jid carry two transverse ri<Iges, may be as many as eleven 
aw, although it is seldom that more than six arc present 
ne time. The Manatees ate inhabitanu of the mouths 
tries of the great rivers discharging into the two sides of 
Kic No fossil forms are known which can be referred 
ainty to this family, aJthough the type generic name has 
lied to certain teeth from the Eocene of Egypt already 


mians of uncertain affinity may be mentioned Chrprrozcum 
later Tertiary of New South Wales ; JlemUaufodon from 
M of Shark River ; Paehynfanihui from the Miocene of 
and Troihytfurium (with which Khytiodus may perhaps be 
&om that of FraiKc 



CLASS MAMMALIA— contituied. 

Order Ungulata. 

Order VI. Ungulata. — The Ungulata, or Hoofed Mammal^ ( 
stitute the largest and one of the most important orders into «t 
the class is divided \ all the included groups being so coniM 
together as to preclude their division into well-defined «]M 
orders. This order comprises at least seven suboiden, of ^ 
three are totally extinct, while all the others, with the exceptiia 
the Hyracoidea, have lost a large number of family types at I 
present day. The two first suborders — Artiodactyla and Poi" 
dactyla — present several features in common, and are acconJini 
brigaded together by some authorities under the names of Ungul 
Vera, Clinodactyla, or Diplarthra ; while all the others are iodu 
in a second division under the name of Subungulata. 

All the members of thb order are adapted for a terrestnal 
and in the main for a vegetable diet, although a few are mo' 
less omnivorous. Their dentition is heterodont and dij^y 
and the milk-set is well developed, and not changed till la 
life ; and in the Fenssodactyla alone among Mammals do W 
certain instances where the whole four premolars are precede 
milk-teeth. The cheek-teeth of the more typical forms have t 
crowns, with either tuberculated or ridged surfaces ; and thar cr 
are very frequently interpenetrated by deep folds of enamel, » 
produce a complicated pattern on their worn sur&ces. Exce 
lypoiherium, clavicles are always wanting. The toes, with th 
ception of Chalico/herium, are provided either with blunt, \ 
nails, or with hoofs more or less completely encasing the ten 
phalangeals. The feet of existing types are digitigrade, and 
number of the toes varies from five to one. In all existing I 
the humerus has no entepicondylar foramen. The scaphoid 


of the carpu3 are always distinct ; but the radius and 

great majority of instances the cheek-teeth are tooted, and 
Artiodactyla, I'erissodactyla, and IVuliosctdca, wliich 
most specialised members of the order, a gradual in- 
Ae height of the crowns of tlicsc tuLth may be tracird 
geaenliscd to the specialised genera. Those teeth in 
cromis are low-, and ihcir whole structure is visible from 
,ng surface, being known as ^fKif-A^rfow/ (compare fig. t tgs); 
with high crowns, in which the bases of the enainel-folda 
ible from the grinding surface, are termed Ayfsi^d-ynl (fig. 
The change from a brachydont to a hypsodoni di-niiiion 
panicd by the production of a nearly flat and horizontal 
surface in the cheek-teeth, in place of a more or less 
ridged one j the itiore specialisi-d type ht-ing adapted for a 
grindiag action of the upper againnt the lower teeth, while 
nofc generalised type the action is to a gri;at extent a snap- 
Examples of the former type arc shown hy ihc Horse 
and of the latter by the Pig and Hyiax. Hypsodoiitism 
confined to this order, aa will be noticed in the nequel. 
e remarks on the probable origin of the order are made 
isMkr the head of the Condylarthia. In the specialised 
tkne is rer)- often a tendency to a suppression of the anterior 
acre especially in the lower jaw, 
DKtt I. Aktiodactvla. — This and ihc next sulxtrder 
eertoJn structural modifications of the extremities, by which 
distinguished from the remaining five suborders, and on 

r^uxount, u already mentioned, they are grouped together by 
miiers under the name of Ungulata Vera, or l)iplanhr.-i. 
^tbe feet are n«ver plantigrade, and the number of functional 
'"fccs not exceed four. In the carpus (fig. 1 185) the scaphoid 
Poptwl by and largely articulates with the magnum ; while the 
('Ogethd with tbe unciform) supports the IuiilU-, and lia:* no 
tion with the cuneiform. In the Uirsus the cuboid extends 
*y to articuble with the astragalus, which is deeply grooved 
> 3S6|. All the component bones of both the carpus and 
'trongly interlock, which makes the slnjcturc of these joints 
onijilex than in the other suborders. Finally, the jugal forms 
' pan of ihc zygomatic arch (tig. 1187), and the brain is 
ly brge size and complex structure, 
(oborder, so far as at present known, is dtstingubhcd 
PerisKMJiactyla by the surface of the astragnlus (fig. 
**iag (inglymoid, by the third and fourth digits (lig. 1 185) 
'Qua] in size, and arranged symmetrically on either side of 
"^wn between them ; by the absence of a third Irocbantct 



to the femur ; and by the articulation of the fibtda with dK 
caneum. In all existing fonns the number of the dorsofa 

Fig. iifis-— L«ft in«nmof (ijPigWiii), (i)Wfo™>K*iu, (3) Cb«Toi»in lTrm£almii 
buck {.Cafrralufi, (5) Sheep (Oeii), and {6> Cmnel (Cniwi/iii). In the tmat. tl» tatUi 
ihe upper tow ij the lunar, Ihe left the scaphaid, uid the right Ih* cnneilonn : in ibe K 
Ihe lefi bone is the munum, and the liiihi Ih* UDaTona. Reduced. (After D» 


vertebrae is nineteen ; and the nasals are not expanded post4 
In the dentition the first tooth of the cheek series never 
deciduous predecessor. The upper premolars are vei; gei 



[)(e in siruciure than the true molars ; while the last lower 
nearly always has a third lahc, the same feature being 
tin the last lower tooth of the milk scries. 

Pecon the pclris (fig. it2&iis) is characterised by the 
of the ilia, which are not much expanded ; but in the 
ramus these bones arc shoncr and more expanded, and thus 
me to those of the Perissodaclyla. The symphysis of the 
pubes ia much elongated. 
molars in llie upper jaw 
four or five main columns ; 
specialised Turms such as 

(figs. 1194, 1195)1 ^^^^ 
' in both uf^cr and lowvi 
tarn low subconica! tubercles. M j - ■ 

le dentition is termed dutii> { \._ J|y{J 
at in others, such as Kporco 
F isoi) and the Ruminants 
e). the outer pair of cuiumns 
I tran»ven«l)' flattened, and 
tr pair crescent-sbapcdr and 
^tion is then termed itUno- 
tl tbc lower molars of the 
W of dentition it is the inner 
' that become flattened, white 
$r ones a.'isumc a crescent 

■There is, however, a complete passage from the one type 
Kher ; the mo&t specialised form^ with a hypso-aelcnodont 
i being of comparatively recent origin. The existing niem- 
[this suborder are divided into the sections Suina (I'i^ 
tpopotamu!;), Tylopoda (Camels), Tragulina (Chevrotains), 
pra (tyirical Ruminants) ; but since such divisions will not 
Del for the fossil fonns, it will be convenient merely to 
[k suborder into families. 

Mild, moreover, be observed that it is by no means certain 
Ihe above-mentioned characters will apply to some of the 
iwn earlier members of the suborder, since some of these 
I in the AnaphtHeriida preaeiU certain remote indicattonb of 
fith the Perissodaayla. 

tdvancc from a bunodunt to a hyp^^u-aclcnudont dentition is 
Uiird m this suborder by a tendency in ttiesecutid and third 
pals to coalesce into a cannon-ione, and also by a change in 
\oi the odontoid process of the axis venebra from a pcy-tike 
Id to a spoul-bkc demicylinder. 1'hc earliest form which 
I referred to ibis suborder is the small Pantalestes, from the 
EocCDe of the United States. Thii gcnui U still very impcr- 

•Hnl AninUci)'!* (/&}. R«ilu»d. 


fectly known ; but Professor Cope, who makes it the 
linci fAniily, »iaies thai il has tritubcrcular upper moli 
digits in the pes, and lookx upon it as the ancestn] 
Camels. Or Schlosscr b, howct-er, not satisfied that ihe^ 


F1(. 118;.— LcAaipcctcf tbc Hlp[«pouunii>Ctf.HimtlMta4 MaA i 


really irituberciilar, and KUggcsts that it may indicate a ' 
type liClwccn the Artigdnctyla and the Condylailhra. 

Tawilv Hii'POi^^'rAMiD*. — This family contains the fiogle' 
W'orld genus JJifijx^potamut, now represented by two sjxoa i 





FIi< iiM.— Left waniu af tUft^Mtiunmt 

Tig. lib. — LcA upHr m 
■Mbf of Hifftftfmnt mmflf 
Htu. Two-Ouijh mhmI itab 

bulky animals, whioh spend a lai^ proportion of thttt titne In tl 
tvatLTs of lakes and rivers. In the skull (lig. 1 187) the angle of ll 
insjidiblc has a descending flange ; the facial portion is mudt do 



the orbits are tuljuhr and very prominent. The dental 

■^ 7^. C- '. ■'*«• -y -^ -• The cheek-lccth arc 

in structure ; the tnic molars (fig. 1 1 89) having four 
[which [ircwnt trefoil-shaped dentine surfaces when wom ; 
biars arc simpler. The upper incisors arc short and ver- 
li 1S7), but those of the lower jaw arc pTocumbcni, and 
tfry ixpge (fig. itgi)- The carineii of the upper jaw are 
pwards like those of pig:^ while ihc lower onts nre of 
I siw, and have their extremities obliquely worn to a cutting- 

-Ulff*^mmta itr^tmtii: from Ax. Pliocene of I Ik S;*a!il Killi. a, P»l3i.-il 
1; #, TliM rlipil upper Inic [niil.u: t, Syin[itiy»i*o(maniji>l<. Kcducnl. 

biting against the outer surfaces of those of the upper jaw, 
((fig. 1 188) are short and massive, and furnished with four 
ivhich the terminal phalangeals bear nail-like hoofs. The 
icarly naked and or great thicknesit; while the ears and 
frciy small, and the tail is short. 

.more Rcneralised forms like H. iivaUmis (fijf. iw), of the 
pf ih« Siwalik Hills of India, there are three pairs of indson in 
■| all of trhirh are of subcqual sije. This hcx.-iprolDclont LTOup 
Med in the I'liocene of Burma by H. {r4t^'.iliciti, and in that of 
ff. boHaritntii ; while il« laie«t member i^ //. mtmaiUctn, of 
ceae ot the Narbad* Vgtlley, in India. In the latter deposits 



also occurs H, patainiiicvst in which the second .town i"*"* 
minute, and perhaps di%;tppe»r( in the aduh : while in die cd 
amp/iibtHS of Africa, which is found fossil in Europe fron tb 
Pliocene- of Ihc V;il trAmo lo the laic Pleistocene, there Be 
psirsof incisiiri in eiich jaw, the innennnst piiir in the loudS 
of enonnous dimensions (tii'. iigij. li i« evident from the 
prevailing in H- palirinilkut that it xt. the second pair of iadi 
IS missing in the existing vjieiie.'i. Allied to. but smaller thia ' 
are H. Pmilandi and H. miniifus, whose remainB »re found in 
quantities in the cave* of Italy and the Mediterranean iitandu 

in the small H. lihtrietisii, now living in West Africa, the a 
lower incisors is' reduced to a single pair. The resembUnce d| 
dible of Hippopfilamits to that of inc Aalhraeothtriida sua 
both are di^ni-cd from a common ancestor. | 

Family Suio^e. — ^This family may be taken to inclodej 
existing Dkotylidit and PhacsciueriJa and ihc extinct iJitrk 
since foti&il fortns indicate a close connection between all ihq 
The chcclc-tocth arc typically bunodonl and brachydunl. d 
inie molars carrying four main columns, which may be eitM 
(fig. 1 195) or of extreme complexity (Hipfxikyusy, but whicj 
w«nr into distinct trefotk Their outer and inner colutf 
however, coalesce into transverse ndt(e& (6g. 1193). "4 
(fig. 1 193) has a more or less elevated supraoccipiial n*^ 
which the profile slopes away to the muulc, the naisal* 
qucntly much elongated ; while the mandible has tw 
flange at the angle. The amines arc frequently targe ftodj 
(fig. 1 192), the superior ones being curved upwards, and I 
ones biting against a facet on the outer surface of the uf 
but in many of the earlier forms, at>d in the recent Fc 

int of Ihete teeth « much less marked. The digits arc 
IT in number, Ijui in the existing formic (%. 11^5) only 

ddlc ones touch the ground. The denial formula h vcry 
M typical one. 

jict genus Listriodon ciifTera from all the other members 
lily in that the true molars (fig. 1193) carry a pair of 
Sftnc ridges. The dental formula is 

■fliK. -, Af. - i the canines form large 

bust upper premolar is simpler than 
Dlars; and the anterior premolars arc 
'idc. The skull is es^ciilially that of 
Lemains of Usirittdon occur in ihc 
(ooenc of the Continent {where ihey 
descrilicd under the names of LaphiO' 

Tapirothtrium), and also in the Plio- 
,iks of the Punjab and Sind. The 
tent the same relation to thoie of Sut 

by the molars of DiaolktrtHm to those of mail/ species 

this abeirant type, we may turn to the typical genus Sus, 

he normal dental formula is /. -, C. -, Pm. -. Sf. **, 

3 14 3 
le first premolar is absent in some fossil species, and also 
ican Petamotheerus, which cannot be palxontologically 
The canines ate developed into tustks (fig, 119a), 
bey are small in the earlier species. The crowns of 
true molars are oblong, and both the upper and lower 
aobn (tig. 1 194) have a third lobe, although its degree 

a s 

til- . r^j. - Ihc ..t- 

dihI Icf< iipt" inj« mo- 

\m.i of Luirt^tn t^if^t- 
diHi ; tiom tht Middle 
Mlftcenu of Friin-*.- 



of development varies greatly ; and these teeth do not cooi 
use until the first molar has been well worn. The anteiia 
molars are compressed, and there is no diastema between di 

and the second. The 
premolars are simplei th 
true molars, and there i 
siderable difference in lb 
ber of accessOTy tubeic 
veloped in the latter; 
species in which these ai 

Fig. MJ4.— The third right lower init molir numCTOUS shoWine a IDO 
of Sut cntlatut i India, a, d. Middle ctJumiw , ° , 

of talon. plex pattern on the 

crowns. The molars ( 
of the earlier forms approximate to those of Hyotherium. 

The species with the most complicated molar structure are .S. j 
of the Pliocene of the Siwalik Hills, S.phacocharoidei of the PI 
Algeria, .S*. kamuliefisis of the Pleistocene caves of Madras, 
living Indian S. cristatus (fig. 1194), which is also found foK 
same caves ; the last lower true molar of the first-named specie 
a decided approach to that of Pkacockarus. In the Europe 
boar, which is found fossil as low down as the Norfolk Forest 
hind lobe of the last molar is of moderate complexity. S. fila 
giganUus of the Siwalik Hills of India, together with S. am 
eryiiiiintkius, and 5, major of the Lower Pliocene of Europe, 
species with comparatively simple molars ; the first being ih 
known species, and fully equal in size to a Tapir. S. kysutiric 
Siwalik Hills, ^. palaociusrus of the Lower Pliocene of Eppelsl 
.S", cka'roides, which has been recorded from France and ih' 
Miocene of Tuscany, arc small species with simple molars lik( 
the living 5. andamaneiisis, to which they may be allied. .S. ar 
of the Upper Pliocene of France is closely related to the living .■ 
[Potiimocna'rus) a/ricanus ; while in the smalt .S", punjabUns. 
Pliocene of north-western India we probably have the direct ai 
the Pijjniy-hog [S. sah-anius) of the terai-lands of Nipal. No 
tatives of the genus occur in America ; and S. c/ueroiaes seems 
earliest species. 

Here it will be convenient to notice the African V 
{Phacockcerui) which appear to be related to some of the sp 

fossil species oi Sus. The dental formula is / , , C - 

(2-3) > 

M- -1 but the whole of the teeth with the exception of the 

and the last true molars may be lost in the adult, thus pres 

very remarkable instance of extreme specialisation. The 
molar is a very peculiar tooth, consisting of a great numbe 
agglomerated columns, or denticules ; but a marked app 
this structure is presented in some of the species of Bus I 



01. I. 

m. z. 

