BULLETIN 
OF 
THE 
TEXAS 
MEMORIAL 
MUSEUM 


15 


DECEMBER 
1969 


Pliocene 
Carnivores 
of 
the 
Coffee 
Ranch 
(Type 
Hemphill) 
Local 
Fauna 


WALTER 
W. 
DALQUEST 


TEXAS 
MEMORIAL 
MUSEUM/w. 
W. 
NEWCOMB, 
JR., 
DIRECTOR 
24th&Trinity,Austin, 
Texas/ThcUniversityofTexas 



Contents 


Abstract 
1 
Introduction 
1 
Acknowledgments 
1 
Geology, 
Occurrence 
and 
Taphonomy 
2 
Taxonomic 
Synopsis 
4 
Accounts 
of 
species 
5 
Vulpes 
cf. 
V. 
shermanensis 
5 
Osteoborus 
cyonoides 
5 
Canid, 
undetermined 
8 
Indarctos 
oregonensis 
9 
Pliotaxidea 
cf. 
P. 
nevadensis 
10 
Pseudaelurus 
hibbardi 
11 
Machairodus 
catocopis 
13 
Speculation 
as 
to 
faunal 
relationships 
of 
the 
carnivores 
26 
Literature 
cited 
43 


List 
of 
Illustrations 


FIGURES 
PAGE 
FIGURES 
PAGE 


1. 
GenerallocationofthemiddlePliocene 
8. 
Rightupperandleftlowerjawsofyoung 
(Hemphillian) 
Coffee 
Ranch 
vertebrate 
adult 
Machairodus 
catocopis, 
in 
lateral 
view 
15 
quarry, 
Hemphill 
County, 
Texas 
2 
9. 
Vertebral 
column 
of 
Machairodus 


2. 
Cranium 
of 
an 
undetermined 
canid 
in 
catocopis 
18 
dorsal 
view 
8 
10. 
Vertebrae 
of 
Machairodus 
catocopis 


3. 
Lower 
jaw 
of 
Indarctos 
oregonensis 
9 
viewed 
from 
above 
20 


4. 
Lower 
carnassial 
of 
Indarctos 
oregonensis 
10 
11. 
Limb 
bones 
of 
Machairodus 
catocopis 
23 


PLATE

5. 
Plesiogulo 
marshalli, 
upperMl 
in 


11 
I 
Part 
of 
the 
skeleton 
of 
Machairodus 
catocopis

occlusal 
view 


6. 
LowerjawofholotypeofPseudaelurus 
inthefieldblock 
Frontispiece 
hibbardi 
with 
P3-M1 
in 
labial 
view 
11 


7. 
Restoration 
of 
skull 
of 
Machairodus 
TABLES 
catacopis 
13 
Tables 
of 
measurements 
29 



lumbar,


Noteblock.
field 


the 


in

41261—8)


ITMM
cotocopis


Machairodus


of

skeleton


the 


of

Part

I. 
Plate 


and

cervical


articulated


right,upper

at

ulna 
and

radiusbelow,

vertebrae


caudal
proximal


and

sacral, 


this

when

removed


been 
had

bones


the 


of

Manyleft.

upper


at

pelvisand

center,


in

vertebrae


thoracic


matrix. 


the 


in

buried


still 


were
others


and

made,

was

photo


Pliocene 
Carnivores 
of 
the 
Coffee 
Ranch 


(Type 
Hemphill) 
Local 
Fauna 


WALTER 
W. 
DALQUEST* 


Abstract 


New 
material 
collected 
in 
the 
Coffee 
Ranch 
quarry, 
Hemphill 
County, 
Texas, 
represents 
eightspecies 
of 
Carnivora. 
Included 
in 
the 
fauna 
is 
an 
apparently 
new 
species 
of 
Pseudaelurus. 
Abundant 
material 
makes 
possible 
a 
detailed 
description 
of 
the 
bone-eating 
dog, 
Osteohorus 
cyonoides. 
The 
saber-
toothed 
cat, 
Machairodus 
catocopis, 
is 
described 
from 
a 
largely 
complete 
skeleton. 
Specimens 
include 
teeth, 
previously 
unknown, 
of 
the 
rare 
wolver


ine, 
Plesiogulo 
marshalli 
and 
a 
very 
large 
bear, 
Indarctos 
oregonensis. 


Introduction 


The 
Coffee 
Ranch 
quarry 
in 
Hemphill 
County,
Texas 
has 
furnished 
important 
fossils 
of 
extinct 
mammals 
of 
middle 
Pliocene 
age. 
This 
local 
fauna, 
described 
in 
1930, 
1931 
and 
1932,was 
so 
distinctive 
thatsimilarkindsofmammalianfaunasfrom 
elsewhere 
in 
North 
America 
are 
referred 
to 
the 
Hemphillian 
fauna 
(Wood, 
et 
ak, 
1941), 
and 
the 
time 
during 
which 
the 
faunas 
lived 
is 
designated 
the 
Hemphillian 
land-mammal 
age 
(Evernden, 
et 
ak, 
1964). 
The 
Coffee 
Ranch 
local 
fauna 
is 
the 
typelocal 
fauna 
of 
the 
Hemphillian 
fauna. 
Potassium-
argon 
dates 
for 
the 
Hemphillian 
stage 
range 
from 
5.2 
to 
10 
million 
years 
BP 
(Evernden, 
et 
ak: 
164).
The 
Coffee 
Ranch 
quarry 
was 
designated 
“Locality 
20” 
by 
Matthew 
and 
Stirton 
(1930a) 
in 
their 
first 
account 
of 
the 
fossil 
mammals 
discovered 
in 
Hemphill 
County. 
The 
list 
of 
28 
fossil-bearing 
localities 
in 
Hemphill 
County, 
of 
which 
the 
Coffee 
Ranch 
quarry 
was 
number 
20, 
did 
not 
appear 
until 
two 
years 
later, 
however 
(Reed 
and 
Longnecker,

1932: 
66). 
For 
an 
account 
of 
the 
discovery 
and 
earlywork 
at 
the 
quarry, 
see 
Matthew 
and 
Stirton 
(1930a: 
171).

The 
first 
paper 
concerning 
the 
Coffee 
Ranch 
quarry 
fossils 
by 
Matthew 
and 
Stirton 
(1930a)
dealt 
with 
the 
bone-eating 
dog 
( 
Osteohorus 
cyonoides). 
This 
was 
followed 
(1930b) 
by 
a 
study 
of 
the 
fossil 
horse 
remains. 
Burt 
(1931) 
published 
an 


* 


Department 
ofBiology, 
Midwestern 
University. 


account 
of 
the 
saber-toothed 
cat 
( 
Machairodus 
catocopis). 
Following 
Matthew’s 
death, 
Stirton 
completed 
an 
account 
of 
the 
rhinoceroses 
(Matthew, 
1932). 
Descriptions 
of 
the 
remainder 
of 
the 
fauna 
were 
promised 
(Matthew, 
1932: 
411) 
but 
never 
appeared. 
Specimens 
from 
the 
Coffee 
Ranch 
have 
been 
mentioned 
briefly 
in 
a 
number 
of 
works 
dealing 
principally 
with 
other 
matters, 
and 
Webb 
(1965) 
has 
described 
a 
new 
species 
of 
camel 
from 
the 
deposits. 
No 
papers 
devoted 
to 
mammals 
Irorn 
the 
Coffee 
Ranch 
have 
appeared 
since 
1932, 
how


ever, 
and 
no 
important 
collections 
had 
been 
made 
at 
the 
locality 
since 
1929, 
until 
field 
parties 
from 
Midwestern 
University 
began 
collecting 
in 
1956. 
In 
the 
summers 
of 
1963 
and 
1964, 
large 
bulldozers 
were 
used 
to 
push 
aside 
the 
overburden 
of 
volcanic 
ash 
and 
clay, 
to 
expose 
fresh 
matrix. 
The 
large 
adlection 
subsequently 
obtained 
includes 
remains 
of 
manycarnivorousmammals, 
which 
furnish 
thebasis 
of 
the 
present 
report. 
Earlier 
accounts 
of 
several 
taxa 
can 
now 
be 
expanded 
and 
one 
new 
speciesrecognized. 


Acknowledgments 


I 
wish 
to 
thank 
especially 
Mr. 
Walter 
Coffee 
of 
Miami, 
Texas, 
who 
permitted 
access 
to 
the 
quarry 
on 
his 
land 
and 
skillfully 
operated 
the 
heavy 
machinery 
that 
uncovered 
and 
made 
the 
new 
material 
accessible. 
More 
than 
thirty 
persons, 
mostly 
students 
at 
Midwestern 
University, 
aided 
in 
the 
excavations; 
Jack 
Cearley 
and 
Thomas 
Becldow 
worked 
at 
the 
quarry 
for 
more 
than 
two 
months 
in 
1963, 
and 
Frank 
Judd 
and 
Richard 
Palmer 
worked 
there 
through 
most 
of 
the 
summer 
of 
1964. 
Claude 
Hibbard 
of 
the 
University 
of 
Michigan 
aided 
in 
identifications 
of 
some 
of 
the 
material. 
J. 
R. 
Macdonald 
of 
the 
Los 
Angeles 
County 
Museum 
has 
confirmed 
the 
identification 
of 
the 
Indarctos. 
Wann 
Langston,
Jr. 
and 
Ernest 
L. 
Lundelius 
of 
the 
University 
of 
Texas 
at 
Austin 
and 
Donald 
E. 
Savage 
of 
the 
University 
of 
California, 
Berkeley, 
have 
offered 
helpful 
criticisms 
of 
the 
manuscript. 
Drawings 
are 
byMrs. 
Doris 
Tischler. 



Geology, 
Occurrence 
and 
Taphonomy 


The 
Coffee 
Ranch 
quarry 
is 
located 
in 
the 
southwestern 
part 
of 
Hemphill 
County 
(Fig. 
1). 
For 
a 
detailed 
description 
see 
Reed 
and 
Longnecker(1932: 
66; 
their 
locality 
20). 
The 
locality 
is 
also 
shown 
on 
their 
geologic 
map 
of 
Hemphill 
County.