OT. 3- 

Fic. Ilos.— The [f/l upper inx moliii* of Iffvlkt- 
hium firimtitu : (mm ()>« flioecne aS India. 

of Algeria and India. Remains of Pkaeofkarnt occur in 
deposits in Africa. Tile last niwlar of this gt-nus com- 
I with that of Jfyotfurtum pretient!; a tliflVrence analogous to 
twccn ihe cOfTc»[>unding tctth of Maslodon and ihc Mam- 
may DOW revert lo Uic consideration of more generalised 
The most importanl is Hyatherium (in which may l)e included 
ttrus and Chtxromorui) which presents chiiratttra tonnect- 
with Dieotylet, Sus, and Chirropolamus, and may have been 

tor of Ihc first 
r these genera. The 
tnie molars {fig. 
have low, .square 
with a rudimeninr)- 
I column, which b fully 
>ped in Cfutr»pola- 
and the last true 
comes into use be- 
the first is worn. In 
ft third upper true mutar 
brc is no third lobe, and 
k lobe is small in the 

Responding lower tooth ; while ocrasionilly the la?l upper prc- 
olar has only a single outer column, as in CharofoUimui. The 
AiiHs axe scarcely Larger than the incisors ; are oval in sectiur, 
>d the lower one i« not received into a notch in the upper jaw. 
i)e Utend melapodials are stouter than in modern pigs. In Europe 
to genus ranges from the Querc}- Thosphorites to the Middle 
!)occne of the Continent; while in India it occurs in Che Lower 
nnliks of Sind, and also in Pcrim Island (fig. 1 195) ; the .Xmcrican 
niks which have been referred to this genus ate regarded by I'ro- 
nor Cope as d).stinci. Hippokyus, of the Indian Sinaliks, ap|>cars 
I be an alhed but specialised form, in which the crowns of the 
olars arc much ullcr, and have lateral infoldings of the enamel, 
bffetiy an extremely complex pattern is |>roduced on their worn 
ifbces. SaniiAerium of the Siwnliks must be placed with this, 
mup; while Dnlitxherrut of the Quercy Phosphorite* is apparently 
lied to Hyttheriitm, although lI may also have alhnity with Ceho- 
tmri: The genus Babiruia, of Celebes, is unknown in a fossil 

In ibe John Day Miocene of the United States there occur pig- 
kc animals apparently connecting Ilyoihcrium n-ith the existing 
tccaries, most of which may l>c included in the genus Chatwhyut. 
Tlese forms agree witli iiyolktrium in having the fourth upper pre- 
firapter than the true molars, but have [he lower canine with 


a triangular section, and received into a notch in the \tppa ji 
in the Peccaries. The typical forms have only three ptmolan^t 
in others, which it has been proposed to separate 
Bothrolabis, there are four of these teeth. Allied, or piob^il 
tical forms have been described as Thinakyus and PerdmrOti 
former having four premolars. The existing South Amerioii | 
DieotyUi includes the well-known Peccaries, and has the 

formula /. -, C. -, Pm. ^ M. ^. The structure of the 

31 3 3 

canines has been already mentioned ; the last upper pranobi I 
four columns like the true molars (fig. 1 1 96) ; while the third 1 

Fig. 1 194. —Grind ing lurfacc orihe riKht upper check-iccth of* Paccuy (Zhiwffifa 
laiiaitu). (ATter Giebel.) 

true molar, which comes into use before the first is worn, hai 1 
distinct third lobe. In the pes the phalangeals of the fifth digit I 
aborted. Remains of two existing and one much laiger extii 
species of this specialised genus are found in the Pleistocerw ( 
posits of the Brazilian caves ; while lai^ Peccaries also occur 
the Pliocene of the United States and the Pleistocene of Men 
which have been described as Plaiygonus, but may be includ 
in the tj-pe genus, although they have rather simpler premolars. 

Family Chceropotamid*. — The true molars of this extii 
family are intermediate in structure between those of the SnUa a 
Anthracoikeriida ; having in the upper jaw very broad and shi 
crowns, which carry five columns arranged as in the latter fami 
The premolars, although somewhat compressed, are not secant, a: 
may be of very large size. In the two best known European ge« 
there is a diastema between the first and second upper premoU 
I'he mandible has no descending flange at the angle. 

One of the most pig-like members of this family is the gen 
C€boch<Erus, comprising animals of the size of Hyothtrium, whi 
have been considered by some as allied to the Lemuridtr, althoi^ 
there is little doubt that their true relationships are with the prese 
group. I'hey apparently possessed the full typical number of ted 
which (especially in the lower jaw) present a great resemblance 
those of Ifyotherium ; and it is highly probable that they are ckwf 
related to the ancestor of that genus. Cebochtems is represented \ 
several species in the Upper Eocene of France ; but the form i 



the Mktdlc Miocene of Bavaria may l>c gcncrically dis- 
«d to this genu)! arc Uemickarus of th(; Quercy Fhos- 
id Ltptcihanis of the North American Miocene, in which 
lars have a v«)' simple siructure. All ihese forms, to- 

. ihc foUowijig gen\J», are placed by some wtilers with the 
he ij-pe genus Charoj>ofamus orcurs in the Upjier Eocene 
and Englind, and had bten erroneously stated al&o to 
e Mioceoe of Bavaria. The upper irue molars resemble 
rtain species tA Anthracetkerivm, but have shorter crowns, 
approach 10 a sclcnodont stniaure. There is a third 
te last lower true molar, and the dental formula n 

J*m, -. M. -. The feet are unknown, but it is probable 

3 3 

wwe furnished wiih four digits. The typi.' species C. 

1 the Paris gypsiim, u-as an animal of the size of a lar^e 
is probable ihat this genus is a survivor of a form which 

ximon ancestor of both the Suidu and the AnthriKoihe- 
vtktriym, which has been also described under the names 
'«, Ar{JurfftAtrium^ OltinotAtrium, and Pelvnax, is a larger 
in many dental characters :o Ckttr&f^tamus ; ar>d is one 
' few members of this suborder in which the last lower 
has no third lobe. The premolars are relatively large 
; the canines recall those of some of the Camivora; and 
•nal digits of the feet arc reduced to two. The tienlal 
the typkal one ; and the genus is placed by some writers 
tthriuotkeriida. Its remains arc found in the Upper 
osphorites of Quercy, in the Lower Miocene of Ronzon, 
and of Hemp»tead in the I:de of Wight, and also in the 
)f North .\mericn. Apparently allied to this genus is 
on of the Pliocene of the Indian Siwalik Hilb. in which 
rer molar haK a third lobe, and the conienl premobn; are 
tia siie. In this nei(;hbourhood must probably also be 
! remarkable North .American Eocene genus Aehatwdon 
dcntical with Pamhyut) which has, however, been con- 
ProfiKsor Cope as allied to the Leinuruiden and Insccti- 
e strtictarc of the teeth is like that obtaining in the pre- 
; and the last lower molai has a third lobe, but the linit 
B wanting in both jaw& 1'hc fJiull presents, however, 
nivorous features, and it is possible that this genus should 
placed among [he bunodont Condylartlua in the ncigh- 
cX Ptrtfityihus. The resemblance presented by the teeth 
^to those >>{ Elvtherium and AthatioJi^n is, indeed, so r<- 
iPio suggest th:it thusc two genera may be descendant*; 
iJtnown member of the Condyiorthra verj- closely allied to 


?'amilv AKTHKACOTHERttD^ — In thii Eamtlf the dentil 

is, with one exception, /. ', C. , Pm. , ^f. ^ \ the 


molars ^fig. 1199) hnve broad, low crowns, ^Sth five oolunm,! 
of which arc situated on the anterior and two on tbc postcriarl 
the columns in hoth upper and lower mobrs ha%-e a morel 
di.stinclly »ek-nodont structure ; uid tbc nutidible hu x 
ing flange ai the angle. 'Itie Anthncolheres were 
appearance eomew-hat tietwccn a Pig arul a Hi( 
doubtless dwell in •iwamps and man^heii. In the 
Ihracothtrium (fig. 119;) the selenodont structure 
le» marked than in the next genus, with vhich it 
four digits lo each foot 

The species which approaches rcaresl lo C^trrof^'lumui ia 
of its molars is ihe tniall A. tilittremt t>\ the lower ^ - "^ 
but the itill smaller A. Crettiyi [tig. 119?) from ih'. -H 

Swi tier land and Hampihiretajfrces vtth thai );cnua iit : „ . i 

dcaiiiieiii (tvniilK U^ipcr Botaot of Hai4mll, Hjm^iliit*, 

I between the lirsi and second upper prcmolani, and in tbc *l^|^| 
first lower premolar. In most of ihc other species all thecheomH 
in contact. This genus commenced in the Middle Eocene o( Pn 
ii) Diitmatix, where it is represented by A. itnlmatinufH, which hv 
ide Ihe type of ihc xenm PrtnaimUkerium ; it wa» abundant 

'Qucrcy PhotplmriieN and Lower Miocene oi Eurujie, where il tr 
prescmed l)y species like A. magnum, A. iiildmu of Lauiannr. » 
i/fyrinitu of Tuscany, which aitamed ihc siie of a Khinocerm. ' 

vKprMentcd in the Middle Miocene of France by A. Cirfitri. aftl 

^il died out in P.umpe, alihou^ih it sunived in India till the I'f, 
cenc, where it \^ Icnoun by the lar^c A. hyoptiiamfidt% and the \ 
tilislrmu. It \i unknown in .\incnca. 



enus Hysptlamus generally has a more completely scleno- 

ptition than AntArawiAerium, but in some sptcies (fig. 1 19S) 

tets axe less marited, and thus indicate a complete 

between the tvro genera. In those species (fig. 1199) 

ive the most perfectly selenodont dentition the columns of 



. MfS.— Th( ihiRi l*fi unitrinw 

Fig. **M **Tlt* ikuJ n^hc 
upprr tnw moki nf HjvftUimui 

i,i-i'aiu, \jl,»tt MifiCtfU. Ill* tt 

i molats are taller than in the others. The first upper pre- 
k separated by an interval boih from the canine and the 
tpnsiitobr. This genus apparently commenced in the Upper 
jif Europe, and is especially duiracteristir of ihe Hempstead 
I the Isle of H'ight, and of the Ronion beds of I-'rance, where 
firesented by the large ff. havinus (fig. 1 1 99). H. vr/aunui, 
t brachydont J/, /ora'nus. In India It survived till the 
llioccne, where it i.s represented by two species, one of 
]6g. 1 198) iii the largest known form ; it 
Curs in the Miocene of North Anicrica. 
hme Uifi/afius has been applied 10 
j from the Upper Eocene of Hiinip- 
(ith only two tjigits to each foot, which 

r referred to this family; since, how- 
dentition is unknown this determinn- 
only prmnsional, and it has been sug- 
iiat it may be a tyUhodon, although it 

too large lor the type SpeClC-S. i«rtiue malatur MtTvt 

,v Mi:RvcomiA«iu*.-This family C^^tTlil^u/™ 
fcgardcd a» an offshoot from the An- 
iida in which the upper true molars (fig. tsoo) have only 
mns on their crowns; the dental formula being the same. 
.We that the feet were tetradactylate ; and the mandible 

Fig. iroo.— A rljlit up- 


has a descending flange. Some writeis include this bn^i 
Anthrcuotkeriiday and perhaps this is really the better ananga 
The type genus is found only in the Pliocene Siwaliks of ! 
and Burma, where it is represented by M. dissimilis (% i 
and two smaller species. An imperfectly known but 6as^. 
form from the Upper Miocene of Sind has been deschbed l 
the name of Htmimeryx. In this and all the preceding bmiliesi 
odontoid process of the axis vertebra is peg-shaped. 

Family CotylopiDje. — This extinct North American fan^ 
usually known as the Oreo^ntida^ is r^arded by Professor Cii| 
as related to the Anoplotkeriida, but with more completely sda 
dont teeth and less specialised feet, which are more like thosei 
the Hippopotamus than those of the Ruminants ; but their afyb 
are probably widely spread. Upper incisors are present ; the In 
molars (fig. isoi) are selenodont, and those of the upper)) 
usually have only four columns on their crowns \ and ^ p 

Fig. 1201. — Orel surface of the ri|hl upper ehcck-lccih of Efarttdrm mmjer. 
Miocene, North America. 

molars are simpler than the true molars, but are not secant 
lower canine is approximated to the incisors, and its functio 
taken by the first premolar, which has a tall crown biting W 
the upper canine. The ulna, radius, tibia, fibula, metapod 
navicular, and cuboid are all distinct ; and each foot carries 
digits. The odontoid process of the axis vertebra is intermet 
in shape between the peg of the Bunodonts and the half-c>'lindi 
the Ruminants ; the lachrj'mal bone frequently has a deep lar 
depression ; but the angle of the mandible has no descending fb 

In the type genus Cotylops {Oreodon) the dentition is / -, ( 

Pm. --, M. ; the orbits are completely surrounded by bone 

4 3 

premaxtlla" are separate, the auditory buUx not inflated ; then 
no vacuities in the bones of the face ; and there is a small polk 

' The name Oreodon being preoccupied by Orodut {.mpra, p. 940I. in» 
the change of ihe family name. Celylepi was originally applied by t*ii1y l» 
species of the lype genus. 



nos. In Eportadofty or Eucrolaphus. ihe above chaiattera 
t«n«, with the exception that the tympanic bulla are it^- 
td ; while Merycothttrus differs from the laittr by the anchyloas 
ihc pfrmftxilla;. There are se\-L'n species in the latter genus. 
"yd^t b distinguished Tiom Mcrycaeharus by the presence 
nnttties in the bchrymal region. In Leptauthemn, which has 
■laced by some writers in the CameJida, such vacuities occur 


-rroiilal(Oiib) Mi lBi*nl(>J Mpt«ti of Ihe cranium oFOviMiWMfur/iKinM; 
fr«ilMMic«m«B(lbtVnltKlSu>la- Reductd. (AfMr C^p*^ 

c to the frontals, and the nns^nls become very!. Still more 
ailubic is the cnormoun development of these vacuities in Catio- 
ns (fig. 1 20i), in which ihe uppei iiicii^ors are wanting. Pirkru'sfes, 
n, differs from all the preceding by the absence of the first pre- 
ai, and has but one p.tir cf lower incisors. In the second dlvi- 
t of this family, which indudes the j;c-nt-r.T Ap-uvfrn-rus and 
the orbit is incompletely surrounded by bone, and the 


fourth upper premolar has two outer columns in place of thi 
one of the typical section (fig. i30t). Colereodan diffei 
Agriocharus by having only three premolars, but it may b 
tioned whether this dtfierence really afTords sufficient grou 
generic distinction. The majority of the genera are conf 
the While River Miocene of North America, but Mtryi 
extends into the Loup-Fork beds, which may be either oi 
Miocene or Lower Pliocene age, A tooth, apparently indist 
able from the molars of AgriocAants, has be«i obtained h 
Pliocene of India. 

Here may be noticed a remarkable form from the Upper 
of the United States, described under the name of JVei 
The organisation is said to be of the Cotylopine type, 
upper molars have five columns, as ia the Anthraeotheri. 
Anophtheriida. This genus probably indicates an ancestral 
the Cotylopida, which should perhaps be referred to a distinc 
it is, however, placed by Professor Cope near the Xiphodon 

Family Anoplotheriid^. — In this family the cheek-t 
imperfectly selenodont ; the crowns of the upper true mo 
1204) carrying Bve columns, three of which are placed 
anterior, and two on the posterior lobe, or hal^ of the aw 
the bones of the limbs and feet remain distinct hoia one > 
and there is no descending flange at the angle of the n 
The functional digits may be either two or three in numt 
the carpus and tarsus of the original genus are of that type 
the name inadaptive has been applied.' The anterior p 
are more or less perfectly secant ; there is generally no dia 
the dental series ; and the canines are short and comixes 
depart very widely from those of the Anthracotken'ida a 
allies, in which they resemble the corresponding teeth 
Carnivora. In the type genus Anophtherium (in which 
included Eurythcrium and Diplobune) the dentition (fig. 

usually / ^, C. ', Pm. -, M. -; but occasionally the fii 

premolar is wanting. The tail (Bg. 1203) is long; the fi 
digits may be either three or two ; * and the third upper pren 
a well-developed inner tubercle. In the typical A. commttti 

' In ihe inadaptive modilicalion {Anoplolhrrium) the carpals of tt 
difijits remain as useless lateral bones ; while in ihe adaptive m 
{Nyoihirrium) they shift iheir position, and take a share in the sup[ 
lar^c persistent digits. 

' I'rof. Cope has suggested that the foTms with two digits should bi 
from tliis family, but it is the type species which presents this feature, 
with three functional digits is indistinguishable by dental chancten 
typical A. commutu with only two. 



lypsutn, the columns of the cheek-teeth arc comparatively 
but in other species, like j4. cayfuxeme of llie Quercy Phos- 
Sfas (t^. 1204), they arc shorter, and the teeth thus appcoxi- 
to those of the brachydont species of Hyopotamui. This 
u con/ined to Euio{»e, and la cluractcriiitic of the Upper 
toe <Lower Oh'goc«ne) and the Miocene of Konion in ?uy-«n- 

, noj,— SW«I«« rfyl*^^***"!-* .*<"Ui«-r.' L'vfci tucu«. turuin. MuchniluivJ 

Ky. Tbc species from the South American Tcrtiarics originally 
tred to this genus is now Itnown as Proterothtrium, and is 
jccd UTxler the I'crissodactj-Ia. I'hc Isrgc&t species wa» ahoul 
Bsc of a Tapir. Here may be noticed five genera from the 
licy Phosphorites, some of which appear, on the whole, to be 
I neaiiy allied 10 Anop/olAtriurn, although iheir teeth present 
ain rcscrabUnce* to those of the Perissodactyla, in vfhich sub- 
the last of the group is placed by some writers. Of these 

tioiL— Til* I»i Dtc ri(Sr uytitt (litek-ianh of Aaif/ftitrittmt^mjtiMt; 
fruni tlir Uppcf EotJtpr of Fr*At« 

tra Adeathtriui» is characterised by ihc extreme coniplesiiy of 
tasi upper picmoUr, which resembles the first true molar : 
ri^tkehum is only known by the mandible, in which [he pre 
m are simpler than in the type genu»( : and the true mulars 
» sonae resembbnce to those of Lcpkiomeryx ; Mtxtclhtrium, 
;h is devribed from the pabie, and b probably identical vith 



the preceding, also has simple premolars ; Myxoekarus she 
completely selenodont teeth ; while those of Tapintba appc 
to the molars of the Tapinda. In another direction i 
Dacrytkerium and PUsidacrythtrium, from the Upper Ee 
France and England, in the former of which the first upper 
of either side are separated from one another by a wide 
while the first three premolars are more completely sccanl 
the type genus. The dental type of Dacryilurtum leads « 
to that of Xiphodon, which, in accordance with the ^iews 
fessor Riitimeyer, is therefore placed in the same £unily, \ 
some writers make it the type of another family, whidi is 
include either CanotluriMm (Fk 
DUhadon (Schlosser). The tnji 
are like those of the type genui 
therium, although more completd 
dont ; but the 6rst three premc 
«, , , ,. i2oc) are much elonsated ■ 

Fig. ijoj. — The lul two left -■' j i_ r - i j- 

upper prcmoim and fint irue presscQ ; and the lunctional dl| 
S^'tJpifg^S^^'^r reduced to two in each foot 
typical forms there was no dii 
the dental scries (which comprises the full typical niunbc 
certain smaller forms, separated by some writers under the 
Xiphodtmiotherium, a distinct diastema was developed. Tl 
donts were animals of slender build, with limb-boiKS 
sembling those of Anoplotkerium, and partly those of 
specialised Selenodonts ; they are characteristic of tl 
Eocene of England and the Continent, the largest spe< 
X. magnus, and the smallest X. {Xiphodontolherium) m 
of the Quercy Phosphorites. This genus, although not in 
line, shows how a transition can be effected from the highe 
cotheriida to the Diehadontidir. Jihagatherium, from tl 
Eocene of Switierland, is an allied genus. Finally, Dr . 
refers to this family the genus Brachytherium, from the 1 
South America ; while Dr Schlosser would include in it 
Tetrasehnodon, founded on teeth from the Quercy Ph( 
which have only four columns on the crown. 

Family C^notheriid,e. — Following the classificatioi 
fessor Riitimeyer the next family we have to consider : 
which the type genus is Ccenotherium. All the genera ha' 
complement of teeth, and there are usually five columns, 
on the crowns of the upper true molars (which may be eitl 
dont or bunodont) ; two of these columns being place 
anterior and three on the posterior lobe of the teeth, thus 
the arrangement obtaining in the Anoplotheriida. The I 
Canotherium comprises a number of species of small ai 

ime writers. The limb-bonis show characiere connecting them 
with ibc Suida and the Ruinm.ints. In the ii^urcd cr;tnLUii) 
OMOli are peculiar for tcrminnting in a point. 'I'his genus, of 
ti tbe oames Zoo/igus and MurothenMm arc al^io synunyins, 
■s from tbe Upper luicene (Ixiner Oligncene) of V.itjrliit;e to 
jo^KT Miocene (Upper Oligocene} of .Mlier, An allied genus 
otui/iulAerium. of the Quercy Phosphorite*, in which the tliird 
r true tnolar has but four columns ; while yet anotiier allied 
ftOBX the ume deposits has received the name Oxitcran. The 
tpcdn Upper Eocene genus Dich^hunm {Didymodon) is regarded 
tofcisor RUtimcycr as a bunodont form clusely allied to Ornw- 
'itm i Diialhtrium and Sfnniolfuhum being kindred genera 


from the Quercy Phosphorites. The limb-bones, allhoujj 
four digits are developed, show considerable resemblances w 
the Ruminants, and it has been thought that Dichobunui i 
been the direct ancestor of Gelocus. Here also may be m 
the peculiar Upper Eocene European genus Acothentlum, 
the general form of the skull and teeth of the one know 
seems to indicate affinity with Dkhobunus, while the absen 
third cusp on the hinder lobe of the upper true molars, and 
completely bunodont structure of these teeth apparently 
affinity with Ceboeheerui and its allies, among which son 
prefer to place this genus. 

Family Dichodontid^. — According to the views of 
Riitimeyer this family is taken to include several genera wit) 
dont dentition in which the upper true molars have four 
the type genus Duhodon presenting affinities with Xipha 
Gelocus and its allies are closely related to the TraguUd, 
Cervida. Other writers, however, who do not attach sue 
ance to the structure of the molars, place Dkhodon with 
and make Gelocus the type of a distinct family.^ In Di 

dental formula is / ^, C -, Fm. ^, jW. * : the upper ti 

31 4 3 

having concave outer surfaces somewhat like those of H^ 

while the earlier premolars are elongated and secant, ant 
proach those of Xiphodon ; there is no diastema. The 1 
are unknown, but it is probable that there were only two 

digits. This genus 
the Upper Eocene 
shire In Lophioi 
1207) from the Up| 
and Lower Mioce 
Continent, the hii 
crescent of the u 

Fig. laoj.—Thtlaii four righl upper cbeek-t«lh molarS IS imperfcCli 
of Lefkiemfyx Clutltmiali; from ihc Ouercy . j ^t e . i 

phosphoriii*. eo, and the first k 

lower true molars 
that of Anoplotherium, while the second is Ruminant-li 
lower molars also resemble those of Metriotherium (p. 1 
the family position of this genus, which is referred by I 
to the Tragulidcv, is still doubtful. L. Gaudryi from tl 
Phosphorites has been made the X-j^ o*^ ^be genus Cr 
In Gelocus, from the Quercy Phosphorites and Lower \ 
Puy-en-Velay, it is not known whether upper incisors wen 

' I'rof. Cope would include in this fnmil}' ihe type genus and th 
Agriocharus and C'tlereoiioH. 


e upper motars have low columns, n-ith wide and open 
tbc external surface of each lolic resembling ihat of DirA' 
The navicular and cuboid boni» of the tarsus were united, 
Ibe tiictAiarsaI& fused into a cannon-bone, although the incta- 
ib «ere diuinct Allied to ihis genus are J'hantromeryx 
Frotamtryx of the L'ppCT Eocene of France ; while CA<rr»- 
T of the Siwaliks of India nuy be provistonall}' referred to this 

HtLT Tbaguud^ — The fo&sti forms included in this family 

itc a transition from the t>'pical genus on the one hand to the 

^ntidtt, and on the otlier to the Cervida, 'I'hc upper true 

% have four columns, and the earlier premoUrs arc more or less 

tetely secant. None of the genera were furnished with anilers, 

i* probable that upper mci»ois were likewise always aUtenl. 

e existing forms 

iper canines of the 

(fi%. i3o8)are in 

apeofttulu; there 

long diastema in 

^ws ; the third 

ih, or 'psallcnuin,' 

ing ; and the pla- 

^ diffuse The 

re supplementary 

md the mctacar- 

ihe third and 

digits either unite late in life to form a cannon-bone, or re- 
ks in JiyonioseAus) peniianenily separate. Backilherium of 
BTCy Phosphoritet, in which there are three premolars in each 
lows resemblances both with Gf/iKus. Hyumoie/ius, and I*ru- 
\tritim , the upper teeth being ver>- !>im)lar to those of the 
In Horrathtrium, with which the existing Hyomosehui of 

tppcars gcncrically identical, the premolani arc .—^ .< and arc 

cant tjrpc, and thus indicate ai)inity with the Dkhadantida ; 
nus is found in the Middle Miocene and Lower Pliocene of 
ntincnt, and also in the Pliocene of India. I't-agutus, which 
by the fusion of its nieupodials into cjmnon-hones, and by 
Bving more than three lower premolars, is now coiilined to 
JenULl rc;gion, aiKl is represented by one sipecres in the Flio- 
if India. Ltftamtryx, from (he Miocene of Nurlli America, 
Vt prenrxrfan, of which the first three are simply t;ecant as in 
4us, while the fourth has an inner tubercle \ there are four 
te meUcarpals, but the third and fourth metatarsals form a 
i-t)onc, as in Gtlvtus, III Vntdremvlherium, of the Qucrcy 

Jit, iioI.-Ri<li>cnl*Ucvirwotlhe*kiill«r 


Phosphorites, the dentition is almost, if not quite, indistisgi 
from that of Lepiomtryx, but cannon-bones are found in bot 
although the union of the factors in the anterior ones is less c 
than in the others. These two genera apparently connect G 
completely with the cervine FalaomeryXf that we can have 
tation in regarding them as representing the direct Uiu 
descent of the Cervida from the Dichodontida ; while we n 
sider the existing Tragulida as lateral offshoots from son 
allied primitive stock. Hyfertragulus, from the Miocene 
America, appears to be a form closely allied to Lep/omeryx^ 
the metatarsals separate. 

Family Po&brotheriid£. — We must here leave for a s 
the connection between the Tragttlida and the Cervidte, to 
the Camels and their allies, whose nearest existing relado 
be found in the former family. The Poibrotfuriida are rq 
typically by the genus Poebrolherium, of the North Ameri 

cene, in which the dental formula is / ?, C -, /*«.-, Jtf. 

3 > 4 

structure of the cheek-teeth is selenodont ; in the feet the 

fourth metacarpals remain distinct, the second and third bi 
mentar)-; the carpus has a trapezium ; and the navicular ar 
are not fused together. The structure of the cervical ve 
the same as in the Camelida, of which this family may be 
as the ancestral type ; but in other respects there are 
affinity with the Tragulidte. The type species of Poebtvthe 
scarcely larger than a Fox. In the John Day Miocene o 
an allied form has received the name of Gomphotfuriu> 
Lepiotragulus of the Upper Eocene of the United States 
the ancestral form of both the Miocene genera. 

Family Camelid£. — In the Camels the cheek-teeth ai 
dont and quadricolumnar, but of somewhat simpler struct 
those of the following families. The navicular and cuboit 
distinct, but the metapodials unite to form a cannon-bi 
least one pair of upper incisors is present; and in the cervii 
brx the arterial canal passes obliquely through the anteric 
the pedicle of the neural arch, and is thus confluent posteri 
the neural canal ; a similar condition prevailing in Meuf 
among the Perissodactyla. At the present day this family 
sented by Canulus of the Old, and Auchenia of the New 
but it appears to have originated in the latter, where a large 
of forms have been found. The most generalised mf 
Protolabis, of the Miocene of the United States, in which tl 
tion is numerically the same as in Poebrotherium, on which 
Professor Cope makes it the type of a distinct family. 
eamtius (fig. isto, fi, c) of the Lower Pliocene or Upper 



incison were reduced (as in all the other forms) 

ybt. Ibcre were still four premolars, allhough the first is isolated 

|.the Kcoad very small. Pliauchcma (which is perhaps identical 
lemxamttus) has onlj^ tliree lower premolar^ and occurs in the 

^—\jA tMoal view oT th< iliull oT iIhc C^nwl (OuhAh futttrtiaaifi, ]t«<lDM<I. 
uf^r ■■■rite* : (, r, OuiBOi: jhw. luiUiei] [ircmolu'. Thamaiitlo-pniiiiuilkry Miruie duulJ 


bup-ForV-hc<]s of North America. In Camtlut a^^ain (%. 1209) 

K normal adult formula of the cheelc-ieeih k Pm. ?, ,V. ^ ; the 

3 3 
nt upper premoLiu- being cai)ine-like, and separated by a long 
Itervat from the t^wiiultlmaie tooth of tlui series. This genus ix 
Ijckfincd to the Old World ; the earliest known species occurring in 
i« Pliocene of India, and another form (C TAomasi) in the Pletsto- 
tnc of Algeria. The molars of the Siwalik specie*! show characters 
ow only found in Auckenia. The latter i^cncric tenn Is here iiikeii 

»'itU toft lown Itoc UKiUl of litu^itut ktlUrma, l'lei»liKC"r, CiHfotniB; •, 
r pi^ntoUr aiid l>rtl Inke niolj 
ncbi \trwtx tiua moUi uf du> 

1 i^\mm< praoiolir auA B«l line nio!»t vf Prtttrntlmt virrffuu. Miocent, nonb 

_. lude a number of fossil New World forms some of which 
lave been gcnerically separated under the names of J/fmiauthenia 
rkmia, Pratauchcma, Ilalomeniscus, and Eichniius. In the 
II. 3 F 



existing species, which occur fossil in the caves o( Biazil li 
molars arc normally ? in number, but there is oftn but 


these teeth in the lower jnw ; in A. (Pa/avehenia) Magna b< 
Pleisioccne of Mexira, there were always nro and ocaiianall 
lower premolars ; while in a Pleistocene South Amencaa 
{//emiaa^Aenia) there were three premolars in both Jaws. 
{/M&meHtseut) hi4Ur»a (fig. 12 ro, a), from the Pleistocene o( 
America, on the other hand, the premolars were reduced 10 

each jaw ; while v^. (£*( 



vitaJieriana has the 

ber, but is distinguished 

simpler nature of the npf 

The ranjorilyof the spec 

of comparatively small \ 

A. magna and A. JusSm 

equal in bulk 10 the Ca 

the present day. 

Famii-v CmvinA.— T 

ent and three following 

of the suborder col 

constitute ihc Pecora 

cent Zoology ; — a git4 

defined at the preMl 

but, as already nU 

conneacd in past epw 

intimately by this (am 

the Trof^lida. The i 

of the entire group ifi 

.0 (0-1) s 

/ ., c. — . y>.. J 

upper ineuors being \ 
atjsent. The true tn 
ixrrfccily selenodoni, 
upjjcr one* carry four 
thf second premotar 
in contact with the itui 
is a loi^ interval, or 

Pig. wii.~*. t^nalHp«ioflk<ri(hiDuniH bctWCCn the loWCT 

Xivn v^tmu rta^mt-i- Kejuod «,Cupu>i and caoine ; and tne 
».T»»u.;f.M.up«iiu» (««>«.*««» approsimalcd to, 

resembles the ina 
third and fourth mctapodials (fij{. isii) coaIc»% into & 
bone; ihe navicular nnd cuboid of the lamui are likrwi 
and the odontoid process of the axis ii'crtcbni forms a 



[iodcr. Moreover, in ill existing iiieiiibers of this gruup 
tefal meupodiaJs are either incomplete or totnlty wnnting 
111); the stomach Js composed of four complete cavities; 

pbcenta tlei-elops structures known as cotyledons. On the 
(ithcT horns or antlers arc very gencraUy present, at Icaiit in 
lies of recent rorms. 
the Ctrvida upper canines arc generally present, although 

few exceptions they are of comparatively small st/c; the 
Ktth. and more especially in the earlier forms, are very 
Jly of a more or less brachydont simcturc, the first true 
being invariably of thi« type ; and the upper premolars 

have both an inner and an outer cuhitnn, and are 
simply secant like those of the existing TraguMa, !n 
mil there is always a large vacuity in front of the lachrj- 
rhich prevents that bone from aniculaiing with the nasal, 
large number of forms antlers arc present in the male, 

Jtamgi/er also in the female. .Antlers, it may be ob- 

are outgrowths of true bone arising from the frontal 

:imi~ll«dMBdMllueialwpeciorihcikull«rihc Xocbnek (rajMrWM c^^m), 

of the skull (fig. i3ii), which during their developmenl 

Sled with a vascular, hairy :^kin. On the completion of 

tth a constriction of the lilood-vessels near their base is 

brou);ht about by the fumiution uf u Imrr, and above 

Milt the skin peels off and leaves the bone bnre and in- 

aftcr a time the antUi is sheet, leaving a mure or less 

pedicle attached to the skull, from which a new antler 

Joped. In young animaU the antlers arc simple, and in 

Ipecies in which they finally attain a great complexity, this 

gradually in successive annual growths. Each antler 

^consists of a main stem or Nam, and usually of one or more 

or tiHU ; of which the one luiiucdiatcly abm'e the burr 


is termed the irow-tine. The lateral digits are nearly 
present, and the distal extremities of the metapodials may 
served. The existing Deer have been divided into the Pie 
carpalia (Cervus and Cervulus), and the Telemetacarpali: 
Capreolus, Cariacus, and Rangijer); the former, which m 
habit the Old World, characterised by the retention of the f 
and the latter of the distal extremities of the lateral met 
As in many analogous instances, the development of the a 
the individual is paralleled by their development in the 
since we find that many of the earlier members were to 
provided with these appendages, and that their extreme co 
in the more specialised forms was not acquired until a U' 
in the geological scale. 