The 
Coffee 
Ranch 
fossils 
accumulated 
in 
a 
lake 
or 
bog 
of 
moderate 
but 
unknown 
area. 
The 
lacustrine 
sediments 
were 
later 
buried 
under 
30 
feet 
or 
more 
of 
sandy 
aeolian 
deposits 
and 
are 
now 
exposedin 
vertical 
section 
in 
the 
face 
of 
a 
steep 
hillside. 
To 
the 
south 
the 
sediments 
continue 
into 
the 
hill 
for 
an 
undetermined 
distance, 
while 
the 
portion 
that 
once 
extended 
northward 
has 
been 
destroyed 
by 
erosion 
of 
Red 
Deer 
Creek. 
The 
exposure 
is 
lenticular 
in 
profile 
and 
some 
300 
feet 
long.

A 
measured 
section 
of 
the 
quarry 
is 
given 
byReed 
and 
Longnecker 
(1932: 
59-60). 
Their 
section 
was 
apparently 
measured 
near 
the 
edge 
of 
the 
lenticular 
deposit. 
Closer 
to 
the 
center 
the 
sediments 
measure: 


1.Overburden 
of 
huffy, 
sandy 
clay 
and 
soil 
25.0 
feet 
2. 
Volcanic 
ash 
9.0 
3. 
Compact 
bentonitic 
clay 
2.0 
4. 
Greenish 
gray 
sand 
with 
some 
clay 
2.0 
5. 
Reddish 
brown, 
sandy 
clay, 
variable 
in 
thickness 
in 
the 
deposit 
and 
with 
sharpbut 
contorted 
contact 
with 
beds 
above 
and 
beneath 
1.0 
6. 
Greenish 
sand 
and 
sandy 
clay 
with 
some 
pebble 
bands 
and 
thin 
calcareous 
sandstone 
layers 
5.5 
7. 
Slick, 
hard, 
reddish 
brown 
clay 
with 
calcareous 
crusts 
and 
nodules 
0.2 
8. 
Buff-colored 
aeolian 
sandy.. 
sediments 
of 
the 
Ogallala 
formation; 
bottom 
not 
exposed 
in 
this 
area. 


Vertebrate 
fossils 
occur 
in 
all 
beds 
of 
the 
lake 
deposit. 
Some 
rich 
concentrations 
were 
found 
in 
the 
volcanic 
ash, 
but 
in 
most 
of 
the 
ash 
layer 
bones 
are 
few 
and 
scattered. 
In 
the 
thinner 
lamina 
of 
ash, 
near 
the 
bottom 
of 
this 
bed, 
ripple 
marks 
and 
footprints 
of 
birds 
and 
mammals 
are 
common. 


The 
dense, 
bentonitic 
clay 
just 
beneath 
the 
vol-

Fig. 
1. 
General 
location 
of 
the 
middle 
Pliocene 
(Hemphillian)
Coffee 
Ranch 
vertebrate 
quarry, 
Hemphill 
County, 
Texas. 
For 
further 
discussion 
see 
text. 


canic 
ash 
contains 
an 
abundance 
of 
bones, 
but 
many 
of 
these 
are 
crushed 
and 
broken. 
A 
few 
goodspecimens 
were 
collected 
here, 
however, 
and 
most 
of 
the 
fossils 
of 
small 
mammals, 
such 
as 
mice 
and 
shrews, 
were 
obtained 
by 
washing 
the 
clay.

Most 
of 
the 
greenish, 
sandy 
clay 
beneath 
the 
bentonitic 
clay 
holds 
relatively 
few 
fossils, 
and 
these 
usually 
are 
poorly 
preserved. 
Some 
good 
specimens 
were 
found 
in 
the 
lower 
part 
of 
this 
bed, 
justabove 
the 
contact 
with 
the 
hard, 
brown, 
sandy 
clay 
of 
Bed 
5. 


The 
brown, 
sandy 
clay 
bed 
yielded 
only 
scattered 
fragments 
and 
a 
few 
poorly 
preserved 
fossils. 
Over 
extensive 
areas 
it 
is 
completely 
barren. 
The 
bulk 
of 
the 
fossils, 
and 
most 
of 
the 
complete 
bones, 
were 
foundinthegreenish 
sandofBed 
6. 


Little 
evidence 
of 
stratification 
is 
apparent 
in 
Bed 
6, 
but 
there 
are 
occasional 
layers 
of 
pebbles, 
caliche 
gravel, 
or 
indurated, 
calcareous 
sandstone. 
The 
layers 
are 
often 
extensive 
but 
rarely 
exceed 
an 
inch 
in 
thickness. 


Bones, 
bone 
fragments, 
and 
teeth 
may 
be 
found 



throughout 
Bed 
6 
but 
are 
most 
common 
in 
the 
following 
zones: 
free 
in 
the 
sand 
about 
one 
foot 
beneath 
thecontactwithBed5;inand 
justabovethe 
pebble 
or 
gravel 
layers; 
just 
above 
the 
sandstone 
layers; 
and 
at 
the 
bottom 
of 
the 
greenish 
sand 
bed, 
resting 
on 
the 
slick, 
reddish, 
calcareous 
clay 
at 
the 
bottom 
of 
the 
deposit. 
Some 
depressions 
in 
the 
ancient 
lake 
bottom, 
a 
yard 
or 
more 
in 
diameter 
and 
two 
feet 
or 
more 
in 
depth, 
were 
filled 
with 
jumbled 
massesofribs,limbbones,vertebrae, 
jaws,andother 
elements, 
derived 
from 
mammals 
of 
several 
different 


genera. 
The 
specimens 
in 
such 
pockets 
were 
often 
so 
interlocked 
that 
the 
individual 
bones 
could 
be 
separated 
onlywithgreat 
difficulty.

The 
history 
of 
the 
deposit 
may 
be 
interpreted 
as 
follows: 
sand 
and 
dust 
settled 
in 
the 
basin 
of 
the 
shallow, 
seasonal 
lake 
to 
form 
a 
sandy 
mud. 
Bones 
of 
animals 
that 
died 
in 
or 
near 
the 
lake, 
and 
that 
escaped 
destruction 
by 
scavengers, 
settled 
throughthemudtorest 
nearthebottom.Otherbones, 
which 
may 
have 
accumulated 
on 
the 
surface 
when 
the 
lake 
was 
dry, 
settled 
through 
the 
mud 
when 
the 
lake 


again 
filled 
withwater. 


The 
lake 
must 
have 
been 
dry 
for 
extended 
periods, 
for 
the 
sandstone 
layersprobably 
were 
formed 
when 
rainwater 
leached 
carbonates 
downward 
through 
a 
dry 
substratum. 
The 
pebble 
and 
gravellayers 
doubtless 
formed 
when 
heavy 
rains 
washed 
rock 
debris 
from 
nearby 
caprock 
hills 
and 
cliffs 
out 
onto 
a 
firm, 
sandy 
flat, 
not 
onto 
soft 
mud. 
Sandstone 
and 
pebble 
layers 
were 
later 
buried 
under 
accumulating 
sandy 
mud, 
but 
each 
hard 
layer 
served 
as 
a 
trap 
to 
catch 
bones 
sinking 
down 
through 
the 
soft 
sediments 
above 
them. 


The 
hard, 
brown, 
sandy 
clay 
deposit 
(Bed 
5) 
may 
have 
formed 
under 
somewhat 
different 
climatic 
conditions. 
The 
amount 
of 
clay 
and 
carbonate 
in 
this 
bed 
suggests 
slower 
deposition, 
and 
the 
reddish 
brown 
color 
suggests 
that 
the 
sediments 
were 
usually 
exposed 
to 
the 
air. 
Slow 
deposition 
and 
swift 
drainage 
or 
evaporation 
would 
also 
account 
for 
the 
scarcity 
offossil 
bones 
in 
thisbed. 


After 
the 
brown, 
sandy 
clay 
accumulated 
there 
was 
a 
return 
to 
original 
conditions, 
and 
greenish 
sand 
again 
formed. 
Vertebrate 
bones 
sank 
throughthe 
sand 
to 
a 
level 
justabove 
the 
hard, 
brown 
bed. 


The 
new 
period 
of 
deposition 
of 
greenish 
sand 
was 
brief. 
The 
site 
then 
became 
a 
meadow 
or 
shallow 
bog 
where 
mice, 
shrews, 
and 
rabbits 
lived, 
and 
whose 
remains 
were 
preserved 
in 
the 
bentonitic 
clay.
Dust, 
fine 
sand, 
and 
fine 
volcanic 
ash 
form 
the 
bentonitic 
clay. 


Increased 
volcanic 
activity 
to 
the 
west 
then 
showered 
the 
hills 
about 
the 
lake 
basin 
with 
a 
dust 
of 
volcanic 
ash. 
Each 
heavy 
rain 
washed 
the 
ash 
into 
the 
shallow 
water 
of 
the 
lake. 
As 
the 
ash 
became 
waterlogged 
it 
sank 
to 
form 
a 
mud 
which, 
when 
the 


lake 
periodically 
became 
dry, 
retained 
ripple 
marks 
and 
footprints 
of 
wading 
animals. 
Additional 
ash 
settled 
and 
rains 
washed 
it 
into 
the 
basin 
to 
refill 
the 
lake 
and 
cover 
the 
hardened 
layer 
of 
ash 
with 
a 
new 
layer 
of 
mud, 
preserving 
bones 
and 
footprints.
Layer 
after 
layer 
of 
such 
volcanic 
dust 
was 
washed 
into 
the 
lake 
until 
nine 
feet 
had 
accumulated. 
The 
hard 
ash 
bed 
completely 
filled 
the 
closed 
depressionof 
the 
lake 
bed, 
sealing 
the 
fossils 
and 
sediments 
beneath 
like 
a 
cork 
in 
a 
bottle. 
The 
capped 
deposit 
was 
then 
buried 
in 
the 
continuing 
accumulation 
of 
aeolian 
sediments 
of 
the 
Ogallala 
formation, 
until 
exposed 
byerosionofRedDeerCreek. 


Osteoborus 
was 
doubtless 
the 
major 
scavenger 
at 
the 
fossil 
site. 
On 
one 
slab 
of 
volcanic 
ash, 
ten 
feet 
long 
and 
six 
feet 
wide, 
a 
small 
ungulate 
rib 
was 
exposed 
when 
the 
slab 
was 
split 
open. 
Surroundingthe 
rib 
were 
numerous 
footprints 
of 
bone-eatingdogs. 
A 
few 
feet 
away 
the 
trackway 
of 
a 
saber-
toothed 
cat, 
the 
footprints 
evenly 
spaced 
and 
unhurried,
wentpast 
theclustereddogfootprints.One 
can 
only 
conjecture 
that 
the 
cat 
frightened 
the 
dogs 
awayfrom 
theirmeal. 