The least specialised members of this family form th 
allied extinct genera Amphitragulus and Palaomeryx. In tl 
there are four lower premolars, and antlers were entirely ah; 
crowns of the molars being low. The largest species was s 
bigger than the Musk-Deer, and the genus is characterist 
Lower Miocene of the ContmenL Palaomeryx {Drem 
Dicroceros, and Micromeryx being included) has only thi 
premolars, except in one species ; and the upper true m' 
1213), like those of Amphitragulus, were brachydont, an 
distinct accessory column between t 
crescents. In one species {P. Feign 
lateral metacarpals were perfect, alth< 
slender, and the males had long uppe 
like those of the Musk-Deer, but m 
this species being the earliest, and 
in the Lower Miocene of France. I 
calus {Dicroceros ekgans), of the Mi( 

FiB,~-Le(i upp« *^"«' s^P'^ '^"t'«" '^^^> however, 
trur molar of PaUromirrx and the caniues wcrc apparently sn 
lira iHsis. loccDE, n- ^gj^fii^ of the Frcnch Middle Mioe 

P. sivalensis (fig. 1213), of the PI: 
India, were as large as a Red Deer ; and the latter species, 
with another from the Pliocene of China, were the last rt 
tives of the genus, of which the latest appearance in Eui 
the Middle Miocene of Sansan, in France. Plalyprosopus, 
latter beds, is distinguished by the great projection of th< 
the mandible. To the existing Oriental genus Cervuhts, 
the molars are more hypsodont, and simple antlers moui 
long pedicle are present, may be provisionally referred C. 
ceros (fig. 1214, a), of the Pliocene of Eppelsheim. Then 
uncertainty as to the earliest appearance of the genus Cerv 
not improbably dates from the Middle Miocene, and was j 

b«cr Pliocene ; at the present day it ranges tluoughoui the 
ic r^'on, but is unknown in the Ethiopian rcj^ion and the 
ort of America, This genus, as wc have already observed, 
to tiM PlcMomctacorpAlian section ; the antlers may be of 

t, Amlci of dn Red D«ci (ffraot tlaftuii in ih* ircond t^ax ; u. Anilcr of iht 
W f -f»a»n ctftditioB : o, Anil*r *iul Uwy prdii:!* of iW rnxiul boncofclic Munljak 
MObUl r.Aoikrof tlK r>Iloi> Xi«aitfvm4Ama% 

complexity; and the hinder molars are cither brachy- or 
It, there being frequently an inner accessory column in the 
e molars. 

Ill) is divided into x number »f groups, nf which ilic diMribution 
be bricflj" noticed. The most aberrant is ihc TtlrmieroliMe 
mcd by Cerrus Utnutrof of ihc Krcnch Pliocene, in which ihc 
>pTo\iinaic lo those o{ Caiituus. The AiriMe group, now con- 
he Orirmal reicion, in which the molars arc more hypsodoni 
U nnd the anilcn iirc rounded and cmnparativdy simple, is 
■d by several species, such as C fianiinrMsii {Sg. 1214, ni, m ihc 
>f Europe, while the living C <Ln's occurs tn the Plciiiiuccnc nf 
The Oiicntil Ruifnine group has a reprcscnlaiivc in C. siiti- 
the PHocenc of India ; while the allica RMsine group, of the 
on. in whifh ihe rinilcm arc still compiirativcly simple, and 
beam nftcn smmt^ly ifiouvcd, is knnwn in ;i fiiiMl «liiie \>y the 
f i;xisung species ftom the Pleistocene of India. In the AVn- 
iPt which includes ihc Canadian Wapili \C. ittnatfetnis), ihc Red 
lapkut) of Europe and North Africa, logcthcr with some large 
»m the ral»rarciic region, the antlers (fig. 1114, c. d), although 
led, arc often cupped at ibeir KurnitiiH, and carry a second, 
r, immediatcty above the l>row-tinc. In ilii> ymup remains 
n the existing Red Deer {C. elaphuH are tif common occuaence 
istocene of Europe. Some of the fossil antlers and jaws indi- 
»cr, much larger anhnala than any Ked Deer now existing. 


nnd it has been suggrM«ri tlini thew renuinc belong eiARll( 
martU nt I'rnia, or to the Wapiti (C. iamtdftuit) <M Nanh/ 
Both the«e f:>nns are. however, closely allied to the Red Dar,l 
ajipeart preferable to rcjtard all ihc European foisils ai 
single species from which the three exisiinfj ij-pM are (tal- 
is, however, the name C. tfitSaut for the laige fossil form i 
earlier dAie than the nnm^ C'. tttanW}, if that be really ijiilli 
tiaphus. Kem;tin£ r>f the Wapiti are recorded from the llewo 
the United States. 

The EucliiJoniraUm group comprisea C. Sedg^iH of the 
Foresi-bcd and the Upper Pliocene of Italy, io which the iMh 


: 1 >> 

fix- u>3-— SUktoaef ibcIfiitiUea(LVmufuH«ratl; ftomtta PtoMacsMrfj 
Cre*ily rwjoocd. <AftH Owib.) 

more complex than io any other speciu. In tbe Damim 
intlers (lie. 1114. F) have their lenninationt paliiiaied, and (hel 
iimple ; ine cxi»tinK Fitllow-Decr {C. dama) occurs in ihe bou 
Gibraltar. *hile allied fnrms arc found in the Ent;hsh FoteM- 
Cra^ ; (he most notewonhy b<Mn|; C. vtrtuemis, in which the btv* 



t ikitjjy dofL-nwards. The last group which tl is necessary to 
ta here k ihc Xft^meroHtu, whiirb coninins only ihc Irish Deer 
fgigaiitfus, fig. I3IJ), rh^antctcriud by its enormous palTrmied 
1(^.1116), vhich diverge at ri>,-ht angieK from the plane of the 
^ud have a dUttnct brow> [Sr) and bei- (St) tine. a.nd a small 
9 li« [/'H on the opposite side of the beam (o the bw-linc. 
f of this, fine species arc found in the Pleistocene of Northern 
■od t,r*i especially ab^indam in the bogs of Ireland, where speci- 
lire bftn f<>und with a tpiead of more than eleven feci between 
gj the Antler*. 

ig to ihe Telemctacarpalian genus Jian^/er, which is at once 
rised by the peculiar form of the antlers and their presence 
CKS, wc find remains of ihe existing Ktiindcer (A. tarandus). 





L— Satl tai mltcn of CrmH gitanttut. Rnlucol. ilr. firaw-i 8»,\ht.; FT, 
feucrioc tint, (Aflcr .Soiii.) 


to tbc higher latitudes of the Northern Hemisplicrc, 
t in ihe Pleistotenc of a large portion of Europe. In Aius 
«r Moose {A. mcuhiis), of the northern parts of Europe and 
i occurs in the Pleistocene of the same regions ; while an ex< 
KMS, has Iwm described from the Norfolk Korestbed. The 
|fig. 1317) have no bcz-, and apparently no brow-tine, but 
led into an anterior forked branch {A) and a posterior pal- 
ne {F), A very remarkable form from the Pleistocene of 
imcrica, described under ihe name of Cerva/cts, appears to 
Afm with Orvui, although it belongs clearly to the Telc- 
palian wction. Thu* the antlers (fig. i j 1 8) are superiorly 
itito an anterior {A) and posterior {P) branch ; but below 
tr there occur two tines (/?s and PT), which Dr Scott 
ji6 probably corresponding to tlic bez- and posterior tiscs 



of the Irish Deer. And we may likewise tntce an intennt 
type in the vertical height of the skull, and the form and a> 
tions of the nasal and premaxillary bones. In Capnoha, «be 
antlers are simple and rounded, the existing Roe (C. a^ 
1 2 1 2) occurs in the European Pleistocene ; while C. eusams, 
French Pliocene, is regarded as the ancestor of that specie 
the peculiar C. Matheroni, of the Lower Pliocene of both 1 
and France, is provisionally referred to the same genus. Ct 
again, which is peculiar to the New World, and is cbaiat 

Fig. 1117.— SkuJ[ and anllen of the Elk {^Akii maiJUa.) Reduced. j1, Anta 
B, Poileriot branch. {Alter Scott.) 

either by very simple prong-like antlers, or by a more compl 
totally unlike those of any existing European members of the 
is represented by several existing, and perhaps by some 
species in the Pleistocene of South America. Lastly, it sh 
observed that antlered Deer occur in the Teitiaries of 
America, and the name Biasiomeryx has been applied to 01 
which is regarded as the ancestor of Cariacus. 

AH the preceding existing genera belong to the subfamily C 
but the Musk-Deer {Afoscftus), of the Himalaya and regions 
northward, is the type of a second subfamily — the Mosehina. 



, to which there are no antlers, and the upper canines of the 
tiOain an excessive development, not improbably occurs fossil 
tniocene of the Siwalik Hills. 

ILV GiRAFPiDiC — In this family, which is taken to include 
■ SiBtUkeruda of some authors, the cranial appendages, when 
a{^>ear to be intermediate in character between those of 
%pmda and Cervida. The teeth are more or less brachydont, 

I invested with a rugose enamel ; their number being / -, C. -, 

t, — , Af. -. The type genus Giraffa {Camehpardalis), which at 

I present day is represented only by G. camelapardalis of Africa, 
dosely allied to the Cervida, in which family it is included by 
ifessor Riitimeyer ; the frontal appendages consist of a pair of 

;. 1118 Skull and MalWnot CtrtMtktt amtricattu : from the Pleutocene of Nonh America- 
Reduced. Leitcna* in Ggt. i>i6 and 1317- (Afi« Scolt.) 

ort, erect, bony processes, at first connected by suture, but subse- 
eatiy anchylosed to the skull, which are covered with hairy skin, 
d are present in both sexes. Anteriorly to these there is a median 
xess on the frontals and nasals, which is sometimes termed a 
rd horn. There are no traces of lateral digits ; the humerus has 
double bicipital groove ; there is a lachrymal vacuity in the 
mium ; and the neck and limbs are enormously elongated, 
ssil species occur in the Lower Pliocene of Greece, Persia, 
dia, and China. VishHutherium, of the Siwaliks of Burma and 
dia, appears to be an allied genus, with shorter limbs, but the 


cranium is unfortunately unknovn. The; next place to iht i 
is occupied by fftJ/aJotfitrium, of the Lower Fiiootnc oi i 
India, and perhaps Persia, in which ihe cranium is dcroid i 
peiidagcs and the molar tectli become more like thoie of i 
The hnibs are comparatively shon and «toui ; the cnnium I 
lachrymal vacuity ; and the one known species was of 
greater bulk than the GiralTe. rt uli Hydaspilktrium, of ihe 1 
of North-western India, we enter the group in which ihd 
was provided with large branching antler-like appendagtv i 
the exact nature of Ihcir covering is unknown. These ip 
in this gcnuK ri^ firom a common haiic situated imn 
advance of the occiput, hut their form is not known; 
lachrymal vacuities in the cranium. An apparently dlicd^ 
from the Phoeene of Persia, has l)e<n named Vrmialk 
Br&malherium, of the Pliocene or Western India, llieit 

- / 


Fig. tn^^—fiVvS^tX Shxtthfium tic»xttMmi radaccd. raocvM. tudib 
■lit liiiidar anilan tlwiiU prakoUy b« ri^w— I. 

pairs of these anilcr-Iike appendages, the anterior pair . 
a common base, and being of large siie. Lastlv, we Iuny .* 
ihtrium (fig. 1319), of the Pliocene Siwaliks of Konh-ca»icm Ii 
in which Ihe neck and limbs were not develt^xx) beyond the doi 
proportionn. There arc two pairs of cranial appendages, the \ 
of each being separate. The anterior pair are conical, like dM 



'; vhilc the posterior oiiea arc palmate, and resemble the 
. of the Elk. The latter pair are marlcecl by the impressions 
blood-veucU, as in the CtrviJa, but do not show the burr 
tic of that fannly, from which it ts inferred th;il they were 
There is no lachrymal vacuity in thccntnium ; the nazals 
; and arched ; and the bones of the skeleton approximate 
lUre (o iboee of the Cervi/ia. Some authorities regard this 
\ti tDOal closely allied to brachydont Antelopes^ like Sirtpsi- 
\hai it appears to be so intimately connected with the preced- 
ihat i[ seems imperative to place it in the same family, 
I it may indicate an approximation to the Bsvida. 
remarkable genus Sttmolfurium, of the Lower Pliocene of 
of SamoK, i« referred by I>r Forsyth -Major to the present 
f, although its skull mnkcs a remaikable approximation to that 
ameiopotd (;enuj Palaolragus. The females were homleKS, 
males had a pair of small boms immediately above the 
t. The molars arc descnbcd as being very like those of the 

JiMlLV Ajstilocaphid.k, — This family is now represented only 
be American Fiong-buck {AniilMaJrra), in which ihc horns ate 
le same ruture a* in the /iirvidir, but differ in beiny bifurcated, 
in the shedding of their »heatJi. Remains of Antilo<apra occur 
le Pleutocene of North America ; and it is thoujjht that Cosoryx, 
Ik Pliocene of the same country-, ma>' have l>cen the direct an- 
W of the eiiHCing genus. 

UiiLV BoviD£. — In this, the last, family arc comprised the 
[ specialised members of the whole suborder, such as the 
slopes. Goats, Sheep, and Oxen. 'Jhe genera] characters of 
greater part of the skeleton are the same as those mcntioiwd 
»■ the head of the Certfida ; but a Temarknble difference is 
d in respect of the frontal appendages. These appendages 
I3J2) are paired, and consist of persistent bony processes, into 
h the air-cells from the frontal diploe often extend ; they are 
imlly subconical or triangular, and often twisted, but never 
cbed. These " horn-cores," as they arc termed, arc covered 
the true horns, which are composed of an epidermal fibrous 
rturc, and arc never shed. 1'hc males of all existing genera in 
wild state are furnished with these horns, and they are also 
eni, although of sniailer site, in the females of the great 
irity. In certain domesiiciiled races of so-called /c//f^ Sheep 
llizo), Goats, and Cattle, the horns are, however, wanting in 
uexes; and this peculiarity it; with great probability r^arded 
Hnstance of reversion, since these appendage:^ arc also wanting 
Ikne allied Tertiary' forms of two of these groups. In the 
iam there is gcneiially no lachrj'nial vacuicyg and the lachrymal 




consequently anirulates largely wiih the nn£al (fig. il»};l 
certain Amclopt-s this vacuity is present, when ibe rchtioBirfl 
bones are the snme as in the Cervide. AnMher very i 
feature of the Btrvida, u a whole, is the hj-psodont 
ihfir check-tlen[ition. In many of the Antelopes (fig. null 
feature is only moderately dcvcluped, but in the Sheep, Goatti 
Oxen it is carried tu an excessive degree ; and in the tuie . 
the valleys of ihc teeth arc filled up by a coating of cemeaL 
feature, like that of the relations of the lachrymal, is, farafR^I 
absolutely dtsttnctive of the BoviJa, since vc find xaart) K 




Fig. iT3D>~-SkuUaf« homliH SS<«|i{<M>V Radood. i, liKhon ] t, CbbIboI «( 
l<clh: Mai, FrcmiuillA: mj, UuiIIb; im, IfuaJ: t, Laduym*] ; >■. iMal: yV, rn«^:J 
ruisul. [AfUrOwM,) T 

(and especially those having a 1achr]rmal racuity) with a 
Iirachydont dentition. The upper true molars frequently 
large accessory inner column (fig. 1231). Kunctiunal canios : 
wanting in all cicisting forms. The lateral digits may or may 1 
be present, but in no living form is there a di:ttal Tcttuunt of thi' 
lateral metapodinls. 

This family does not apparently date Further back than tbt 
Middle Miocene, where it k represented by members of its lost 
s|>ecia]i^ed ^roup, the Antelopes ; the Sheep and Oxen not ip- 
pearinjf till the I'Hocene. It has been suggested that the toulf 
orij{tnatcd from the same ancestral fomifi as the Cenida, 

Commencing with the Antelopes, and confining our attcntitn <» 
those forms in which the genus has I>een more or less oxamU^ 
determined, we find in the Akflafhint section, which ts ooii cob- 



Africa and Syria, the exuting genus Ahefaphus represented 
cf Tcniahcs of Algeria, and also in the Pliocene Siwalik^ 
; the species from the latter deposits being apparently 
th to the Hartebeast [A. Mama) and the Rontehock (A. 

b These antelopes have recurved or lyre-shaped horns ; 
has no supraorbital pits, ,ind Tn.ty have a very long 
"Ac molar teeth are very narrow. In die CefiAuUfiiiu 
compn'sing Indian and African species ot compsiatively 
le, the eiistiiig Four-horned Antelope {Tefraceras ifuadn- 
found in a fossil state in the cave-deposits of Madras ; 

$nt liairinil itirtirrlr-i nf tht twond leri u|ipi-r iroi moluorilic NLUhtl 

I extinct species occurs in the Siwaliks. The African genus 
ims may perhaps also occur in the latter deposits. The 
W« section is now confined to Africa, ami includes some 
» antelopes in which the females arc hornless. Cof'us, in 
\k *kall has welt-inarkcd supraorbital pits, appe;ir«. 10 tte 
\tA in the Indian Siwaliks by species as large as some of 
can forms. In the Pliocene PiVerini beds of Greece an 
with round lyrate horns, desrribed as fUlui>ph^ra} appears 
(ied in some respects with Cobus, alih^ju^h it !ias lachrymal 
^ but no distinct Kupraorbilal pits. In the* t)-pical Antehpin^ 
which is allied in many respects to the preceding, there arc 
tretl-developed supraorbital pits, and the molar teeth are 
W, and resemble those of the sheep. The type genus 
, which has round and spirally twisted horn.'i, is known in 
Itocene of India by remains of the one exiating species A. 
It (Blaek-buck) ; the existing Siberian Sm^a larlarua is 

OHicisaJly dochbed under the picoccupied mine of fltiinurm. 



found fossil in the cave-deposits of Europe ; while the Aft 
genus Gatella, in which the lyrate horos are laterally coi 
occurs in the Pliocene of Europe, Africa, and India, as ' 
the Norfolk Forest-bed. The Hippotragitu section maybe 
include the existing African genera Oryx, Addax, and Hi} 
and is characterised by the long and straight, or backward 
horns, the absence of supraorbital pits in the skull, and 
and hypsodont upper molars, which resemble those of 
In a fossil state this section is represented by ffi^dra, 
Indian Siwaliks ; and also by the extinct Palaoryx, of 
Pliocene of Greece, Italy, Samos, and France, which a 
have been closely allied to Oryx, although showing some 
tfippoiragus. The last section into which the true axis 
lopes may be divided is the Tragelaphine, comprising Boi 
India, and Tragelapkus, Strepsiceros, and Oreas in Africa 
Indian genus, of which the Nilghai is the only existing t 
tive, the horns are short and upright, and are not presi 
females, while the dentition is hypsodont (fig. 1321); fo 
occur in India from the Siwaliks upwards. In the Afri 
the horns are spirally twisted, with two more or less wi 
longitudinal ndges, the skull has deep supraorbital pits ai 
mal vacuities, but no pit in the lachrymal itself, and the 1 
broad and brachydont like those of the Cervida. ^ 
(Kudu), in which the anterior ridge on the horns is 
stronger of the two, apparently occurs in the Indian Siwal 
may also contain a representative of the allied Oreas (Elai 
extinct Pa/aoreas (fig. 1222), of the Lower Pliocene of E 
Algeria, appears to have been allied to both the precedin 
while the so-called Antilope torttcorms, of the Pliocene ( 
has the posterior ridge of the horns the most developed 
existing Tragelaphus, to which genus it has, indeed, beei 
The remarkable Protragtlaphus, of the Lower Pliocene c 
differs from all the preceding genera in that the horns h; 
posterior longitudinal ridge, in the absence of supraorbita 
in the development of lachrymal depressions like those o 
vid<E. With the Rupicaprine section of this family we come 
showing characters connecting the true antelopes with I 
but the only definitely known fossil remains belong to th 
alpine Chamois {RupUapra), which occurs fossil in the cav 
of the Continent. Under the name of the Palaotragii 
may be included three extinct Tertiary genera having th 
compressed horn-cores of the goats, but the upper molai 
less like those of the brachydont antelopes. The earlies 
genera, and indeed of all the antelopes, is Protragocer, 
Middle Miocene of France, one of the species having 



described as Antihfe c/avata {sansani'titstt). These were 
I ftfms with short horns, and the crow-ns of the molars very 
Iffid moderately wide. In Pa/aotniffiis and Tragoceres, of the 
PHocciw of (Greece and Samos, the horns were larger, and 
' JUlars wider. The former genus is considered to be allied to 
\tritnn, noticed among the Giraffida. The tnie (>oat5 and 
, collectively forming ihc seciion Caprimr, iire charactcri.-icd 
more or less laterally compressed and ofitn aiijulatcd horn- 
!iich may be either curved backwards, as in the Ibex, spir- 
ited, as in the Marlehoor, or with a peculiar outward curvaitire 
I twist, as in the Sheep ; the horns themselves being frequently 

rrt- — \jt^ Uwnl tIvw of th* oaDhiBi ot J'ainrrr^ t.tudmKitjnrl: fram ihc Lomr 

on the anterior surface by transverse ridges. In all the 
waa the dentition is markedly hypsodonl, and in existing forms 
accessory inner oolumn of the upfwr true molars is wanting, 
one of thcra show a lachrymal vacuity : but in the Sheep there is 
ly a deep depression (larmier) in this bone, which is absent 
. tfae Goats. Capra may perhaps occur in the Upper Pliocene of 
it is represented in the Pliocene of the Indian Siwaliks by 
ic species IC. sita/ensis), which is probably the ancestor of the 
Himalayan Thar {C. jfm/oira) ; by another species equally closely 
allied to the Matkhoor (C> I^tontri), of the same region ; and not 



improbably by a third allied to the Himalayan Ibex (C. si 
Remains of the Pyrenean Ibex (C /yrenaiea) are found 
Pleistocene cave-deposits of Gibraltar ; and those of the a 
Goat (C. hireus) in the turbaries and fens of England. T 
remarkable hornless genus Buea^a, from the Siwalilu of Ini 
a skull presenting a great resemblance to that of the Go 
cheek-teeth like those of the Oxen. The true Sheep {Oris) 
to be a group of very late origin, and are scarcely known in 
condition ; a large species has, however, been described 6 
Norfolk Forest-bed as O. {Caprovis) Savigni, which was ap; 
allied to the existing Argali. The Musk-Ox {OvtAoi) of tbi 
regions, which forms a connecting Unk between the Capr 
SovintB, occurs fossil in the Pleistocene of Europe and 
while two closely allied forms, from the Pleistocene of K 
and Arkansas, have been respectively named O. {Boothtrittm 
frons and O. cavijrons. 

The members of the Bovine section, comprising about 
recent species distributed over the greater part of Eurofi 
Africa, and North America, agree with the Caprine section ii 
no lachrymal vacuities, but differ from the recent members 
section in having the crowns of the cheek-teeth extremely t 
large accessory columns in the upper true molars, and thei 
filled up by a large quantity of cement The hom-cores 
either rounded, flattened, or angulated, and are frequently 
more or less outwardly, but are never curved spirally inwar( 
the " cork-screw " shape characteristic of many Goats ; w 
horns themselves are not marked on their anterior surface b 
nent transverse ridges. The most aberrant genus is Lepto. 
the Pliocene and Pleistocene of India and the Upper Plit 
Italy, in which the frontal portion is broad, with widely si 
horn-cores placed far below the level of the occiput, Tl 
cores are sometimes absent \ and this genus is r^arded as 
Bostlaphus. Bubaius, typically represented by the Buffaloes 
and Africa, but which may also be taken to include the dii 
Anoa {B. depressicornis) of Celebes, is characterised by its ai 
hom-cores, which may be directed either outwards or upwa 
by the great convexity of the forehead in the more tj-pica 
Among the more aberrant species may be reckoned three ( 
Siwalik Hills of India {e-g., B. occipitalis), which are close 
to the Anoa, and (together with that species) by some wri 
termed Probubalus ; the hom-cores are frequently compleW 
gular in section, and the forehead is not decidedly con\ 
platyceros from the same deposits is intermediate between 
mentioned group and the existing Buffaloes. B. antiquus \ 
Pleistocene of Algeria is regarded by Professor Riitimeyer a 

he existing African Buffaloe*, (ilthoujjh Dr P. Thomas 
I more nearly rclulcd to the living Indian sptcies. The 
iit/fAtt) a found tn a fossil state in the Pleistocene of tKc 
'alley. India : while an apparently closely allied form also 
the Pliocene of the Siwalik Hills. In Bison, now repre- 
ihc Aurochs of Lithuania {/?. iiomutts) and llic nearly 
ed North American Hiton ameriianus^ the skull is chninc- 
in great relative width and shortness, the tubular orbits, 
iiely convex forehead, and the curved, round, hum-cores, 
pbced considerably below the level of the occiput. The 
uropean speetes is represented by a \-3rieCy {l>riseus) in 
cene of Eurojic and Arclic America ; while the gigantic 
* of the Pleistocene of Texas may urohably he looket) 
■c progenitor of the recent apcric.') of that country. A 
om the Pliocene of the Siwalilc Hills has been referred to 
with the name of B. sivalensis, and appean to be allied to 
rms. The genus: 7:^^, which is confined to the Old World, 
it spedali^cd representative of this se<.'tion, anU may be 
to the BUhn-im and Tiurinr groups. In the former are in- 
wild Oxen of India and Bunna, which are characterised by 

HuEh raductd. 

Battened horn-cores, and hy certain peculiarities in 
occipital region. The earliest representative of this 
W tirusaii from the Upper Pliocene of the Continent, in 
hortvcores are pLiced very low down on the fronuls ; this 
i^ considered 10 be nearly related to /!. l^anftng of Burma, 
iljerfecily known member of this group is B. falirosaurus 
fitovcne of India, which iua.y turn out to be identical with 

a G 




B. gaurus now living in the some tegiomt. In the 
the frontali (I'lg. laaj) are extremely elongated, and the 
which in the type species arc rounded, aic placed inuni 
the occiput i'o this group may Irc referred B. fifamfiv 
(Uutijrvns of the Phoccnc of the Siy-alik Hills ; the UtlCTi 
markable for its sharply angulated frontals and its enoni 
cores, which liave n pyrtform section. Another membet d 
dUui of ihc Pleistocene of Central India, which present! 
proxiination lo the Bibovim group ; but the hevt known H 
the Vrus of tlic European Pleistocene {fiy. i aa^), which a 
be only ji larger form of the existing Ox {^Bps fauna), aoi 
the descendants of wild races an: still preserved in Chillii^ 
some other Briiinh parks. A still smaller race, whoic rcij 
been found in the Turbaries and tens of England, .iihI 1 
described under the names of B. longifrons and B. 
seems lo be only a stunted variety of the same specie*, fra 
is probable that the small cattle of Wales and Scotland 

SunoHDER ». — pRBissoDACTYLA. — The Characters pa 
this suborder in common with the Artiudactyla arc noti 
that head. The disiincii^e Teatures of the Peris<iodactyb 
found in the truncated di&tal surface of the astra^calu^ ({ 

ihe circumstance that the 
in tioth the fore and hind II 
metrical in itself, and li 
either of the others (fig. i 
presence (except in Ckal 
of a third trochanter to 
(fig. i2]6): and tlie non-a 
of the fibula with the ci 
Other characters very gei^ 
setvahle in this suborder ai^ 
whole of the scries of cheel 
in contact with one another! 
upper premolars are near!; 
as complex as the true mt 
the last lower true molar frequently has no third lobea 
when such third lobe is present, it is absent in the last fi 
molar ; while the first tooth of the cheek series is sooil 
ceded by a milk-tooth. In .ill existing fomts the n 
dorso-lunibar verlebr.e is never less than twenty-twof, 
twenty-three; while the nasals are expanded posi 
stomach is simple, and the placenta dilTuscd. In exiai 
the cervical vertebra; are markedly opisthoccelous. 'Hie < 
molus are constructed on some modification of what is 

Mtriii-iliit I'F [lie licitc (Afiiiu Oit*i- 



WpUn (Rg. 133^); that ii, th^re is an outer longitu- 
fcpm wliich two transverse ridges proceed at right angles 
Bner border of the crown. In the bnchydont forms 
re is perfectly simple, but in those genera with very 
leeth it U so comphcatcd by foldings and involutions 
t iilwA>'s easy lo trace the original plan. The crowns 
true molars consist in their amplest stnicturc of two 
idges (as in the Tapir), bm these ridges may Ijc cun'cd 
(as in the Rhinoceros), or complicated by foldings 

bhl BUM at (At Tm^rut. iri (■) 
a*. lUduoed. (AAti flowcrO 

rif. nni, — Donal or Mitirior 
vinr of iti* lift fanur of KUmf 
etrti. Tat mKtian pniJKiiiin on 
tli« rifhi «<It of ilie Bku'i it ihv 
Ihinl UDchumcT. Kedu«d, 

itions (as in the Horse). The transition from the aim- 
^dORt to th« most specialised hypsodont dentition is 
i by a reduction of the number of the dij^its from four 
one ; thai one being ilic third, or middle, of ihc typical 


tsodactyla have RufTcrcd considerably more in proportion 
ibers than the Artiodactyla by the extinction of generic 
types ; the existing genera being at the present day 
EhTCC, which are the types of as many difTeient families. 
s bare suggested that this extiiiciion of types is owing 
lOdonl plan of molar structure beinjj less readily suscep- 



tible of modification than the Bunodont type upon which the Arftt- 
dactylate molar is constructed, but this is really a pure assumptiaa. 
Family Tapirid-e. — This family is represented by the sin^ 
existing genus Tapirus, now found in the widely separated areas cf 
the Malay Peninsula and South America, and thus aiTording an 
excellent example of what is termed discontinuous distribution. Tie 

dental formula is /. ^, C.-, Pm. ^, M. ^ ; the cheek-teeth ae 

3 ' 3 3 

brachydont, and of the simple Lophodont type ; the hinder pre- 
molars being as complex as the true molars, and the last Iowa 
molar having no third lobe. The first upper premolar has a dead- 
uous predecessor ; and in the existing forms there are four digib 
in the anterior (fig. 1225, a), and three in the hind foot. The 
cranium (fig. 1227) has its cerebral portion much vaulted, and the 

Fig. i»7.— Side view of the AaW OT Ta/ina amerKmnia. Reduced. (After GiebEL) 

nasals short and arched ; in one American species {which on this 
account is regarded by some authors as generically distinct under 
the nair\e of £/asmog-na/^us) the narial septum is largely ossified; 
and there is a short proboscis. In Europe this genus is found in 
the Middle Miocene of France (T. Poirrieri), and continues to the 
Upper Pliocene {T. arvernefisis) ; it also occurs in the Pliocene of 
(;hina (7". sinensis), and in the Pleistocene cave-deposits of Biaiil, 
one of the forms from the latter being indistinguishable from ihe 
living T. americanus. The North American Miocene forms named 
Tapiravus are probably not generically distinct. An imperfectly 
known form from the Middle Eocene of France, described as 
Palieotapinis, is referred by Dr Filhol to this family. 

Family LoPHioDONTiDiE. — This Eocene family presents diai- 
acters allying it very closely with the Tapirida, PalaotheriidJ, and 
Mhinacerotidie^ and probably contains ancestral forms of all those 
families. The upi>er true molars (fig. 1228) are brachydont. and 
always more complex than the premolars ; the last lower true mobr 



ncially has a third \o\x ; and there arc usually four digits in ihc 
oaus aiwl three in the pes. The type genus LofihWffH (fig. 
I3&, a) comprbeii some specie^s which attain a bulk tivalling that of 

EhinoctTosts, and has the dental formula /. ■, C - , J^. -, M. -. 
found in ibc Middle an<l Upper Eocene of Eurofw, and is 
ally regarded as having died out without descendants. In 
c dentition here figured ihc inner crcseciil of the fourth upper 
iBDolar is incomplete, and the ridgex of the tower molars are 
Ki{)le; the last lower true mohr always lias a third lobe. Allied 
I iepkiediin Ik ffdaitUt {DeiTnalotherium), of tlie Upper Kocene 
[ North America, characterised by the more roundt-d upper true 
lolars and the absence of a diastema in The lower jaw ; it is le- 

-«• Tba iue live nsbi up|tcr chscli'imih 4( LifAia4an ttitimiti, Irani the Miditic 

■•■ of Piaac*: M> Th« rtchi upp^t thcfli-iwih of HyraiKymi arrmriui. !toax Ihc toc«i» of 
nk AMwta. lUdaccd. 

rded by Prtrfessor Cope as an ancestral stock of the Jihitttxtretida. 
itb fomu from Ihc North Aracnc^n Eocene dcficribcd as htetclo- 
tu and Pr&tkyratodon^ rn the first of which the last lower true 
}Iar has a third lobe, wc come to the contideiatiun of Lophio- 
nts showing more affinity with the Tapir*, but it docs not appear 
itc certain tliat both these forms are really entitled to generic dis- 
cxion from the next. The genus Hyrachyus (fig. 1 33S, b), which 
cuts ty{>ic3]ly in the Uinta and Bridger Eocene of the United 
Ues, is lalcen by Dr Filhol to include European Lophiodonts 
iging from the Middle Eocene of Trance to the Lower Miocene 
St Girand'lc-I'uy. 'ITiis gcnas has four premolars, of which the 
« is somewhat simpler than the first true molar; while there is 
I third lobe 10 the la<.i lower tiue molar. In the type species the 



Upper true molars (fig. laaS, b) resemble those (^ Ha £k 
ida ; but there appears to be a gradual transition in this 
towards ff. prisms, of the Quercy Phosphorites, in whii 
teeth have rounded angles, and approximate to those of the 
from which, however, these forms are distinguished by thi 
fourth premolar. This transition has induced Dr Filhol t 
the genus Protapirus, which was proposed for H. prise 
latter species is evidently allied to the American Iseetolop 
Dr Filhol suggests a transition from this type towards th 
The typical American species of Hyrackyus is rc^arde 
ancestor of the genus Hyracodon^ which is classed in the t 
tida. An American species which may be provisionally 
in Hyrackyus is said to have an attachment for a denna 
each nasal, on which account it is separated by Professor 
Colonoceros. The American Eocene genus Dilopf^don a 
be allied to this group. 