The 
name 
“bone-eating” 
dog 
may 
be 
validated 
by 
other 
than 
anatomical 
features. 
Found 
in 
the 
bentonitic 
clay 
were 
numerous 
small 
clusters 
of 
bone 
chips. 
These 
sometimes 
were 
evenly 
but 
densely 
distributed 
through 
fist-sized 
areas 
of 
claythat 
were 
not 
otherwise 
marked 
off 
from 
the 
surrounding 
matrix. 
The 
bone 
chips 
were 
rarely 
more 
than 
10 
mm. 
in 
greatest 
dimension, 
and 
presumably 
represent 
the 
disintegrated 
feces 
of 
Osteoborus. 


The 
dogs 
were 
possibly 
cannibalistic, 
as 
the 
hunting 
dogs 
of 
Africa 
are 
reputed 
to 
be. 
The 
species 
is 
the 
most 
common 
carnivore 
in 
the 
deposit, 
represented 
by 
more 
than 
50 
skulls 
and 
jaw 
fragments.
Yet 
very 
little 
postcrania! 
material 
was 
recovered 
and 
much 
of 
that 
is 
damaged. 
For 
example, 
the 
humerus 
of 
Osteoborus 
is 
represented 
by 
eight 
distal 
portions 
but 
not 
a 
single 
proximal 
end. 
Doubtless 
the 
scapulae 
and 
meaty 
proximal 
ends 
of 
the 
humeri 
were 
devoured, 
but 
the 
distal 
ends, 
where 
there 
is 
little 
flesh, 
were 
less 
attractive. 
Lower 
legand 
foot 
bones 
are 
more 
common, 
but 
vertebrae 
with 
protruding 
neural 
spines 
and 
transverse 
processes 
appear 
to 
have 
been 
heavily 
gnawed 
and 
largely 
destroyed. 



The 
Osteoborus 
series 
included 
only 
two 
specimens 
with 
milk 
teeth. 
Fifty-four 
adult 
dentitions 
were 
arbitrarily 
separated 
into 
four 
age 
categories; 
young 
(little 
or 
no 
wear 
on 
teeth), 
medium-aged 


(moderate 
tooth 
wear), 
old 
(considerable 
wear 
on 
all 
teeth), 
and 
very 
old 
(teeth 
worn 
almost 
flat). 
Upper 
dentitions 
were 
separated 
as 
follows: 
7 
young, 
10 
medium, 
2 
old, 
and 
3 
very 
old. 
Lower 
dentitions 
included: 
9 
young, 
13 
medium, 
8 
old, 
and 
2 
very 
old. 
Maximum 
mortality 
seems 
to 
have 
occurred 
among 
the 
fully 
mature 
animals. 


Vultures, 
too, 
probably 
aided 
in 
scattering 
bones 
at 
the 
quarry. 
On 
a 
number 
of 
bones, 
protruding 
processes 
and 
sharp 
edges 
appear 
to 
have 
been 
damaged 
by 
nibbling 
or 
pecking, 
but 
there 
are 
no 
distinct 
tooth 
marks. 
This 
is 
especially 
apparent 
in 
the 


skeleton 
of 
the 
saber-toothed 
cat, 
described 
later. 
The 
broken 
edges 
of 
the 
cranium 
of 
this 
specimenhave 
a 
pecked, 
nibbled 
appearance, 
while 
many 
of 
the 
transverse 
processes 
and 
neural 
spines 
of 
the 
vertebrae 
are 
damaged 
or 
missing 
on 
one 
side 
only. 
No 
actual 
remains 
of 
vultures 
were 
discovered, 
how-
ever.Theonlybird 
tracksfoundinthevolcanicash 
appear 
to 
be 
of 
a 
heron-like 
form, 
and 
the 
only 
largebird 
bone 
found 
was 
identified 
as 
a 
hawk 
by 
Dr. 
Pierce 
Brodkorb 
of 
the 
University 
of 
Florida. 


Many 
bones 
were 
broken 
or 
crushed 
by 
the 
feet 


of 
large 
mammals 
wading 
in 
the 
mud 
of 
the 
shal


low 
lake. 
Rhinoceroses 
were 
probably 
responsible

for 
much 
of 
this 
damage. 
One 
rhinoceros 
skull 


showed 
clearly 
that 
a 
heavy 
foot 
had 
crushed 
the 


frontal 
area. 
The 
skull 
lay 
in 
the 
matrix 
in 
three 
pieces: 
cranium 
and 
two 
separated 
rnaxillaries, 
with 
a 
mass 
of 
bone 
splinters 
between 
them. 
The 
three 
parts, 
reassembled, 
showed 
a 
gaping 
hole 
where 
the 
foot 
had 
descended. 
The 
skull 
of 
the 
saber-
toothed 
cat 
also 
was 
clearly 
damaged 
in 
this 
fashion. 
The 
articidated 
skeleton 
(Plate 
I) 
lay 
in 
greenishsand 
matrix 
a 
few 
inches 
above 
a 
layer 
of 
sandstone. 
Atlas 
and 
axis 
were 
separated 
from 
the 
other 
cervical 
vertebrae 
by 
three 
inches 
and 
the 
basicranium 
of 
the 
skull 
lay 
three 
inches 
ahead 
of 
them. 
The 
main 
weight 
of 
the 
crushing 
foot 
had 
come 
to 
rest 
on 
the 
anterior 
left 
part 
of 
the 
skull. 
The 
right

maxillary 
with 
canine 
and 
carnassial 
had 
been 
pressed 
down 
until 
it 
lay 
on 
the 
sandstone 
layer. 
A 
right 
premaxillary, 
doubtless 
of 
the 
same 
skull, 
was 
found 
a 
short 
distance 
away. 
Both 
glenoid 
processes 
were 
broken 
free 
and 
lay 
in 
the 
matrix 
beneath 
their 
natural 
positions, 
but 
the 
remainder 
of 
the 
zygomatic 
arches 
were 
splintered. 
The 
remainder 
of 
the 
skull 
(left 
maxillary) 
was 
not 
found. 
The 
toothless 
skull 
of 
a 
wolverine, 
Plesiogulo, 
also 
appears 
to 
have 
been 
crushed 
by 
a 
wading 
mammal, 
but 
the 
damageis 
less 
distinctive. 


In 
the 
accounts 
that 
follow, 
all 
measurements 
of 
specimens 
are 
in 
millimeters. 
Specimen 
numbers 
are: 
MU, 
Midwestern 
University, 
Wichita 
Falls,
Texas, 
and 
TMM, 
Texas 
Memorial 
Museum, 
Austin. 
Figured 
specimens 
are 
in 
the 
collection 
of 
the 
latter. 


Taxonomic 
Synopsis 


Order 
Carnivora 
Bowdich 
SuborderFissipeda 
Blumenbach 
Family 
Canidae 
GraySubfamily 
Caninae 
Gill 
Vulpes 
cf. 
V. 
shermanensis 
Hibbard 
Subfamily 
Borophaginae 
SimpsonOsteoborus 
cyonoides 
(Martin)
Family 
Ursidae 
GrayIndarctos 
oregonensis 
Macdonald 
Family 
Mustelidae 
Swainson 


Subfamily 
Mustelinae 
Gill 
Plesiogulo 
marshalli 
(Martin)


Subfamily 
Melinae 
Burmeister 
Pliotaxidea 
cf. 
P. 
nevadensis 
(Butterworth)

Family 
Felidae 
Gray 
Subfamily 
Nimravinae 
Trouessart 
Pscudaelurus 
hibbardi 
DalquestSubfamily 
Machairodontinae 
Gill 


Machairodus 
catocopis 
Cope 



Accounts 
of 
Species 


Vulpes 
cf. 
V. 
shermanensis 
(Hibbard) 


Two 
edentulous 
lower 
jaws 
and 
a 
number 
of 
isolated 
teeth 
belong 
to 
a 
fox 
approximately 
the 
size 
of 
an 
adult 
red 
fox, 
I'ulpes 
vulpes. 
Jaw 
MU6866 
lacks 
the 
ascending 
ramus 
but 
has 
the 
alveoli 
of 
the 
canine 
and 
cheek 
teeth. 
When 
this 
jaw 
is 
placed 
beside 
the 
jaw 
of 
a 
red 
fox 
from 
north-central 
Texas, 
from 
which 
the 
teeth 
have 
been 
removed, 
the 
alveoli 
of 
the 
canine, 
premolars 
and 
carnassial 
match 
closely, 
but 
the 
alveoli 
for 
M 
2 
and 
M 
3 
are 
more 
widely 
spaced, 
and 
the 
jaw 
is 
longer 
in 
the 
corresponding 
region. 
The 
distance 
from 
the 
anterior 
edgeof 
the 
canine 
alveolus 
to 
the 
posterior 
edge 
of 
the 
alveolus 
for 
M 
3 
is 
79.0 
mm., 
or 
slightly 
greater 
than 
in 
the 
jaws 
of 
several 
red 
foxes 
from 
Texas. 


FourisolatedML’s(MU4362,4363, 
6406, 
6497) 
are 
similar 
to 
the 
lower 
carnassials 
of 
red 
foxes 
from 
Texas 
in 
size 
and 
morphologic 
details, 
but 
in 
each 
tooth 
the 
metaconid 
is 
distinctly 
larger 
and 
more 
prominent.Except 
forthisdifference,and 
thelargerM 
2 
and 
M.„ 
the 
lower 
jaw 
of 
the 
Hemphill 
fox 
is 
quite 
like 
that 
of 
modern 
red 
foxes. 
Canid 
lower 
jaws 
are 
usually 
conservative, 
however, 
and 
if 
complete 
upper 
jaws 
or 
skulls 
were 
available, 
greaterdifferences 
between 
them 
would 
probably 
be 
found. 