The imperfectly known genus Ribod^n, from the infra 
beds of Patagonia, is apparently nearly allied to ffyratky 
genus Triplopus, of the Upp>er Eocene of the United Stai 
tinguished by having only three digits in the manus, 
account Professor Cope makes it the type of the family . 
idtE. In Hyracotherium, with which the forms describ 
the names of Pliolophus and Orohtppus, and not im[HO 
htppus, are identical, the upper true molars resemble those 
odon, but their anterior ridge is incomplete, and the transve 
of the lower cheek-teeth have a tendency to assume a creso 
In this genus there is a diastema behind the first premol 
an allied form from the Lower Eocene of North America, 
been named Systemodon, all the teeth are in contact, TTie 
species of Hyracotherium indicate animals not larger thi 
the dentition is of the full typical number, and the structi 
fore-foot is shown in fig. 1236, a. This genus occurs in t 
Eocene ^ both of Europe and North America, and, as wil 
fully noticed below, it is regarded as the ancestral sto 
Equida. Heptodon, from the Eocene of New Mexico, \ 
originally identified with Pachynohphus, appears to coi 
latter with Hyracotherium, although it is placed by Profe 
next to Hyrachyus. In Pachymhpkus the dental formula 
full typical number ; but the upper true molars have tal 
than in Hyracotherium, and are intermediate in structun 
those of the latter and of Anchilopkus ; the lower mol 
subcrescentoid. The largest species is P. isselanus, but 

• It has recently been recorded from the Middle Eocene and the I 
or Fiance ; but at least one or the species from tb« latter deposits 
form more nearly allied 10 Anckilophm. 



|] otiiCT forms from ihc Middle and Upper Eocene of the 
Ban ; Pr<>f«liK0th(riHm (in which Lephhlkertum may be in- 
I) does nol appear gencrically separable. 
IniY pAL.r,OTHFRiin^ — Wilh the Palireihiriida we enter 
uftlhcr cxlintl family of ihin suborder, Ihe iy|)e gcniis of which 
Kg b«en known from (he classic labours of Cuvier. In this 
the upper premolars may be cither simpler or quite as com- 
t the true molars ; the lower molars have cresccnioid crowns, 
I the last tooth of this series the third lobe may be cither well 
ped or olmo-it warning ; all the ehcek-tetrtli xk braehydom, 
ben cement is present it docs not fill their valleys. There 
lays three digits in each fooL The type genus J'aUtotktrium, 
th may be included Paioplothtrmm (or Piagiolopkus)^ ranges 
Middle Eocene of the Paris b.^ to the Lower Miocene 
lion, but b especially cha^acll;ri^tic of the Pnrisian stage. 

lUl formulft is / ^, C. !, Pm. *i=i?, M. ^ ; and the last 
3 ' (i-4) 3 

molar has a third lobe. In the more typical species (fig. 7129) 
ptemolars arc present in both jaws, the fourth upper [ire- 

Opiwi EwKiw of Pui*. fteduccd. 

r ts as complex as the first true mutar, and the third lower pre* 
[ OS the fourth lower premolar ; while the diastema is compara- 
fshort, and the canines are not large. The upper true molar!; 
Itbe species exhibit an expansion of the inner extremities of the 
■CISC ridges, foreshadowing the slruriure of the teeth of itoine 
e E^ida. The cranium is Tapiroid in character, especially in 
irotm ncncc of the nasal hones ; from which it is deduced with 
>iUty thnt the nose posses-<c<l a short movable proboscis, 
fomi may also be supposed to have been like that of 

[(ir», and the retioration of P. maptvm given by Cuvier (fig. 

exhibits to us an animal closely similar to the existing Tapir. 

particular instance, however, we know that the restoration is 
rcct, since the discovery of a complete skeleton of this species 
ibovn that it was a more slender and longer-necked animal, 
ibling in its general iigurc a Llama. 

Ibat group, which ii considered by some writers as jcenc^Hcally dis- 
uodcr the njune of PalopiaiJurium, the last upper premolar ha» its 



hinder lobe more or less completely aborted, the third lower pr 
less complex than the fourth, the first lower premolar is absent 
corresponding upper tooth may also be wantmg ; when the b> 
is present it is sometimes preceded by a milk-molar. In this 
yavali, from the Quercy Phosphorites, is remarkable for the p 
a considerable quantity of cement in the cheek-teeth, and for u 
mation of the upper canine to the premolars and its apparent 
from the incisors. P. minus from the Paris basin is the small< 

Fix. i>3o. — Reduced ralmlion cX Pat^clktrium magMum, mlui Car 
Upper EoccDC, France. 

of the genus ; while P. eodidense, from the Middle Eocene of F 
its fourth upper premolar as simple as in Lophiodon and Hyri 
thus shows how extremely intimate is the relation between all 
forms of the suborder, and how very diflScuIt it is to give an; 
tinctivc characters of ihe families into which it is convenicni 

In Anehilopkus, of the Upper Eocene of the Continent, 

formula is / ^, C. \ Pm. -^, M. ^ ; and the cheek-teeth 

intermediate in stracture between those 
nolophus and Anehitherium. The last 1 
molar is as complex as the first true mo 
are well-marked ridges on the outer o 
the upper true molars ; the last lower t 
has a large third lobe ; and the diastea 
gated. l"he genus Epihippus from tl 
Eocene of North America, which is 
by some writers as being on the dired 
of the Horse, has been placed her 
Schlosser. The genus forming a step ii 
of Anchihphus is Atukitherium, typic 

the Middle Miocene of Europe, but with which the con; 

North American forms described under the names of J 

KiR. I2JI. — A left 
upper true molar of 
An<kithcrit4m aureli- 
atitruf, from the Mid' 
die Miocene of France. 



K^^fits may be united. The dental fonnula is the typical 

^; die gpper premolars are 09 complex as the true molsrs (fig. 

, (he external surface being without a mctlian vertical ridge; 

lower premolar U coraparativeiy small ; and the third lobe 

siBt lower true molar is reduced to a siiial! talon ; while the 

have no infolding of the enntnci at their sutnniits. Some 

»hciw a " larmial " deprcs^ian in the 

il. The typical j4. aurthammt (fig. 

\\), of the NUddIc Miocene of the Con- 

Bl, b the largest sjiecies, and shows no 

of the fifth metacarpal \ while the 

and entocuneiform of the tardus are 

In the smaller A, liairJi of the 

of Nonh America the lateral digits 

leUtively larger, the fifth mclacarpul is 

:nted by a splint, and the me«o- and 

cuociform are sepoiatc. Allied to this 

ins are Af>thippus, Parakippui, and Hyt.*' 

Mi of the North American Miocene. 

[i'uiiLY ?RarHKOTHKKiiu*. — In this place 

ly be convirnicnt to notice the ^cnus Pro- 

therium from the Tertiary of Patagonia, 

1 ■»-»$ at fir^l r^arded by Ut Ameghino 

belonging to the Artiodnrtyla, but was 

quenily made the tj-pe of a distinct 

of thi* suborder. One speties wa>, 

l>y Bravard to Anoplothtrium, while 

and was subsetiuently described by I>r 

neister first aJi Ancfiitkerium, and then 

AmisalofihMS. The upper premolars arc 

\y as eomplex as the true niulari. The 

ThoaifKrium, Diadophertii, and Li- 

•ium have been applied to allied forms 

the same deposits. The lowei molars 

four diMinct roots. Il may be ques- 

ned ■whether these forms arc really cn- 

to form a distinct family. 

! Family EQUlD.e. — The division between 

family and the Pa!tTorfunid>t is a more 

f kas arbitrary one. In the preeent one tl]c 

premolars are as complex as the true molars, and all tlie 
•teeth arc usually of an exucmcly hypsodont type, with their 
Heys filled with cement ; the crowns of (he lower molars are cres- 
ccntoid, with complex folds of enamel, and there is scarcely any 
dtstmct third lobe to the la&t lower molar. The digits may be 

Jig. 1131. — Laitnil lipin- iif 
ihc i>kIii iii'ind^ n( (ha KoMa 

irt. Cnr|)u->; •". jd M«taf»r. 

lJ*t lUlllllllll-tllMKK t,\-Utttii 

At:: 1. 1. 1, IliaUnstak oT 



either one (fig. nja) or thicc in number ; the ulna and 
incomplete; and the meso- and entocuneifonn of the 
unitctl. The plane of vrcar of the cbcck-tecth becomo 
smooth, insteAd of being raised into ridges a in the 
fomilifs; and the summits of ihc inciBors ha*"* an infoldiftj' 
enamel exlt-nding some distance into the crown. One of i 
generalised forms is Protohipf^i or Meryihipfxs of the 
Pliocene of North America, in which the pennancin nietai] 
ficmhlc those of the generalised species of E^vut, but have i 
crowns, while the milic-molars ajjproximaie more n<urly la dKl 
molar* of AmhUherium. The next genus '\i Hiff^rion \^ 

ikerium). The dentition is / ^, C ', Pm. -, M. ^ ; bnt Ael 

3 1 3 } 

upper cheek-tooth, which has no predecessor and oppean i 

the milk scries, is shed before the animal is adulL The Bf 
cheek-teeth (fig. 1233) at first sight seem to differ rery n 

f d * i 

T\%. iij).— Thrac ri|ht amm cittrk-inih of Mifftwtf* ; tnm ih* Pbicaa rf IXt 
PotMrior. ruxl t, AntartM «il«r «fttMi*Ai ; t, AaMtfor, aMl d, Aucn«t Innn ooCMi : t. 
IMC, mtiA/, PosicriM pltlar. 

from those of the Paiaothcrinm type (fig. ^^^^\^ but a c 
examination will show that the outer portions marked a soAi 
respond to the outvr wall of the more ger»eralised tooth ; whA 
portions < and t, and d and / respectively represent the fit* 
second transverse ridges of the name These ridge* ha>-<, bOW 
united together in a crescent-like form, and enclose between I 
and the outer wall a pair of islands surroundnl by a plicated «i 
enamel and filled with cement. The terms which it b coofd 
to apply to the Equine molar arc irtdicatcd in the accompti 
figure; and the distinctive feature of the upper teeth of Itif^ 
is thai the anterior pillar (c) is disconnec-tcd from the auRrior) 
ctcscciit (r) for at least three-quarters of its height, so that il 
mally appears on the worn crown as an isolated oval (fi^ il 
There arc normally three digits to each foot, but in the PIk 
Indian Ji. anlthpinum they aT« apparently reduced to ooej 



I Wing been made the tj-pe of the geniii; Hif'podadyhs. The 
kkiiown species is //, grmiU of the Pliocene of Europe, Samos, 
,ind Algeria; but the genus is aljui well represented in the 
of India {H. antihfmum, H. TheaMdi), China [H. Ruhl- 
. and Xonh America. All the sjx-ne* retain the primitive 
of a depression in the larhrj-m.ii. With £^1110: (in whirh 
be included the American Hipptdium. otherwise Pltohippus) 
10 the most specialised of all the PeriBsotUictyla. The 

is /. ^, C. -' , Pm. lill\ M. ^ ; but the first upper check- 

3 I (3-4) 3 

ah i« usually absent in existing forins, and the corresponding 

one is only occafeionally developed in sonic axtincl species. 

' crowns of the cheek-tetlh ate higher than in Jiipparion, and 

' antcnor inrvcr pillar of the upper ones, except in a very early 

of wear, is connected with the adjacent inner crescent (fig. 

J4)- There is but one functional digit to each fool, altliuu^jh the 


iximil portions of the Intenl metapodi-ils remnin (figs, la^a, 
^36, d). and in the so-called Hippidtum the terminal phalangeals 
reprtnenled by claw^ A maxillo-bchrytnul fossa ik present in 
extinct E, andtum and E. stvelensis, but is wanting in all cxist- 
species. At the present day this gcnuH is confined to the Old 
Id, and is especially charactcrisUc of Africa, hut in ihc Pliocene 
Pleisioceiw it was spread over both North and South .America. 

those South American Pleistocene forms rcfcnrd by >onte wriler* 
'ippiiitifn, the molars are shorter ;in«l more cun-td than in exi^tin^ 
ict, and Ihc grindiag surface uf the antcnor pillur uf die upper ones 
;0o« wider than in Hipparicn ; £. primipulis h a large 6pccics 
lis lypc. In E. SttHonis of the Upper Pliocene of Italy, Kos, and 
ria, and the Norfolk Forett-bcd, the molars ore taller, but th«y still 



have a n.irrow ajiterior pilUr in the upper jaw, and thus chowl 
n«clion with Hippurion. In E. si\<altniis, of the Pliocene of I 

A". quageintUt of that of Italy, this pilUr becomes rather more ■. , 

Mid in inc Pleistocene E. nttmadian of India, as well «s Lti E. nvnl 
nf thai »r liratil. and all the existing members of the genus (fig: | 
(he |;rindiii){ lurlace of thiii pill»r b«cumeft greaily tndened ■) 
antero-poxieriflr direction. It is noiewonhy that E. sif-Ufiuu '» I 
in the same beds as thoae coniatmng Hippari^H; and that th« efl 
E. cabnltus is apparently' the common species of the European Hi 
rene, althou^'h it is not improbable thai (he Asiatic E. ^hmt n^ 
iiccur in (ho saniv dcpoGils. In Soiithem India, wt>ere bo Uiioc* 
are found, the remainft of the existing Afiican E. ast'tms and Of «N 

r-„ V 

>'•€- ii)t--SI'«II «f III* HOTnlffMUfaMMuV RubtaA. 

undetermined form occur in the Pleistocene cave-deposits . _._ 
Finally, ii should be mentioned that some of the South AmcficaiiTe 
forms have been separated under the name oi Haf^okipptii. 

OKNt^AUJCV Of THE HoKSF-— Allusion has been incidentally 
in the preceding parajfiaphi to the fjenealofiy of the genus E^tm 
since this is one of the best instances of evolution among the 1 
Mammals yet worked otii it is adi-isablc that it should be ndntrd < 
more fiiily. The tup of the scries is the F.^uut itttitUus f,--- ■ i 
the dentition i* of the moKl specialised type, and which de-- 
to the topmost I'liocene; then we have the E. St/fiams and Ji pnm 
(■roup, in wTiich the molars become more like thoac of /flpparim 
tinilly the so-called Hippidium. The siruciur« of the foot l< iho 
tig. 1136, [> : the ^3t site of the plialan^'cals and th« metapodial 
functional digit being very noticeable. The eariiesi ftcnirrrnce of the 
is in the Pliocene of India. From the E. Sten-trnt emn^ to Ht^ 
it but a step, i\ie transitional «pr«ies beinj; N. 4iH//fi'/nmitm of t&i 
cene of the Siwallk Hills in which the laienl digits were apfs 
wanlini' 1 in the other ApccieN (tig. I3J6, C) the lateral dijpls , 
small SIM, and the middle one is relatively more slender than ra i 
Prp{okippnt connects E^uui by ibc structure of its mtlktrwilw 
the Miocene .■Iruit/^critrm ■ and in the latter the teeth hawe b 
brachydont, the ihiiU lower moUr has a »mall tliird lobe, iuid : 


the fool rfig. 1336, 1!) have become tarter and the tnidittedi^l 

; iracc* of the fifth metacarpal bciny retained in ihe American 

The next step k probably made by some form allieci 10 AkM' 

ror F<tikYiia!ofkui nt the Uijper Eocene ; and from such a type 

Btition r* easy 10 the Lower Eocene Hyrn(r>lherium and SycttMO- 

twhich till the species are verv small, i1ie dentitron i« nf the simple 

Bl type, with a lar(;e third fobc to the b*t lower Inie molar, and 

"fisol (fig. 1136. Ai has foiir complete digits, which are of sub- 

wie I while (here may be [Eciufipus) a rudim«ntal metacarpal of 

(poUcT, Kinally, the earliest stage of this series is formed by PnmaC' 

lot the Lowest or I'xiereo Kocene nf North America, in which there 

hit digiti to each foot, and of which the stnicture will be mor« 

reoticed under the head of the suborder Condylarthra. It should 

erred, however, that Professor Cope would introduce an inter- 


3 — 

Htff^rim (c), and Sfnui (d> 

Icdiate Maife between Sytttmodoti .md PhemKodus : and ilint (he came 
kdiOTTty also introduces the genus Palaothirium between IfyrMO- 
^Atm and AnMt&trium, although mgst writers regard itut ffenus as 
lofftlic line 

conncciiMi may be noticed the rcmailtable circumstance that 
line of evolution culmmntinx in the modem Horse a parallel 
i of jrtncrically identical or tloady allied forms occurs in the Ter- 
ries of 00th Europe and North America, from which it has been sug- 
mcA that in both Continents a parallel development of the same Kcncra 
b simultaneously taken place— /.r., that in boih rci;ion5 AHihilk<rium 
IS (ivCD rise 10 lli^tarion, and Ilipporiait or an alJicd type to E^hus. 
em, seting it 15 evident that in the ciLie of siierics nf a single genus 
C O'oluiion han taken place in HCparalc Hiiva. — that b to say. that the 
lasting Indian aperies of <.'.j«/j arc probably derived directly from the 
Bocenc fiirms of the same region, and the Brazilian species of thai 
EOiM have their predecessors of the cave-epoch of that country, — there 
jipears no logical reason for refusing to admit an analogous parallel 
rotuiion in the case of genera, and there is accordingly a i^unsidcrablc 
rohability thai the Iiy|xnhcsii in queilinn may be a true one. I'ro- 
Bsor Cope considcpt (hat in one country Protohtppuif and in the other 
M, was ihu immediate ancestor of E^uiti. 