Osteoborus 
cyonoides 
(Martin) 


The 
bone-eating 
dog 
is 
the 
commonest 
carnivore 
in 
the 
Coffee 
Ranch 
Quarry. 
Four 
skulls 
(Table 
1), 
18 
maxillaries 
or 
maxillary 
fragments 
with 
teeth,
32 
lower 
jaws 
or 
mandibular 
fragments 
containingteeth, 
numerous 
isolated 
teeth, 
and 
isolated 
bones 
representing 
most 
of 
the 
skeleton 
were 
recovered. 


In 
spite 
of 
the 
abundance 
of 
cranial 
fragments,
postcranial 
material 
is 
uncommon 
and 
fragmentary. 
No 
useful 
remains 
of 
pelvis 
or 
sacrum 
were 
obtained, 
and 
only 
the 
proximal 
portions 
of 
two 
scapulas. 
A 
femur 
is 
nearly 
complete 
but 
only 
fragments 
of 
humeri 
were 
found. 
Lower 
leg 
and 
foot 
elements 
are 
more 
common. 
A 
number 
of 
vertebrae 
were 
taken 
but 
only 
some 
of 
the 
cervicals 
and 
posterior 
thoracics 
can 
be 
identified 
with 
confidence. 


DESCRIPTION.—An 
excellent 
skull, 
jaws, 
and 
some 
postcranial 
bones 
ol 
this 
dog 
were 
briefly 
described 
and 
figured 
by 
Matthew 
and 
Stirton 
(1930a), 
but 
only 
a 
few 
measurements 
were 
given.
The 
following 
data, 
based 
on 
newly-collected 
specimens, 
augment 
the 
work 
of 
Matthew 
and 
Stirton. 


.Skull.—MU7408 
is 
the 
broken 
skull 
of 
a 
youngdog. 
The 
permanent 
teeth 
are 
in 
place 
and 
are 
scarcely 
worn. 
The 
skull 
lacks 
the 
tip 
of 
the 
rostrum, 
the 
teeth 
anterior 
to 
the 
second 
premolars, 
and 
the 
right 
M 
2, 
as 
well 
as 
part 
of 
the 
anterior 
dorsal 
skull 
roof. 
The 
skull 
had 
been 
broken 
into 
three 
pieces;
basicraniurn 
and 
two 
maxillaries. 
No 
point 
of 
bone-
contact 
between 
the 
three 
pieces 
remained. 
The 
missing 
parts 
have 
been 
restored 
with 
plaster, 
but 
measurementsofbreadth 
are 
not 
usable. 


MU3229 
is 
the 
skull 
of 
a 
young-adult 
dog, 
with 
the 
teeth 
scarcely 
worn 
except 
that 
the 
tips 
of 
the 
fourth 
premolars 
show 
some 
wear. 
"The 
right 
11,I1 
right 
andleft 
Px’s,right 
zygomaticarch, 
andright, 
posterior 
portion 
of 
skull 
are 
missing. 
The 
skull 
is 
embedded 
in 
a 
hard 
calcareous 
matrix 
and 
has 
not 
been 
completely 
prepared.

MU5049 
is 
also 
the 
skull 
of 
a 
young-adult 
dog, 
a 
bit 
older 
than 
MU3229. 
Missing 
are 
all 
teeth 
anterior 
to 
the 
P2’s, 
the 
right 
M 
2, 
the 
left 
M 
1 
and 
M2and 
left 
zygomatic 
arch. 
There 
is 
also 
some 
minor 
crushing 
andbreakage.

MU5862 
is 
from 
an 
older 
animal, 
and 
a 
mod


erate 
amount 
of 
wear 
is 
present 
on 
even 
the 
canines 


and 
incisors. 
Missing 
is 
the 
right 
Pl 
right 
and 
left

, 


P3 
and 
left 
M 
2, 
as 
well 
as 
all 
of 
the 
skull 
posterior 
to 


the 
palate. 
Additional 
material 
includes 
18 
maxil


lary 
fragments, 
some 
with 
the 
premaxillaries 
at


tached. 


The 
four 
skulls 
are 
from 
animals 
considerably 
younger 
than 
the 
ncarly-perfect 
specimen 
figuredby 
Matthew 
and 
Stirton 
(1930a, 
Plates 
21, 
24, 
25, 
26), 
but 
they 
are 
essentially 
similar 
in 
shape. 
The 
braincase 
is 
narrow 
but 
the 
remainder 
of 
the 
skull 
is 
broad 
and 
powerful. 
The 
zygomatic 
arches 
are 
stout 
and 
broad 
posteriorly. 
Rostrum 
and 
palate 
are 
broad. 
The 
breadth 
is 
accentuated 
by 
the 
short



ness 
of 
the 
muzzle, 
which 
results 
from 
the 
shortening 
of 
the 
premolar 
region 
of 
the 
skull 
and 
reduction 
of 
the 
premolars. 
The 
frontal 
region 
is 
greatlybulged, 
in 
the 
manner 
typical 
of 
hyaenoid 
dogs. 
1 
he 
zygomatic 
arches 
are 
narrow 
anteriorly 
but, 
posterior 
to 
the 
orbits, 
flare 
widely 
to 
permit 
passage 
of 
the 
enormous 
masseter 
muscles. 
The 
orbits 
are 
small 
and 
diagonally 
constricted 
by 
the 
bulged 
forehead 
above 
and 
flared 
arches 
behind. 
In 
life 
the 
eyes 
must 
have 
been 
strongly 
slanted. 
The 
sagittal 
crest 
is 
very 
heavy, 
and 
the 
occipital 
crest 
is 
strong 
and 
projects 
posteriorly 
above. 
It 
is 
transversely 
truncated 
posteriorly 
rather 
than 
ending 
in 
a 
point, 
as 


in 
Canis. 
The 
head, 
in 
life, 
might 
have 
resembled 
somewhat 
that 
of 
a 
modern 
bulldog, 
but 
the 
expanded 
frontals 
and 
powerful 
muscles 
of 
the 
temporal 
region 
and 
of 
the 
jaws 
must 
have 
appearedquite 
different. 


Superior 
dentition.—There 
is 
a 
greater 
amount 
of 
individual 
variation 
in 
the 
series 
of 
upper 
jaws 
of 
O. 
cyonoides 
than 
in 
a 
comparable 
series 
of 
skulls 
ofthemoderncoyote,Canis 
latrans.Variationinthe 
relative 
size 
and 
crowding 
of 
the 
teeth 
is 
especially 
apparent. 
In 
skull 
MU3229 
the 
teeth 
are 
small 
and 
widely 
spaced. 
The 
individual 
teeth 
are 
separated

by 
short 
spaces, 
and 
the 
long 
axes 
of 
the 
premolars 
are 
parallel 
to 
the 
axis 
of 
the 
tooth 
row. 
The 
alveolar 
length, 
from 
P2 
to 
M 
2, 
is 
relatively 
and 
actuallylong 
(63.8 
mm.). 
In 
skull 
MU5862 
the 
teeth 
are 
relatively 
large 
and 
the 
toothrow 
short 
(P2-M2 


58.5 
mm.). 
The 
teeth 
are 
in 
contact, 
and 
the 
anterior 
premolars 
crowded 
until 
they 
are 
rotated 
out 


of 
the 
axis 
of 
the 
tooth 
row. 
Even 
more 
extreme 


crowding 
is 
seen 
in 
some 
maxillary 
fragments.

There 
is 
considerable 
variation 
in 
the 
size 
and 


shape 
of 
the 
small, 
anterior 
premolars. 
Unworn 


teeth 
may 
be 
elongate 
or 
broadly 
oval. 
There 
may

be 
a 
single 
central 
cusp 
or 
three 
distinct 
cusps. 
There 


is 
an 
equal 
amount 
of 
variation 
in 
size 
of 
the 
car


nassial 
and 
upper 
molars. 
In 
some, 
the 
upper 
P 
4 
’s 


are 
more 
than 
20 
percent 
larger 
than 
others. 
The 


difference 
is 
so 
great 
that, 
were 
only 
the 
largest 
and 


smallest 
teeth 
available, 
one 
might 
place 
them 
in 


separate 
species.

There 
is 
little 
variation 
in 
the 
shape 
of 
the 
carnassial 
and 
first 
molar, 
however. 
The 
parastyle 
is 
well 
developed 
and 
present 
on 
every 
specimen 
at 
hand. 
Vanderhoof 
and 
Greggory 
(1940, 
footnote 


p. 
160) 
state 
that 
the 
parastyle 
on 
P4 
is 
“a 
demonstrably 
variable 
character, 
especially 
in 
Osteoborus, 
where 
it 
ranges 
from 
total 
absence 
to 
large 
size,” 
but 
earlier 
in 
the 
same 
paper 
(p. 
144) 
these 
authors 
list 
the 
parastyle 
as 
a 
character 
of 
the 
genus 
Osteoborus. 
Canid 
dentitions 
are 
variable 
and 
that 
of 


O. 
cyonoides 
unusually 
so. 
It 
would 
not 
be 
surprising 
if 
occasional 
individuals 
lacked 
the 
parastyle.
In 
the 
more 
than 
20 
specimens 
available, 
however, 
the 
parastyle 
is 
present 
and 
strong 
in 
all. 
The 
first 
incisors 
of 
O. 
cyonoides 
are 
small, 
the 
second 
only 
slightly 
larger, 
and 
the 
third 
strong 
and 
stout, 
resembling 
the 
canine 
in 
shape 
but 
only 
about 
half 
its 
size. 
Poorly 
developed 
basal 
cusps 
are 
present 
on 
P 
and 
at 
least 
sometimes 
on 
the 
posteriorbase 
of 
13.I3 
. 
The 
huge 
cusps 
of 
the 
upper 
incisors 
of 
Aelurodon 
are 
not 
even 
approximated 
by 
O. 
cyonoides. 
Few 
measurements 
of 
the 
upper 
incisors 
are 
possible 
for 
I 
1 
and 
12 
are 
rarely 
retained 
in 
the 
fossils, 
and 
I 
3 
is 
only 
slightly 
more 
common. 
The 
canines 
are 
stout 
and 
short 
but 
otherwise 
doglike.
Their 
roots 
are 
heavy, 
and 
the 
tooth 
is 
often 
retained 
in 
the 
fossil 
jaws.

The 
three 
anterior 
premolars 
are 
small 
and 
grad


uatedin 
size,withthefirstthesmallest.Thistoothis 
usually 
lost 
from 
the 
fossil 
jaws. 
P2 
is 
two-rooted. 
It 
is 
almost 
always 
oriented 
anteroposteriorly 
in 
the 
jaw 
and 
is 
firmly 
fixed 
and 
usually 
present 
in 
the 
fossils. 
P3 
is 
larger 
than 
P2 
but, 
if 
the 
tooth 
row 
is

, 


crowded, 
this 
tooth 
is 
rotated 
or 
forced 
out 
of 
alignmentbetween 
P2andP4.Itislessfirmlyfixedin 
the 
jaw 
than 
P2 
and 
is 
more 
often 
missing 
in 
fossil 


, 


jaws. 
Many 
premolars 
were 
lost 
in 
life 
and 
their 
alveoli 
almost 
completely 
obscured 
by 
bone 
growth.

P 
4 
is 
a 
large, 
strong 
tooth. 
There 
is, 
at 
most, 
only 
a 
trace 
of 
the 
protocone, 
but 
the 
tooth 
is 
noticeablybroader 
anteriorly 
where 
the 
protocone 
seems 
to 
have 
been 
taken 
into 
the 
body 
of 
the 
tooth. 
The 
root 
remains 
lingually. 
The 
bulk 
of 
the 
tooth 
consists 
of 
the 
small 
but 
distinct 
parastyle, 
large 
paracone, 
and 
metacone. 


M 
1 
is 
a 
large, 
stout 
tooth. 
It 
is 
well 
fixed 
in 
the 
jaw 
and 
seldom 
lost 
in 
fossils. 
M 
2 
is 
much 
smaller 
and 
is 
quite 
variable 
in 
size. 
It 
is 
often 
lost 
in 
fossil 
specimens. 
Because 
P 
4 
and 
M 
2 
are 
so 
often 
lost, 
they 
are 
not 
included 
in 
measurements 
of 
lengthbetween 
teeth 
(Table 
2). 
In 
the 
measurements 
of 
length 
of 
the 
individual 
teeth, 
the 
longest 
axis 
is 
given 
as 
length 
of 
the 
premolars, 
regardless 
of 
the 
orientationinthejaw.Breadth 
isthegreatestwidth 
at 
right 
angles 
to 
the 
long 
axis. 
The 
length 
of 
M 
1 
was 
taken 
with 
the 
jaw 
of 
the 
caliper 
placed 
againstthe 
flat, 
posterior 
face 
of 
the 
tooth 
and 
width 
was 
taken 
with 
the 
jaw 
against 
the 
flat, 
labial 
side 
of 
the 
tooth. 
All 
of 
the 
measurements 
given 
in 
Table 
2 
weretakenfromjawscontainingP4Ml 
andother

,, 



teeth. 
Measurements 
indicated 
with 
“a” 
are 
from 
alveoli 
where 
the 
tooth 
is 
lost. 
No 
isolated 
teeth 
are 
included 
in 
the 
table. 


Mandible.—The 
lower 
jaw 
of 
O. 
cyonoides 
is 
short, 
deep, 
and 
thick. 
The 
ascending 
ramus 
is 
strongly 
developed. 
The 
lower 
border 
of 
the 
mandible, 
posterior 
to 
M 
2, 
curves 
upward 
and 
backward, 
so 
that 
the 
angular 
process 
is 
well 
above 
the 
inferior 
border 
of 
the 
horizontal 
ramus. 
The 
articular 
condyle 
is 
heavy 
and 
strong. 
The 
coronoid 
process 
is 
very 
high 
and 
strong. 
The 
height 
of 
the 
ascending 
ramus 
is 
as 
long 
as 
or 
longer 
than 
the 
tooth-bearing 
part 
of 
the 
horizontal 
ramus. 
The 
masseteric 
fossa 
is 
large 
and 
deep, 
crossed 
with 
diagonal 
ridges 
and 
rugosities 
for 
attachment 
of 
powerful 
muscles. 
These 
rugosities 
seem 
to 
be 
best 
developed 
in 
animals 
with 
worn 
teeth, 
and 
doubtless 
increased 
with 
age.

Inferior 
dentition.--Wariation 
in 
thelower 
dentition, 
in 
spacing 
and 
relative 
size 
of 
teeth, 
is 
equal 
to 
that 
in 
the 
upper 
teeth 
(Table 
3). 
In 
the 
largeteeth, 
P.i 
and 
M,, 
there 
is 
little 
variation 
in 
structure 
but 
much 
in 
size. 
The 
largest 
carnassials 
are 


fully 
20 
percent 
larger 
than 
the 
smallest. 
No 
such 
great 
variation 
was 
noted 
in 
the 
lower 
molars 
of 
the 
coyote.

Osteoborus 
typically 
lacks 
the 
first 
lower 
premolar, 
but 
this 
tooth 
is 
present 
in 
one 
lower 
jaw 
and 
its 
single-rooted 
alveolus 
is 
present 
in 
another. 
The 
P, 
that 
is 
present 
is 
nearly 
as 
large 
as 
the 
P2 
in 
the 
same 
jaw.

Characters 
that 
separate 
the 
lower 
dentition 
of 
Osteoborus 
from 
that 
of 
its 
descendant, 
Borophagus,
include 
the 
stepped 
posterior 
face 
of 
P, 
and 
the 
presence 
of 
a 
metaconid 
on 
Mi. 
That 
these 
are 
always 
lacking 
in 
Borophagus 
remains 
to 
be 
determined, 
but 
they 
are 
present 
in 
all 
of 
the 
34 
Mys 
and 
28 
P,’s 
of 
Osteoborus 
from 
the 
CofTee 
Ranch. 


The 
lower 
incisors 
of 
O. 
cyonoides 
are 
not 
pre


served 
in 
any 
of 
the 
32 
lower 
jaws 
available. 
Their 


alveoli 
are 
present 
in 
some 
cases, 
and 
some 
isolated 


specimens 
that 
probably 
are 
lower 
incisors 
of 
Osteo


borus 
were 
found. 
The 
lower 
incisors 
appear 
to 
have 


been 
small 
and 
simple. 
The 
canine 
is 
short, 
thick, 


and 
strong, 
with 
a 
heavy 
root. 
P, 
has 
been 
lost 
from 


most 
specimens. 
P2 
and 
P3 
are 
small, 
oval 
teeth. 
In 


unworn 
can

specimens, 
three 
small, 
low 
cuspsusually 
be 
distinguished 
but 
these 
vanish 
with 
moderate 
wear. 
The 
teeth 
vary 
considerably 
in 
size 
from 
jaw 
to 
jaw, 
but 
P3 
is 
invariably 
larger 
than 
P2 
. 
P2 
is 
often 
lost 
in 
fossil 
jaws.

P.j 
is 
a 
large, 
strong, 
thick 
tooth, 
triangular 
in 
lateral 
view 
and, 
in 
unworn 
teeth, 
strongly 
inclined 
posteriorly. 
The 
posterior 
face 
is 
invariably 
stepped.
Thistoothisrarely 
lostfromfossil 
jaws.

M, 
is 
a 
very 
large, 
strong, 
crushing 
tooth. 
The 
trigonid 
is 
subject 
to 
heavy 
wear, 
and 
in 
old 
animals 
is 
worn 
down 
to 
the 
level 
of 
the 
talonid. 
Ihe 


tooth 
is 
strongly 
attached 
in 
the 
jaw 
and 
seldom 
lost. 


M 
2 
is 
a 
moderately 
large 
and 
prominent 
tooth. 
It 
seems 
to 
receive 
relatively 
little 
wear, 
and 
it 
is 
often 
lost 
from 
fossil 
jaws. 
M 
3 
is 
a 
tiny, 
almost 
vestigial 
tooth 
that 
is 
usually 
lost 
in 
fossil 
jaws. 
Even 
the 
alveolus 
is 
sometimes 
absent, 
indicating 
that 
the 
tooth 
was 
lost 
during 
the 
life 
of 
the 
animal. 


Measurements 
of 
length 
of 
the 
lower 
tooth 
rows 
do 
not 
include 
M 
3. 
Measurements 
of 
length 
of 
individualteethwere 
takenparallel 
tothe 
long 
axisof 
thetooth,regardlessofitsposition 
inthetoothrow, 
and 
breadth 
was 
measured 
at 
right 
angles 
to 
the 
length.

Postcranial 
skeleton.—Most 
vertebrae 
are 
so 
fragmentary 
that 
recognition 
of 
their 
place 
in 
the 
spinalcolumn 
is 
uncertain 
or 
impossible. 
The 
following 
are 
identified 
withreasonable 
certainty: 
Atlas 
(MU7560), 
Axis 
(MU5693), 
third 
cervical 
(MU3872),
fifth 
cervical 
(MU4504), 
seventh 
cervical 
(MU5692), 
eleventh 
thoracic 
(MU5503), 
twelfth 
thoracic 
(MU5686), 
and 
thirteenth 
thoracic 
(MU5507). 
These 
vertebrae 
are 
slightly 
stouter 
and 
heavier 
than 
vertebrae 
of 
the 
modern 
coyote 
( 
Canis 
latrans) 
but 
arc 
otherwisequite 
similar. 


Some 
measurements 
of 
the 
atlas 
are: 
length 
of 
topofneuralarch, 
14.5;lengthofbottomofneural 
arch, 
11.3; 
least 
breadth 
just 
posterior 
to 
transverse 
processes, 
39.5; 
greatest 
breadth 
across 
transverse 
processes, 
93.7; 
maximum 
breadth, 
internal, 
of 
articular 
surfaces 
for 
occipital 
condyles, 
40.6; 
breadth 
acrossarticular 
surfacefor 
axis, 
35.1. 


Some 
measurements 
of 
the 
axis 
are; 
length 
of 


centrum 
to 
end 
of 
odontoid 
process, 
48.0; 
breadth 


across 
articular 
surfaces 
for 
atlas, 
31.9; 
breadth 


across 
posterior 
zygapophyses, 
35.2; 
breadth 
of 
pos


terior 
epiphysis 
of 
centrum, 
19.0. 
The 
anterior 
part

of 
the 
neural 
spine 
has 
been 
broken 
away 
and 
lost. 