Family Riiinocp.rotid.«- — With this family we 
considtraiion of another branch piohalily derived trooi tl 
live I^phiodont stoclc, which attainiti great dcvdopment 
tiary times, and is still represented in Asia and Africa b 
five wcll-dcfincJ species. It is not eaay to diMinsuish || 
from tiie Lofhiedofaida,, as represented bjr /fynuhiut (' 

■■» c 


Fk. i*J}.—Lrft hair of itie paUnl nirfiKic o( llw cruian, tmi M IMM 
of MttmmymtJam /JjM,/nmi : rrom th« WhiM Hinr Mlofan* of K«d> 

Tulutal tiie. A'a NwuI i /^r, l-nnuJ 1 /'it. PaiKu! ; Sr. 5uc>aaiEitanl ; : 

eIbmIcI and pMnyinrun ii! procotct : 1, i. ), Almti or inciun; >:, Ahnol^ll^ 
PiOHitmi ■• >> ji Tf ite o)oW>. (ATlcr SmIi Mill Otbcim.) 

Schlosscr includes in the RhinoerrotiJir), tiut llie uppa I 
(6g- 1 »39) generally have a very thick outer wall, whic 
produced in advance of the firil ridye ; their trattsverK 
but slightly hent, and are intimately connected with the i 
ihe upper premolars are usuidly nearly or quite » coni| 
true molars; the lower cheek-teeth are more or less • 



And in all ihe forms in which that tooth is known 
ijrd lobe to the last lower true molar. The height of 
-leeth vari« consideiablj', their crowns being tallest in 
jfenwMr. One or more dcnml horns may be attached to 
ito-naul region ; 2nd when twu of these appendages are 
Bicy may l>c either placed one behind another in the middle 
b a pair on cither side of this line:. The digits of the [je» 
jTcntly always three, but there may he either three or four 
|Bnu5. One of the mo»t generalised forms is Hyracodon, 
Lower Miocene of Nebraska, in whieh the dental formula 

n. -, Pm. -, Af. -. ThcrewereappaTenllyonlythreedigils; 

and limlx^ were slender and Horse-like; and there was no 

a nasal Iwin. This gcnui> was in all prolMhtliiy n dc- 

of llie Lophiodont JJynukyus, but docs not appear to 

in the prDgeiiitor of the true Rhinoceroses. In auiiie re- 

jU more generalised is the genus Amynadon {OrtfuKynoilnn), 

I Middle and Upper Eocene of Nortti America, in which 

(al formula was tlie s:ime as in Hyra(odim. Thi; lower 

^crc nearly upright ; there was a short diastcmn ; the prc- 

■ere unlike the true molars ; and it is Iwlicvcd that the 

id four digits. Allied to this genus is Atefamynodcn, from 

jeae of the United States in which the skull (fig. 123;) 

ng sagittal crest, the premolars arc reduced to -, the lower 

lavc become somewhat proclivous, and ihe upper premolars 
I more like the tnie mobrs. I'hese t«'0 genera are regarded 

! of the American 

plogists as indicnt- 

istinct family — the 

PMtida — and aa> 

iked upon as the 

B of the true Khi- 

^ In the Old 

(here is, however. 

[US Cadunotherium 

[nercy Phosphorites, 

pnay possibly lay 

I this position, al- 

lit may indicate a 

tranch allied to the 

lltia. Unfortunate- 
detached teeth arc at present known, so that the dental 

cannot be detennined. The upper true molars (fig. U38) 

Fifl. itjl^ -^ A left upucr iruc pioUi of CAiiur' 
rMMTiim aifiinfiui i imta tbi tJppcr Eecwt 

of PfMtM. 



are Rhinoccroiic in structure, but are extremely narro* 
verse direction, and the ridges of ihc lower chcck-tnih 
fecily cresccntnid. Apparently nearly related to the pi 
HoatahdonkilkcnHmy from Tertiary strata of unknown a 
gofiin ; the dental formuln is the lypical one, and there bn 
but the skeleton is unknunti. VX'c now come to the ot 
of iho^e nnimnts which we may tcnn [rue Rhinoceroses-' 
which rcry dtvcnc rvcm as to the limits of generic tci 
valent Among zoologistB and paleontologists. By sonw 
five existing spccicii arc referred to at teast three ditt 
and if this view be adopted, it nitl be necessary to n 
number of genera for the extinct forms ; the Lnglish s 
ever, now generally include all ihc living species* in one 
from tht» point of view there teems no good reisotn foi 
separating any uf the extinct specie^ which form a se 
mately connected that it would be very difficult to dc 
{^eneia into which they are divided by the American scho 
then, the lerrn Rhincifr<fs in its widtfst sense, the variat 

number of teeth may be expressed by the formula J. 


4 3 

Pm, ■ , M.- ; the aUscn<:e of upiKr canines is a distinc 

** 3 

the upp*r true molars (fig. 1*39) haw Iheir crowns rdi 

their transverse ridges well dei'clopcd, the hindtx lobe 

^^ of Knitonil. Unc-htlT auunJ mr. ^^ 

tooth partially aborted, and frequently a more or l^ff 
tress ai their antcro-cxtcmal angle. The teeth rcprea 
1 »j9 are the most generalised type; and it is evident thai 

■ III thi&ARil nihcr inttinca the number of jcnnk <livukxiswhl 
(lii|iosed to iiJoui it solely n mniier <tf conveiuencc. From (be wi 
view the muLtlplIcaitua of eennic icmti, which w obi )i>mw1ci1k« i 
Ijecmnc lew and leu miKcptible of cx/ict dcfibitian, trn>li to drowv 
■ MB of tumea, whicH (onn ■ gicat burden to ibc m«notf , Mk l ifcl 
•Irajr the vei7 object of dudMalion. 



awm of each molar wouUJ carry Iwu isolated foseetics 

I by enamel (fig. 1344). Tlie worn crawn-smface is trans- 

[ed i and there i.i a pToccss ptojc:ctiiig fmn) ilic hinder 

he middle valley leniifd tlie ciochet, which U ;ihsent in 

es. llic hinder prcmulur:^ arc as complex as the tru« 

id the crowns of ihe chtek-Ieelh. though varj-tng in 

jievcT very tol), and their vallejTi are alwnys open. In 

wek-l«vth the ridges form compleiL' L-rescents, with their 

liKClcd inwardly (ilg. 1240). 

canines are always proclivous. 

m and akull are very iiia&ai^c, 

being idcaI maiVcd in the more: 

species. This genus may he 

> several groupie, of n-hich the 

■ is (he mcMt generalised. In 

there is usually no horn, and 

jones (fig. I J4 1 ) are conse- 

nll ; culting-tceih arc always 

lODgh thciv is some variation 

Via. >na.— T Iw third Mi la*w 
litic iiiol-nr of JtAfvtvtrvg mtgur* 
tiHmt: ?lali.UK>fi(. Tuu-lhioli 
eiuucdI tiim. 

ibet, which may be expressed by the formula / 


?. indtivui (which is the Ij-jic of the so-called Ai^ra- 

Sre are four dibits in the inanu» ; but in many of the 
rican forms (which on this account are separatee) hy 

-Cnaium </ XAiivHrTV imliini; hum ihc Pliucene a( <M-muny. 
Oiu.f*VTnih nuiunl lire. (Aflu Kivup.) 

Vpe under the name Aphehps^ fig. 1241^ the number 
} reduced to three; and these forms were lhu& similar 
ate examples of the existing R. wndiiUus, in which the 
aL In Europe this ^oup ranges from the 1 .owcr Mio- 

3 H 



cene to ihc Loner I'lioccnc ; Jt also occurs in ihc Upper Mi 
and Pliocene of India, and id itii; Upper Mio^-cne for 21 
uf Nonh America. In ihc Ifuera/Acrtne group {Db 



J:* "^i 

FJK- IW.—Skt\la( Xliimttm mcThMm; frodi ihe Upficr itia 

■ orCobafc 

Mnr^h) there was a. transrvrscty-pkced pair of sniall nasil botoi 

the formula or Ihe cheek-teeth being / , C. - z it is rcpreeaH 

in the Ixmer Miocene of Europe bjr X. miiuifus, and bywHh 
specic-s in North America. At this stage of erolution the go 
disappeared from the latter country, 

The RkiKiKtrittinf grouji is charvicteriscd by the prcKlxv of 
single well -developed nasal horn,^ and of eulting-tecth in boihilo 
Ii is represented at itie present day by the Asiatic R. umjaiaa a 
A", itnie^rnis (fig. i 243), the upper true molars of the former W 
or the type of those of ft. mrgarhinui (fig. r 239), while those rft 
Inlter ore of the more specialised type of R. aMfi^MUntii (fig. m 
The ancestor of R, sondaUus is probably lo be found in R. iivaiim 
of [he Plitit'i'ne of India : while Ji. fialmndimt appears to be I 
species from which R. ttnuormt has spning, The CrmAmt 
group, represented by the existing .\siatic R. sumaiw>tnm^ isA^ 
European Ixiwer Pliocene R. SeMeUrmackeri, £\f[eti from the p 
ceding by having two homs, placed one l>ehind Ihe other in I 
median line, but still retains cuiting-leeih in buih jaws ; ihc ap| 

' The horn nr the RhinoccfMn, ir should be obMrvcil, comut* aeidf < 
bundle of cloKrljr agglomcmiol bniilct, soil hu no boav UttduBrnt M 

ftkull. r 



boih species being of the type of fig. 1239. This group 
lohahly be separated from the next, with vrhich it is con- 
jy H. Persia, of the i'liocene of Mar.agha in Persia, which 
caiiino, allhough apparently allied to if. platyrhinns. 
specialised, or Aftiodint. );roup is repreacnicd <tl the 






I lac^—Woni sfi iipfwi JcoUtiao of ffJMMtvfw m*kentii : Iwlii. Much itdiicnJ. 
(Aflar Cttvur.) 

^b>' the AfTi«:an f{, simui and H. bicornis, in which there 
_ 1 horns, Iwil no cultin^-tecth in either jaw. Of species 
■ motors of the simpler type of fig. 1 a,^g, wc may mention 
^^ ta£4»f, of the Lower I'liuoene of (Jreece and the isle of 
whkh is closely alhed to the African R. bicornii ; /i. 
(fig. 1144). of the Upper Pliocene of Kurope (in whirh 


|nn> iri'lhs cnnium of KlirmBi^ri itraieft, itlih ilie Inlh niuih vain; 
tJplKf PKocsn*, llaly. Ons^cwnih tiBtiinl ui«. 

are of a brach>'dont stnictuie) ; /?. deemntnsis and R, 

tit, of the Plcistotxnc of Soullicm India; and H. mcga- 

Sg. 1339) and R. ifplnrhinus, of the European Pleistocene. 

attcr there is an ossification of the nas.1l septum. The 

of this group hive their tipper molars (fig. 1345) of 



0. more complex Type ; there bcti^ an absence of a bvinaii 
amcTo-i'Jitcmal anxle. and ihe folds of the crown to amnjd 
when more worn than in the fibred specimen thice »M 
enamel would be formed on their crowns. These teeih m 
charaacriscd by their plane of wcnr being perfectly bona«4 
by their relatively tall crowns. Arv early meml>er of ih» npei 
(ffalyrkims, of the Pliocene of Northern Indb ; from wbicfcui 



ii«.~'n>t weonri tifbi upper inienu Jar i)rff4/«ir«M*««(fra 

it is highly probable that both the existing African Jt. 
the Pleistocene Ji. antiyvitath, of Nonhern Aua and Et 
been derived. 

The tatter ^ecie», of which the »lcull is represented in 61;. 
upper In fij,," '^45! '* soinciimcs kiMiwn as ihc WtjotK^R 
ninrc tt was cohered wilh a thick cait of woiilly hiur. Thr ill 
devoid of the folds which charaacriM: the U[);c Indian ^i" 
front horn was of very large siic- As in some of ihe Pic. ^ 
(he scpium of the nnrc^ ivai completely ossi^cd (&^. iiabi. 1 aa 
is essentially » noMhcm fi>nn, and haH nearly the 5ame diMi 
as the Mammoili, aliti<iui;l) it liirei ntit ap| ici haiT rrossed I 
Siraii inin America. In lime ihi« Khinocctus makci it* fint appi 
in the Pleistocene Drick-canhs of the Thames valley, AaA it very a 
in European cave -deposits, and in the tuH<iras of Siberia. Ci 
carcasses, still cnvricJ wilh the dried desh. skin, and hair, havea* 
qucndy been fnurul w.i^hed oui from the frotrn alluvial deposit** 
tuniinu un the tiank> ai the Yenciti and Lci 



ift animal tMunly censiKcd of th« Waves and twigt of juniper 
lifenntft plants. 

rtpr«€ntativc of this family is the ^gartic Efatmo- 
raKcros) of the Pleistocene of Siberia, in which the 

aU of the adull is /. -, C. -. Pm, -, M. ^. The slruc- 

tkuU and limbs '\% csHcnliatly Rhinorcrotic : and in ihc 

laiial »eptURi vm conipIctL'l)' us^iflL-d. and the frontal^ 

; bony protubenince fur the support of a lai^c horn 

Ig 10 ihe second one of Rhittoctrni aniiquitatii. The 

considerably from those of any species of Hhitwctroi, 

\ ^IfMCt 4f >)>« «klllJ ofb ffjunff individual tiC Jli/wMfrvt JTMfiyuiMfa; 

Irma ibc FUiitactiM of Sibcni- Kcduccd. 

aracterised by their very tail crowns, plicated enamel, 
plane of wear. Their structure is, howcA-er, merely an 
dificatian of the Rhinorcrotic ij^pc, to which ihe nearest 
nong later forms is made by A*, anri^wi/tifis. There is, 
ihesc tccih a marked resemblance w Ihosc of Cudurcih 
3 Homaiodontofhtrium, and it is not improbable that 
vai presents the last rcprescnialive of a ^tock de^ccI1ded 
irmef genus which has remained altogether apart from 

Luni>oniEKilD.c. — With the LamhdothMida we enter 
insidcration of the first of three cxtinrt families in which 
eeth have remarkably short (brachydont) crowns, and 
. certain extent from the more t)'pical Lnphodont form, 
tnie molars (fig. 1 J47) may he described as consisting 
imns, of which the two hindmost are frequently eon- 
tn obli4]ue transverse ridge ; while there may also be a 
iplete ADterioi ridge. When ibese teeth are worn two 



niolu of CAMlktiitrirm iimtmu; 6oai 
iht PUoecn* of China. 

V-«hape<J surfaces of dentine appear on ibe crowa "nm 
tooth may be derived from tlic Lophodonl by the xsxn 
complete abortion of the middle portions of the nanswn 
The upjTtT premolars are simpler than the inie molantii 
hm a single iniK-r toliimn ; while ihe lower d»cek-lc«th art 
loid. the last tiuc molar usually not hav-irtg a conipktc thi 

There un^s ;Ll«-ays a dvutcs 
dcRUl scries, and the fknl 
void of bony proiubmtK 
the preset! I North Araewa 
the femur has a tliird tmchi 
the feet are of the nomul 
dactyJate iy|ic, ihc maiius I 
vtded vrith four, and the 
three digits. This (amilr 
sented by Falao^fs an 
lktrii»m, from the Upper I: 
which tliere are four [weni 
the last lower true molar h 
lobe ; the eanines lieing 
resembling those of the < 
iMmbdotkerium is anothei 
bter age; while in the White-river Miocene of Candull 
ffapluavion, wilh only two pairs of lower inctsorK. 