Measurements 
of 
other 
vertebrae 
are 
given 
in 
Table 


4. 
The 
proximal 
portions 
of 
two 
scapulae 
apparentlybelong 
to 
Osteoborus, 
but 
only 
one 
of 
these 
(MU6021) 
belongs 
to 
an 
adult. 
Measurements: 
anteroposterior 
diameter 
of 
glenoid 
fossa, 
33.0; 
transverse 
diameter 
of 
glenoid 
fossa, 
21.8; 
least 
constriction 
of 
neck 
above 
glenoid 
fossa, 
26.4. 



The 
limb 
material 
is 
sufficient 
to 
show 
that 
the 
humerus 
and 
femur 
were 
considerably 
longer 
and 
much 
stouter 
than 
the 
bones 
of 
a 
coyote 
of 
comparable 
size. 
Radius, 
ulna, 
and 
tibia 
(no 
complete 
fibulae 
wei'e 
found) 
are 
almost 
the 
same 
length 
as 
the 
comparable 
bones 
in 
the 
coyote, 
but 
are 
much 
stouter. 
Relatively, 
the 
humerus 
and 
femur 
of 
the 
coyote 
are 
shorter 
than 
those 
of 
Osteoborus. 
1 
he 
humerus 
of 
Osteoborus 
differs 
radically 
in 
structure 
from 
that 
of 
Canis. 
The 
distal 
end 
is 
greatly 
expanded. 
The 
olecranon 
fossa 
is 
very 
wide, 
and 
a 
large 
entepicondylar 
foramen 
is 
present. 
This 
for-
amen 
is 
absent 
in 
Canis. 
The 
proximal 
end 
of 
the 


humerus 
of 
Osteoborus 
cyonoides 
is 
unknown. 


The 
other 
long 
bones 
are 
typically 
canid, 
exceptfor 
their 
unusual 
thickness. 
The 
metapodials 
are 
relatively 
short 
and 
thick 
but 
otherwise 
similar 
to 
the 
metapodials 
of 
the 
coyote. 


Canid, 
species 
undetermined 
(Fig. 
2) 


Among 
the 
specimens 
obtained 
at 
the 
Coffee 
Ranch 
is 
the 
partial 
skull 
of 
a 
large 
canid. 
The 
skull 
is 
broken 
across 
the 
preorbital 
region, 
just 
anterior 
to 
the 
termination 
of 
the 
nasals. 
Both 
zygomaticarches 
are 
missing, 
though 
the 
left 
glenoid 
processis 
present. 
Part 
of 
the 
parietal 
of 
the 
right 
side 
is 
broken 
away. 
The 
remainder 
of 
the 
skull 
is 
well 
preserved.

The 
intcrorbital 
region 
is 
broad 
and 
the 
post-

orbital 
processes 
are 
wide 
and 
strong. 
The 
post-

orbital 
region 
is 
strongly 
constricted. 
The 
lamb


doidal 
crest 
is 
long, 
reaching 
maximum 
height 
at 
the 
anterior 
margin 
of 
the 
parietals 
and 
then 
decreasing 
in 
height 
posteriorly, 
becoming 
obsolete 


just 
ahead 
of 
the 
occipital 
crest. 
The 
occipital 
crest 
is 
strong 
and 
broad, 
transversely 
truncated 
posteriorly. 
Preserved 
cranial 
foramina 
are 
similar 
to 
those 
of 
Canisand 
Osteoborus. 


The 
preserved 
part 
of 
the 
skull 
is 
extremely 
slender 
and 
elongate. 
It 
bears 
a 
distinct 
resemblance 
to 
the 
skull 
of 
Tomarctus 
euthos 
(McGrew) 
from 
the 
lower 
Pliocene 
Burge 
fauna 
of 
Nebraska. 
The 
skull 
is 
clearly 
of 
a 
young 
animal, 
and 
cranial 
sutures 
are 
distinct. 


Donald 
E. 
Savage 
has 
examined 
this 
specimenandbelievesittobealargelate 
juvenileOsteoborus. 
The 
specimen 
is 
as 
large 
as 
the 
comparable 
part 
of 
an 
adult 
Osteoborus 
(length 
from 
end 
of 
nasals 
to 
end 
of 
occipital 
crest 
124.2 
vs. 
135.0 
in 
Osteoborus).
The 
posterior 
and 
ventral 
parts 
of 
the 
specimen 
resembled 
the 
skull 
of 
Osteoborus 
except 
that 
the 
lambdoidal 
crest 
is 
much 
lower. 
In 
canids 
the 
lamb


doidal 
crest 
increases 
in 
size 
and 
prominence 
with 
age, 
especially 
in 
male 
animals. 


The 
greatest 
difference 
between 
the 
fragmentaryskull 
and 
the 
skull 
of 
Osteoborus 
is 
the 
nature 
of 
the 
frontal 
region. 
In 
Osteoborus 
the 
interorbitalregionis 
much 
wider 
and 
the 
frontals 
are 
greatly 
bulged.
In 
the 
fragmentary 
skull 
the 
interorbital 
region 
is 
narrower 
(38.9 
mm. 
vs. 
50.9 
in 
Osteoborus 
) 
and 
the 
frontal 
curvature 
is 
as 
smooth 
and 
gentle 
as 
in 
a 
modern 
red 
fox 
( 
Vulpes 
vulpes). 
If 
a 
straightline 
from 
the 
posterior 
ends 
of 
the 
nasals 
to 
a 
point 
on 
the 
lambdoidal 
crest 
100 
mm. 
posterior 
to 
the 


Fig. 
2. 
Cranium 
of 
an 
undetermined 
canid 
(TMM 
41261—4), 
in 
dorsal 
vtsw. 
X 
1.0 
natural 
size. 



nasals 
be 
taken 
as 
a 
base 
line, 
the 
frontals 
of 
Osteohorus 
(young-adult 
specimen) 
arch 
19 
mm. 
above 
the 
base 
line 
while 
the 
frontals 
of 
the 
fragmentaryskull 
rise 
only 
7 
mm. 
above 
the 
base 
line. 


If 
the 
fragmentary 
skull 
belonged 
to 
Osteohorus, 
it 
suggests 
that 
the 
distortion 
of 
the 
frontal 
regionof 
the 
skull, 
typical 
of 
this 
genus 
and 
Borophagus,
of 
the 
upper 
Pliocene, 
developed 
rather 
abruptlybetween 
the 
late 
juvenile 
and 
early 
adult 
stages. 
I 
know 
of 
no 
comparable 
changes 
in 
the 
skull 
during 
ontogeny 
of 
modern 
canids. 
If 
the 
fragmentary 
skull 
is 
not 
a 
juvenile 
Osteohorus, 
it 
may 
represent 
a 
late 
surviving 
species 
of 
Tomarctus, 
otherwise 
unknown. 
It 
is 
not 
Canis. 


Indarctos 
oregonensis 
Macdonald 
(Figs. 
3-4) 


Remains 
of 
a 
very 
large 
bear 
are 
among 
the 
rare 
elements 
of 
the 
Coffee 
Ranch 
local 
fauna. 
Matthew 
and 
Stirton 
(1930a) 
listed 
Hyaenarctos 
from 
the 
Coffee 
Ranch 
but 
the 
material 
has 
never 
been 
figured 
or 
described. 
The 
present 
collection 
includes 
a 
fragmentary 
lower 
jaw 
with 
P4 
to 
M 
3, 
an 
isolated 
unworn 
lower 
M,, 
and 
two 
fragmentary 
lower 
molars. 


DESCRIPTION.—A 
small 
part 
of 
the 
ascending 
ramus 
is 
present 
in 
the 
jaw 
(TMM41261-1),
and 
only 
a 
small 
amount 
of 
the 
upper 
margin 
of 
the 
mandible 
holds 
the 
molars 
in 
place. 
P, 
was 
in 
placein 
the 
matrix 
but 
free 
of 
the 
bone. 
The 
teeth 
are 
beautifully 
preserved 
and 
almost 
unworn. 
4'hey 
are 
huge,even 
whencompared 
withthoseofan 
Alaskan 
brown 
bear. 


The 
central 
cusp 
of 
P 
4 
is 
high. 
The 
anterior 
cusp 
is 
low 
but 
trenchant. 
The 
posterior 
cusp 
is 
poorlydeveloped. 
There 
is 
a 
low 
cingulum 
about 
the 
posterior 
third 
of 
the 
tooth, 
posterior 
to 
the 
central 
cusp. 
The 
enamel 
has 
many 
faint 
vertical 
striations, 
most 
pronounced 
on 
the 
labial 
side 
of 
the 
tooth. 
M, 
is 
elongate. 
The 
trigonid 
is 
narrower 
transversely 
than 
the 
talonid. 
There 
is 
a 
low 
cingulum 
on 
the 
lingual 
side 
of 
the 
paraconid. 
The 
carnassial 
notch 
is 
distinct 
but 
narrow, 
with 
the 
posterior 
edgeof 
the 
blade 
of 
the 
paraconid 
and 
the 
anterior 
edgeof 
the 
blade 
of 
the 
protoconid 
in 
contact 
for 
a 
vertical 
height 
of 
approximately 
5 
mm. 
The 
mctaconid 
is 
a 
small, 
rounded 
cone 
on 
the 
lingual 
side 
of 
the 
tooth, 
posterior 
to 
the 
protoconid. 
The 
entoconid 


is 
smaller 
and 
posterior 
to 
the 
metaconid. 
The 
postentoconld 
cusp 
is 
still 
smaller 
and 
posterior 
to 
the 
entoconid. 
The 
three 
cusps, 
metaconid, 
entoconid 
and 
post-entoconid, 
form 
an 
evenly 
stepped 
series 
on 
the 
lingual 
side 
of 
the 
tooth. 
The 
hypoconid 
is 
a 
low, 
elongated 
cusp, 
as 
long 
as 
but 
lower 
than 
the 
entoconid 
and 
post-entoconid 
together, 
on 
the 
opposite 
side 
of 
the 
tooth. 
There 
is 
a 
low 
intra-protoconid-
hypoconid 
cusp 
opposite 
the 
metaconid. 
There 
is 
a 
low, 
faint 
cingulum 
on 
the 
labial 
side 
of 
the 
talonid. 
The 
enamel 
is 
finely 
wrinkled 
and 
vertically 
striated, 
especially 
on 
the 
labial 
side. 