I-'amilv CiuuconiKRiiD*. — The Kcond family, 
tAeriiiia:, is found in l>oth Uie Old and tlie New W' 
scnts such a remarkable abnonnnhty in the structure 
as to render it for the future quite unsafe lo predict 
of an animal from a single bone, and to make in 

maxim tx tutgye 
the femur the 
chanter lus been 
in the feet, while 
tnal bones retain 
t'erissodacty lot e 
the phalangeals 
modified to 
of Edentates. 
ric....^-A<.wt<xud<ii««iup«Nof.>«>wi phalangeal (^ 

EhulaiuaJ c/ CAaiictlitrimm tiratimu; frwB (b* \f\a a StPOnslv 

tal trochlea for 
tion of the huge claw forming the terminal joint. 
geals have been described under the names of Midi 
and AncyMherium, and were until quite recently* whea 
found by Dr Filhol in association with the skull 


skeleton of Chalifothirium, thought to belong lo liuge 
lies. The laticr writer has indeed proposed to regard this 
as a Teritable Edentate, but The rescmljUnce of its denli- 
In that of Paitrasyefs, coupled with the es.scntially Peiisso- 
Me chjirarlers of the rest of the skeleton — notably the 
loaelous cervical veaebr* — prevents the acceptation of 
and compels tu to regard thiii strange animal a<i a 
nodilicd and aberrant Ungulate.' In the type genus 
•tkerium, with which Matrofhtrium ■« identical, there were 
in the number of the cutting-teeth analogous to those 
ing in Xhinoaros, which may be exprt'ssed by the formiih 

V, a ^^I^, Pm. ^, M. -1 The tyi>c spcdt-s, which should 
3) < 3 3 

own as C. gigantmai, occurs typically in the Lower Pliocene 
peliheim in Hessen-Dannsladt, and aLso in the Middle Mio- 
rf Saman in Gcrs; the claws were first described by Cuvicr 
the name of Pangolin gigaula^ue, and were Kubnequently 
the type of llic gcnu-s Afua-atktrium. Another species, C. 
mi, occurs in ihc Upper Eocene I'hosphoiitet. of France, to 
probobU' belong some Urge claws described as those of an 
lU'. The genus also occurs in the Pliocene of China and 
the species from the latter area having been referred by 
to a distinct genus Nestiwitkerium^ on account of the absence 
anterior iceth. The phalangeal from the I'Uoccnc of Sind 
nted in fig. 1348 doiibllesv belongs to a small individual 
species, itlthough first deKcribed as Mants, and subsequently 
crothtriHtn. It has likewise been lately recorded from the 
-ii*-er Miocene of Canada and the Loup-i'ork bed* of 
'ITie last lower mular of Chahcolhtrium has no third 
From the Lower Pliocene of Attica the genus /^j>t,>i{afi, 
>ed on the evidence of a lower jaw with a Chal icoiheroid 
on, but with a third lobe to the latii molar, is probably iden- 
fith Attcylothfrium, founded upon the evidence of claws from 
1B€ beds, which are of the snme general type as those of the 
d Afacrm/urium. Anrylolhrnum also occurs in the Lower 
me of the isle of Samos. I^fti^fon has been provisionally 
■d by Dr Schlosaer lo the next family. AforDfus, from the 
Fork of Kanus, and Mornlhei'ium, from the Miocene of the 
d States, which were described by Professor Marsh as liden- 
are probably closely allied to, if not identicjil with, either 
^htrium or An£ylotheniim. Finally, the imperfectly known 
ydiattttnaMherium, from the Kocene of Hungary, is probably 

B(c<Mr Cop« hu reotntir propMod to make CAaiktl^rinm ihv type uf n 
onl^r wuui tbe iMme of Ancylopoda. 

Iheriida, that I>r Schlosscr has propo&ctl to unite ' 
pfcsvnt rami))- at tva&t the founh upper premc 
columns, and is thus a& complex, as the true nu 
forms of which the limbs are known there are foar i 
posterior digits. Ihe third trochanter of the 
I'aired bony pmtuheninces may lie present in the 
of the skull 111 Dipiacvdffit, of itie Eocene of Nc 
the fourth upper premolar is as coinplex u tfal 

canines arc as large as in Paiaotyops; the int 

and ihc skull has no bony protuberances. 

The tyije genuN Tttamttkerium, wliieh Messrs 
consider should include MencjHs (picocoJ|Mcd),J 
tfifherium, Diconodim, and proljably Syndxfrodotiy' 

Fit. IM9-— I<a(l birni upMi af the cnnium of TUttmMlL, 

Miocene of North .\01crica, and comprises ani» 
and it is provable that the forms recently de 
Marsh under the names of Alh'ps, Srvnt&pj (fig. l) 

1 premolars are alike, the incisors are Kmalt, and in 
tha full dentition there is no diastema : while in the 
he canines are small. The cranium (fig. 1149) is 

Family Macrauchenhd.-e. — Here may be p 
able family, which presents extremely generaliset 
vertebrae and limb-bones, such as are unknown 
bers of the suborder, on which account some wri 
not to be included in the Perissodactyla. In t) 
ehenia from the Pleistocene of South Ameria 

/ ?, C. \, Pm. 1, j»/: 3 ; the cheek-teeth are 

3 ' 4 3 

structure, the upper molars showing two externa 
surfaces and two transverse ridges ; while there i 
diastema in the lower jaw. The cervical vertel 
of the Camtlida in the position of their arterial 
articulates with the calcaneum (as in the Artiot 
are three digits in each foot, of which the lateral 
size. The incisors have a deep coronal infolding 
in the Equidte; and Dr Hermann Burmeister thir 
was produced into a short proboscis. The type 
chonicha; an allied form from the infra-Fampean 
named by Bravard Palaotkeriutn paranense, is n 
meister to this genus, but has been made the 
genus by Dr Ameghino, under the name of Seoi 
vardi. M. minula from the same deposits is i 
writer the type of Oxydontotherium ; and the n; 
applied to yet another form from the same area, 
noticed Tkeosodon from the above-mentioned i 
placed by Dr Ameghino in this family, although 
Jiomalodontotherium, which is also placed here b; 

SuROHDP-R 3- ToxoDONTlA. — This group incl 
verj- aberrant and generalised Ungulates from the 
America, which present affinities to the Perissoda 
and Kodontia, and consequently render it almost 



fH>wn. In TMfidvH, ol which llie lypc siwcics is of large 
I is round in the Pleistocene of Argentina, ihv dental for- 
/. ?. C. -. Pm. *, .If. ^. All the tctrth (fig. 1251) grow 
; 3 ' 3 3 

|jcrsis[cnt pwlps ; ihe lower canines are very minute, the inci- 
irge, and the crowns of both the Utter and of the clicck-tceth 
ly currcd. The Mnicture of the latter is a simplilication of 
in obtaining in A'esoJoH. The femur has no ihird trorhanter, 
tola articulates with the calcancum, and the rmnium approxi- 
in some respects lo that of the SuiJit. In the typical T. 
v't the outermost upper inciMr i& the larger of the two, the 

fc-OnI uw fa c M of ill* nch) apM'' (aWikI \Mrr(*yUr>iitlan of ruaJM RttrmuitNri i 
mm PMwocMt of tbe Aii*iitinc Republic. Much nUuced. e. The lower aniuc. 


condition obtaining in T. Burmaiteri {^g. i!5i). From the 

unpeon depo&its l)r Ameghino has recently desoihed vari- 

nains of allied forms under ihc names of ToxodattMhermm., 

W^-tothtHum, and Di/&f>odi*n. A mandible from the Teniary 

tltc Hrmiosa, in vVrgcntina, is characterised by the triaiigvilar 

f thr third incisof. and has accordingly been n;>nied TrigoiiiiH 

ntcghino, Dr Moreno ttiaics, however, that this mandible 

to the .some animal as the leeth descrihed as Toxodontothfr- 

Haphd^ntathcrium ; and he would adopt for their owner 

Trig^don as the one which was alone well defined.' The 

dible is peculiar in havin^j only a single median incisor, 

on the line of symphysis, but this is probably an individual 

Dahty. Apparently more nearly allied to Nesodon (which IJr 

lino makes the type of a distinct family) are Colpi'don and 

^<m ,■ while other allied remains described by the same writer 

fie Tertiarics of .\rgcmina under the names of /ntenifArrium. 

^UfMatus, and Ttmh^thtrium, arc referred lo two distinct 

LT TvPOTHERill>je. — Perhaps still more remarkable is Typo- 
« U reallj pmxcupicil by llie e*ilin TrigfKoJrm uid TtumimAm. 



tkerium {Metolhfriunt) from the infra-Pampcan beds of 

in which the denlilion is /. -, C. -, Pm. -, M. -. The 

grow from persisl«nt pulps ; and the structure of the chcek'l 
recalls that of Tcxoiivn. The iikuU (fig. i3S>) U of ui U 
late typ«; there is a ihird irochanler 10 the femur; anddi^ 

I 4 


Oiif -luir iuiumI tin. 

(which are iiiikno"ini in any other Un^late) are present 
peculiar ^srwa presents duiacterft connecting the ToxeAmtiJa 
the Rodente. Allied forms from the infra-Pampean deposits of 
iame region have been dcscrilMrd as Profypotkerhim, 
and Tcmodus ; while Pachymcus from the lalcr Tertiary of Mon* 
Hermosa is a raiirh smaller form with three premolanL 

SuBORDF-R 4. CoNHYLARTHRA. — This gToup ootnptises a Bumbo 
of very generalised Uiigulaiea ninstly from the Eocefie of N**'* 
America, which are grouped by Profe>:sor Cope, on acratmt of ik( 
structure of their feet, with the Hymroidea in a divi&ton tascd 
TaxcDpoda.* Both the present group and the Hyrarotdea irt 
charactetised by the scaphuid of the car^ius being supported brlfx 
trapezoid and noi by the inaL-nuin, which carries ihe lunar ; «t"If 
in the tanus ihc cuboid articulate:! proximalty with the tala««» 

^ J'roreuar Cope would bIm indiMi« tlie Prinialct among ihc TMMfiii 



There is but sliglit iniitual interlocking of the carpal and 
nes ; the stnicture of ili«.e joinik tteing dimpler thiin in the 
:bordcrB, and lescmbling ihoiv of the Unguiculalc orders. 
Condyliinhra the denial formula is nearly alway>i the typic;il 
be cbcek-tccth arc brachydont, and usually bunodont, flUhou(?h 
Df>htxlom. The premolars arc simpler than the true molars, 
may be tritubciculor Itkc those of many Carnivora ; the can- 
d incisors frequently also recall thoM- of that order. The 
Bs is peculiar among Ungulates in having an entepicondylar 
n ; the feinur has a third trochanter ; the a-siragalus, as in the 
Ora, prtsenis a uniformly convex distal articular <iurfacc ; and 
la no anktibr facet for the fibula tilhcT on ihi:> hone or on 
llcanetim. The feet ucunlly h:ive live dibits, with siharply 
d terminal phalangeals : and the radium and ulna arc distinct. 
luburder may be regarded as containing ihe ancestral types 
■rhich the Aniodactyk and Pcrissodaclyla have sprung;, It 
presents such remarkable signs uf atlinity (especially in the 
ire of the tcclh, the form of the astragalus, and ihe presence 
foramen in the humerus) with tlie Coniivora, that it seems 
probable that we may look u|K>n the CondyLarthra an a hide 
I frotn the original ancestral stock of the Clamivora, which is 
Dcaxly represented hy the nunc primitive Creodonts. 

iLv PERimxHrn*. — In this the most generalised family the 
ion Ubunodont; the digits arc five on each foot; the astra- 


: from Ihc FUoco Eaxim of N(V Htiico. Two-thinli attursl uzc. isr>ri..i|. f 

"ias no trochlea ; and the premolars are very simple. In the 
enus J'iripiyckitf, from the I>owcst, or Pucrco, Eocene of New 

», tlie dental formula is /. l?-5^, C. -. Pm. ^, M. -, and the 

and lower caiiinea are small. The typical f, rhab^dfm 



<fig. 1*53) is characleriscd by its %x'itically grooved 
Other genen from the Nonh American Eocene are /frjKdn, 
fifnas, AniiofKhus, Htmitklmis, /fafifaceityt, nnd &foJi>H ; d 
\he fourth may be not sepcuable frutn tlie third From iha I 
|)r Schlos:«er rcftArds the bunodont Arttodaru-la as dcrmiL 
thinks a direct rtrljt ion ship can be traced from /Vn//yrAwf la , 
ikfan, and thus to ihc other Char^potamiia. Tbc 
short, and much expanded distally. 

KAMtLv J*Ht!*\coi>ONTiD*. — In thw family the brain {fi|.li 
is characterised by the extremely small siu of the heiniv 

\^- ) 

F^ ■>]«.— InfErio I, >.ui«-ior. ud left Uhnl vpeett of Ifcc 1 

(rani IDE Luimf Edmiic af Nortk Anula. (Alia C«ve.) 

which arc only one-fourth longer than the cerclK.-nmn. aid Itet 
indicate a very low type of organisadon. The family i* reaiili 
distinguished from the preceding by having a proitmal trochlcit>) 
the a»itraga1u5. by the longer neck, and less simple premobn, ■'^^ 
are, however, diifercni from thos* of the following famity. 
type genu» PhtMncodus (fig. 1 155) includes several species framl 
Puerco and W'asatch I-Jacene of North America, VArying fwn 
size of a smalt terrier to that of a leopard. The dcnta] Mrrtm-'oK-^ 
prises the full typical niimlwr ; and although the croHTis of the ufpJ 
true molars are of a bunodont struciiire. yet they couW be roidiK 
roodilied into the lophodoni type of //yraealJurimm, and vr mui 
probably regard the latter as a direct descendant of ihc prtxiJ 
genus, with perhaps the inlervention of SytUmadam. Pro(t:^s« 
Kiitimeycr has dcKcrilicd some upper molars from the VfV^ 
liocciie of Switzerland which he refers to PAfMOtvJiu, aUbooc^ 
[hey arc much more of a lophodont type than in the AfBcrio' 

Profesior Cope remarks of one of Ihc species of PA^ManyAs tlui "-^ 
»i» of the animal is about Iliai of a Bull-dog, but the head u tmal'^i 


and the neck rather shorter and not nearly so robust The lim 
about the same proportions to the body as those of a Bull-dof^ 
anterior ones are shorter The proportions of the parts of the lio 
of the fore and hind limbs to eacti other, excepting the feel, x 
as in the Collared Peccary. . . . We can thus imagine the Pii 
yor/mam as an animal of the comparati\'ely slender build of t 
dog, with a head and neck proportioned more as in the Racoon,] 
the rump more elevated than the withers, as in the Peccary, 
resembled those of a Tapir or Rhinoceros, but had an [additio 
of short toes on each foot, which did not reach the ground. To 
a tail much like a Cat's in proportions, and the picture is comi^ 
diet the animal was omnivorous, the proportions of animal foe 
smaller than the Hogs, for instance, use. The food is more likel 
resembled that of the Primates. What means of defence this spi 
is not easily surmised, as the canine teeth and hoofs are not lar^ 
The species represented in the accompanying woodcut was 
dimensions ; Professor Cope stating that it was intermediate ii 
twcen a Sheep and a Tapir. Comparing it to an animal with a 
long tail, we might perhaps take a Leopard as a fair representati 
remarkable length of the tail at once shows a wide diffeicnn 
existing Ungulates. Professor Cope, from the structure of the 
the nasal region, suj^ests that the bead may have had a ihort | 

Other genera from the American Eocene are J'iro/agffm 
odon, and DiacodexU. The former occurs in the Puerco, and 
has but one outer tubercle to the fourth premolar, in plac 
two of Phenacodus. Professor Riitimeyer refers to this g« 
from the Upper Eocene of Switzerland. 

Family MENiscoxHERiiDit. — This family is taken to inc 
genera characterised by their lophodont dentition, which 
dently more specialised than the preceding types. By Dr ; 
they are regarded as allied to the Chalicotheriiday and 1 
perhaps indications of affinity between the European genui 
Hyracoidea. The humerus is longer and less expanded th; 
Periptyckidiz, but the number of the digits is unknown. 
no marked diastema in the dental series. The typical gem 
cotherium is from the Wasatch Eocene of New Mexico, ani 
acterised by its small incisors, and the presence of two outer 
the last upper premolar. Teeth from the Swiss Eocene h 
referred to this genus. In Hyracodontothtrium, from th 
Eocene of France, the upper incisors are large and cur\-ed. 
being especially enlarged, and closely resembling the corre 
tooth of Hyrax ; the canine is small, and resembles the 
cisor ; while the fourth premolar has but a single outei 
tubercle. Two species are known by the skull. 

SunoRDER 5. Hyracoidea. — .A3 mentioned under the 
the Condylarihra, the structure of the carpus and tarst 
Hyracoidea is the same as in that suborder, but the termi 
angeals are truncated, and there is an interlocking articul 



I ihc fibuU and JUtnigatus. This suijorder is represented solely 

Euntly HyradtLf, oonnining the twn existing genera Hyrax 

1156) and DtHdrohyrax ; both of which are confined to Africa 

ami are unknown in a fossil condition. The dental 

of ihc adult is /. 

C. ° Pm. *, M. ^ 
o 4 3 

tlicre are four 

n ; ihe incisors grow from persislenl pulps, and tlie pattern 
eheck-lc«th is of a Rhinoccrotic type. The forefeet have 

nt. 199S1,— l-*Atai«nlai9Mtef ibci)inll«f /OvKxn^Mfic. KeJuoed. 

. and tbe hind ones four dibits : and the terminal phnlangeal» 

in the inner digit of the pes) have rounded liouf-Iike nails. 

feoracoid process of the scapula is well developed ; there i< no 

jndylar foramen to the humerus : the femur possesses a small 

trochanter : and the tibia and Abula are distinct. 

IJBORi'KK 6. AwBLVioDA. — [n chis suborder, which comprises 

Vngulatej of great bulk from the Eocene of Europe and 

.\merk-a. the carpus (tig. 1157, f) 13 chnr.irtcri.scd by the 

>id being t^upportcd by the trapezoid and not by the magnum, 

the lailer and the unciform support tiic lunar ; in the liirsus 

iboid .inicubtw with both the calcineum and astragalus. The 

is therefore of a more primitive type llian the tarsu?. Both 

\\ and the tarsal bones interlock to a slight e