The 
Mo 
is 
somewhat 
triangular, 
w'ith 
the 
trigonid 
much 
broader 
than 
the 
talonid. 
In 
American 
Pleistocene 
and 
Recent 
bears 
( 
Ursus, 
Tremarctos, 
Arctodus, 
etc.) 
this 
tooth 
is 
rectangular 
or 
hourglass 
shaped. 
The 
metaconid 
is 
very 
large, 
largerand 
higher 
than 
the 
protoconid. 
The 
paraconid 
is 
obscure 
or 
absent. 
Inward-reaching 
crests 
from 
metaconid 
and 
protoconid 
meet 
in 
a 
low 
“V” 
to 
form 
a 
loph 
across 
the 
tooth 
and 
separate 
the 
occlusal 
surface 
into 
two 
distinct 
basins. 
The 
anterior 
basin 
is 
closed 
anteriorly 
by 
a 
ridge-like 
cingulum.
The 
transverse 
loph 
on 
the 
M 
2 
of 
the 
Hemphill 
bear 
is 
quite 
similar 
to 
that 
of 
Arctodus 
(Kurtcn, 
1963).
The 
entoconid 
is 
small 
and 
the 
post-entoconid 
is 
equal 
in 
size. 
The 
hypoconid 
is 
larger 
and 
higher.
A 
narrow 
ridge 
or 
cingulum 
connects 
the 
hypoconidwith 
the 
post-entoconid 
around 
the 
posterior 
border 
of 
the 
tooth 
to 
close 
the 
basin 
of 
the 
talonid. 
The 
enamel 
of 
the 
tooth 
is 
finely 
wrinkled 
and 
vertically 
striated. 


The 
crown 
of 
M 
3 
is 
a 
rounded 
triangle 
in 
occlusal 
view. 
In 
the 
specimen, 
the 
tooth 
is 
set 
so 
that 
its 
occlusal 
surface 
is 
parallel 
with 
the 
occlusal 
surface 
of 
M 
2, 
but 
the 
tooth 
is 
so 
far 
back 
in 
the 
jaw 
that 
all 
but 
its 
anterior 
edge 
was 
buried 
in 
the

Fig. 
3. 
Lower 
jaw 
of 
Indarctos 
oregonensis 
with 
P4—M3 
(TMM 
41261—1). 
X 
.50 
natural 
size. 



rising 
curve 
of 
the 
ascending 
ramus 
of 
the 
mandible. 
Considerable 
bone 
had 
to 
be 
ground 
away 
to 
expose 
the 
crown. 
The 
root 
appears 
to 
be 
developed. 
Either 
the 
tooth 
was 
not 
completely 
erupted 
or 
it 
was 
“impacted’' 
in 
the 
manner 
of 
some 
human 


lower 
thirdmolars. 


The 
protoconid 
is 
a 
low 
swelling 
on 
the 
labial 
side 
of 
the 
tooth. 
The 
remainder 
of 
the 
crown 
consists 
of 
a 
wrinkled 
basin 
surrounded 
by 
a 
narrow 
cingulum 
or 
ridge 
that 
is 
continuous 
with 
the 
external 
(labial) 
edge 
of 
the 
protoconid.

The 
isolated 
(TMM4I26I2) 
is 
also 
from 
the 
right 
side. 
It 
is 
smaller 
than 
the 
of 
EMM 
41261-1, 
and 
the 
talonid 
basin 
is 
deeper. 
The 
roots 
had 
not 
developed. 
The 
lingual 
border 
of 
the 
tooth, 
between 
the 
paraconid 
and 
protoconid, 
is 
more 
notched 
than 
in 
TMM41261-1, 
so 
that 
the 
trigonid 
and 
talonid 
are 
not 
so 
distinctly 
set 
off 
from 
one 
another. 
In 
other 
respects 
the 
tooth 
is 
like 
TMM41261-1. 
Metaconid, 
cntoconid 
and 
postentocomd 
are 
similarly 
stepped.

Comparison 
of 
the 
present 
material 
with 
In


darctos 
oregonensis 
Merriam, 
Stock 
and 
Moody

(1916), 
and 
Hyaenarctos 
gregoryi 
Frick 
(1921) 
is 


not 
possible, 
for 
those 
species 
were 
based 
on 
upper 


teeth. 
Indarctos 
nevadensis, 
however, 
is 
based 
on 
a 


lower 
jaw 
with 
unworn 
canine 
and 
P,-Mi. 
It 
is 


directly 
comparable 
with 
the 
Coffee 
Ranch 
teeth, 


but 
only 
P, 
and 
M, 
are 
present 
in 
both 
specimens. 


Theteethinthe 
jawfromNevadaareslightlysmall


er 
but 
are 
so 
similar 
in 
shape 
and 
details 
of 
tooth 


structure, 
including 
the 
stepped 
mctaconid-ento


conid-postcntoconid 
on 
Mi, 
that 
identity 
seems 


probable. 
Individual 
variationsof 
greater 
extent 
are 


seen 
in 
the 
P, 
and 
M, 
of 
modern 
Ursus. 
Whether 


I. 
nevadensis 
is 
actually 
a 
species 
d’stinct 
from 
I. 
oregonensis 
remains 
to 
be 
determined 
when 
lower 
jaws 
of 
the 
latter 
are 
discovered. 
For 
comparison 
of 


I. 
nevadensis 
with 
old-world 
species 
see 
Macdonald 
(1959).
Agriotherium 
schneideri 
Sellards, 
described 
from 
the 
Pliocene 
Bone 
Valley 
formation 
of 
Florida, 
is 
a 
bear 
of 
approximately 
the 
same 
size 
as 
the 
specimen 
from 
the 
Coffee 
Ranch. 
The 
holotype 
is 
a 
right 
lower 
jaw 
with 
P 
4-M2 
and 
the 
alveolus 
for 
M 
3. 
The 
teeth 
are 
worn 
and 
it 
is 
difficult 
to 
determine 
whether 
the 
cusps 
of 
the 
talonid 
of 
M 
4 
were 
“stepped” 
like 
thoseof 
theTexas 
bear. 


The 
Mo 
of 
the 
holotype 
jaw 
of 
A. 
schneideri 
differs 
strikingly 
from 
the 
Coffee 
Ranch 
specimen 
in 
that 
talonid 
and 
trigonid 
are 
approximately 
equal 
in 
breadth 
and 
the 
tooth 
is 
almost 
rectangular. 
In 


Fig. 
4. 
Lower 
carnassial 
of 
Indarctos 
oregonensis 
(TMM 
41261—2) 
in 
lingual 
and 
occlusal 
views. 
X 
1.5 
natural 
size. 


the 
bear 
from 
Texas 
the 
M 
2 
is 
more 
triangular. 
Maximum 
breadth 
is 
near 
the 
base 
of 
the 
enamel, 
and 
not 
affected 
by 
wear 
until 
extreme 
old 
age. 


Savage 
(1941) 
figures 
and 
give 
measurements 
of 
theteethofalower 
jawofabearfromthePliocene 
Optima 
local 
fauna 
of 
Oklahoma. 
The 
measurements 
of 
the 
Mo 
are 
similar 
to 
those 
of 
the 
Coffee 


Ranch 
specimen. 
Anteroposterior 
and 


transverse 


diameters 
of 
the 
Mo’s 
of 
the 
specimens 
from 
Texas, 
Oklahoma 
and 
Florida 
are 
respectively; 
34.2 
x 
25.5; 
32.5x25.1; 
26x20. 


Donald 
E. 
Savage 
has 
suggested 
(personal 
communication) 
that 
all 
of 
the 
American 
Pliocene 
bears 
may 
belong 
to 
one 
genus, 
namely 
Agriotheriurn. 
Certainly 
the 
group 
is 
in 
need 
of 
revision. 
Pending 
such 
revision 
it 
seems 
best 
to 
refer 
the 
Texas 
bear 
to 
Indarctos 
oregonensis, 
the 
species 
it 
most 
closely 
resembles 
among 
described 
forms. 
Savage’s 
specimen 
from 
Oklahoma 
would 
appear 
to 
belong 
to 
the 
same 
species. 
Agriotherium 
schneideri 
appears 
to 
be 
at 
least 
specifically 
distinct. 


Pliotaxidea 
cf. 
P. 
nevadensis 
(Butterworth) 


A 
small 
badger 
is 
represented 
by 
two 
lower 
jaws.

MU 
1334 
is 
a 
right 
mandible 
lacking 
the 
ascending 


ramus 
and 
all 
teeth 
but 
M,. 
The 
alveoli 
and 
roots 


of 
the 
other 
cheek 
teeth 
are 
present. 
The 
molar 
is 


well 
worn 
and 
closely 
resembles 
the 
tooth 
from 
Ok


lahoma 
figured 
by 
Hesse 
(1936:60) 
and 
later 
re



ferredtoPliotaxidea 
nevadensisbyHall(1944:15).
The 
tooth 
is 
10.3 
mm. 
long 
and 
5.5 
mm. 
wide. 


MU8066isaleftmandibularramus,complete 
except 
for 
the 
alveoli 
of 
the 
incisors. 
Only 
P2 
and 
P:i 
are 
present. 
The 
roots 
of 
most 
of 
the 
other 
teeth 
are 
visible 
in 
their 
alveoli. 
The 
distance 
from 
the 
anterior 
edge 
of 
the 
canine 
alveolus 
to 
the 
posterior 
edgeof 
the 
articular 
condyle 
is 
48.6 
mm. 
The 
distance 
from 
the 
anterior 
edge 
of 
the 
canine 
alveolus 
to 
the 
posterior 
edge 
of 
the 
alveolus 
of 
M 
2 
is 
32.1 
mm. 
The 
maximum 
depth 
of 
the 
jaw 
beneath 
M 
4 
is 
5.7 
mm. 


Hall 
(1944) 
has 
shown 
that 
the 
Pliocene 
old-
world 
Parataxidea 
Zdansky 
is 
distinct 
from 
the 
Pliocene 
American 
Pliotaxidea, 
and 
that 
the 
former 
is 


close 
to 
the 
ancestry 
of 
the 
modern 
old-world 
Meles 
while 
the 
latter 
may 
be 
the 
direct 
ancestor 
of 
the 
modern 
American 
Taxidea. 
Long 
(1965) 
presentsevidence 
that 
Taxidea 
and 
Meles 
should 
be 
placedin 
separate 
sub-families. 


Plesiogulo 
marshalli 
(Martin)
(Fig. 
5) 


This 
wolverine-like 
mustelid 
is 
represented 
by 
the 
posterior 
half 
of 
a 
lower 
first 
molar 
(MU5132) 
and 


Fig. 
5. 
Plesiogulo 
rnarshalli 
(TMM 
41261—12), 
upper 
Ml 
in 
occlusal 
view. 
X 
1.5 
natural 
size. 


the 
unworn 
crown 
of 
an 
upper 
first 
molar 
(TMM-
41261-12). 
The 
fragmentary 
lower 
tooth 
is 
almost 
identical 
in 
size 
and 
details 
of 
structure 
to 
the 
tooth 


in 
the 
photograph 
of 
the 
holotype 
lower 
jaw(Martin, 
1928). 
The 
upper 
tooth 
lacks 
roots, 
but 
the 
crown 
is 
perfect. 
Its 
greatest 
dimensions 
are 


19.5 
X 
16.3 
mm. 
The 
least 
breadth 
across 
the 
median 
constriction 
is 
9.3 
mm. 
The 
enamel, 
even 


on 
the 
strong 
cingulum 
and 
low 
cusps, 
is 
delicatelywrinkled 
and 
denticulate. 


A 
skull 
without 
teeth, 
MUBO6B, 
is 
also 
referable 
to 
this 
species. 
It 
is 
crushed, 
and 
has 
lost 
both 
zygomatic 
arches 
and 
other 
important 
details. 
It 
shows, 
however, 
thattheskullofP. 
rnarshalliwassimilar 
in 
shape 
and 
in 
general 
construction 
to 
the 
skull 
of 
the 
modern 
wolverine: 
broad 
through 
the 
rostrum 
and 
across 
the 
interorbital 
region 
with 
a 
large 
infra-
orbital 
foramen, 
and 
narrow, 
cylindrical 
braincase. 


Pseudaelurus 
hibbardi, 
new 
species(Fig. 
6) 


Holotype.—Left 
lower 
jaw 
(TMM41261-3),
lacking 
canine, 
frontof 
caninealveolus 
andanterior 
tipofjawwithincisors, 
andalloframusposteriorto 
Mx.Mostofsymphysis,P3,P,andM, 
present.

Locality.—Coffee 
Ranch 
quarry, 
Coffee 
Ranch 
local 
fauna, 
from 
gray 
sand 
bed 
six 
feet 
beneath 
volcanic 
ash 
layer, 
middle 
Pliocene, 
HemphillCounty, 
Texas. 


Fauna.—HcmphiIlian. 


Diagnosis.—Size 
of 
large 
puma. 
Carnassial 
notch 
open 
in 
upper 
P 
4 
and 
lower 
M,; 
Pt 
absent; 
vestigial 
P 
2 
possibly 
present; 
P:i 
and 
P4 
small 
and 
thin 
but 
Mt 
relatively 
very 
large. 
Ramus 
straight, 
deepand 
stout, 
especially 
anteriorly. 
Symphysis 
strong,
with 
angle 
between 
ventral 
and 
anterior 
surfaces 
of 
ramus 
acute. 
Metaconid 
on 
lower 
M, 
of 
moderate 
size. 


Referred 
material.—Right 
lower 
jaw 
fragment(TMM41261-5) 
with 
part 
of 
canine 
alveolus,
alveolus 
of 
P:! 
a 
well-preserved 
Pt 
, 
and 
part 
of 
the

, 


Fig. 
6. 
Lower 
jaw 
of 
holotype 
of 
Pseudaelurus 
hibbardi 
with 
P3—Ml 
(TMM 
41261—3), 
in 
labial 
view 
X 
1.0 
natural 
size. 



alveolus 
of 
M,; 
isolated 
upper 
carnassial 
(MU 


6389) 
; 
scapula 
(TMM41261—15) 
; 
humerus 
(TMM41261-16) 
; 
radius 
(TMM41261-17) 
; 
astragalus 
(TMM41261-19) 
; 
two 
calcanea 
(TMM 
41261-21,22) 
; 
navicular 
(MU6389). 


DESCRIPTION.—The 
ramus 
of 
the 
lower 
jaw 
is 
thick 
and 
deep, 
especially 
anteriorly. 
The 
alveolus 
of 
the 
canine 
is 
very 
large, 
of 
round-oval 
shape, 
and 
the 
jaw 
is 
almost 
bulbous 
about 
the 
canine 
alveolus. 
The 
angle 
between 
the 
ventral 
border 
of 
the 
jawand 
the 
front 
of 
the 
jaw 
is 
acute. 
The 
diastema 
is 
short, 
and 
contains 
a 
tiny 
pit 
that 
may 
be 
an 
alveolus 
for 
a 
vestigial 
P2 
. 
The 
lower 
P3 
has 
a 
posteriorbasal 
cusp 
but 
only 
a 
slight 
swelling 
in 
place 
of 
an 
anterior 
basal 
cusp. 
There 
is 
no 
posterior 
cingular 
cusp. 
Lower 
P 
3 
and 
P 
4 
are 
relatively 
small 
and 
thin, 
and 
closely 
placed 
in 
the 
tooth 
row. 
Lower 
P, 
has 
anterior 
and 
posterior 
basal 
cusps. 
Lower 
Ma 
large 
and 
heavy 
with 
broadly 
open 
carnassial 
notch 


and 
moderately 
well-developed 
metacone.Referred 
upper 
P4 
with 
broadly 
open 
carnassial 
notch. 
Referred 
postcranial 
elements 
exactly 
comparable 
to 
bones 
oflargepuma.

COMPARISONS.—Hibbard 
(1934) 
has 
described 
Adelphailurus 
kansensis 
from 
the 
Edson 
Beds, 
Sherman 
County, 
Kansas, 
and 
Pratifelis 
martinifrom 
theLost 
Quarry,WallaceCounty,Kansas; 


both 
species 
are 
from 
deposits 
of 
Hemphillian 
age. 


Adelphailurus 
kansensis, 
as 
noted 
by 
Hibbard 


(1934: 
246) 
is 
not 
closely 
related 
to 
Pseudaelurus. 


It 
is, 
instead, 
very 
close 
to 
Panthera. 
A 
cast 
of 
the 


holotype 
was 
compared 
with 
the 
skull 
of 
an 
African 


leopard, 
and 
the 
resemblances 
are 
numerous. 
Dif


ferences 
include 
a 
shorter 
diastema, 
more 
laterally 


compressed 
canine, 
very 
slightly 
more 
open 
carnas


sial 
notch, 
and 
some 
minor 
details 
of 
the 
teeth. 


Adelphailurus 
may 
well 
be 
the 
direct 
ancestor 
of 
the 


leopard, 
and 
the 
genus 
is 
not 
strongly 
differentiated 


fromPanthera. 


Differences 
between 
Adelphailurus 
kansensis 
and 


Pseudaelurus 
hibbardi 
are 
numerous. 
In 
the 
upper 


P 
4, 
the 
carnassial 
notch 
of 
P. 
hibbardi 
is 
broadly 


open 
rather 
than 
almost 
closed. 
The 
diastema 
of 


the 
upper 
jaw 
of 
Adelphailurus 
is 
very 
short, 
and 


when 
the 
lower 
jaw 
of 
a 
leopard 
is 
placed 
in 
occlu


sion 
with 
it, 
the 
canine 
of 
the 
leopard 
will 
not 
enter 


the 
diastema 
of 
the 
maxillary. 
With 
canine 
re


moved, 
the 
leopard 
jaw 
articulates 
fairly 
well 
with 


the 
Adelphailurus 
maxillary, 
and 
the 
teeth 
almost 


occlude. 
The 
lower 
jaw 
of 
Adelphailurus 
was 
al


most 
certainly 
similar 
to 
the 
leopard 
lower 
jaw 
but 


lighter 
in 
build, 
and 
the 
lower 
canine 
must 
have 


been 
much 
smaller. 
In 
Pseudaelurus 
hibbardi 
the 
canine 
was 
enormous 
and 
the 
lower 
jaw 
very 
heavy.
When 
placed 
with 
the 
maxillary 
of 
Adelphailurus,
the 
teeth 
do 
not 
occlude 
and 
the 
lower 
canine 
is 
almost 
exactly 
below 
the 
upper 
canine. 


The 
holotype 
of 
Pratifelis 
martini 
is 
a 
left 
lower 
jaw, 
and 
is 
thus 
comparable 
with 
the 
type 
of 
Pseudaelurus 
hibbardi. 
In 
Pratifelis 
the 
lower 
jaw 
is 
slender, 
light 
and 
curved. 
In 
P. 
hibbardi 
the 
jaw 
is 
straight, 
heavy 
and 
deep. 
The 
lower 
canine 
of 
Pratifelis 
was 
small, 
in 
P. 
hibbardi 
it 
was 
very 
large. 
4 
he 
lower 
P:! 
and 
P, 
of 
Pratifelis 
are 
quite 
similar 
to 
those 
of 
P. 
hibbardi 
except 
that 
the 
P3 
of 
Pratifelis 
has 
a 
posterior 
cusp 
and 
a 
posterior 
cingular 
cusp.
In 
P. 
hibbardi 
there 
is 
only 
a 
posterior 
basal 
cusp.
Both 
P:t 
and 
P, 
of 
Pratifelis 
are 
thinner 
than 
the 
premolars 
of 
P. 
hibbardi. 
The 
Mx 
of 
P. 
hibbardi, 
on 
the 
other 
hand, 
is 
both 
longer 
and 
wider 
than 
the 
molar 
of 
Pratifelis. 
A 
striking 
difference 
is 
the 
broadly 
open 
carnassial 
notch 
of 
P. 
hibbardi. 
Pratifelis 
was 
a 
highly 
aberrant 
cat. 
It 
does 
not 
belong 
in 
the 
genus 
Pseudaelurusand 
shows 
little 
similarity

, 


to 
P. 
hibbardi. 
Both 
were 
cats 
approximately 
the 
size 
of 
the 
puma.

Pseudaelurus 
thinobates 
Macdonald, 
known 
from 
Clarendonian 
and 
early 
Hemphillian 
faunas 
of 
California, 
Texas 
and 
Oklahoma 
(Kitts, 
1958),
resembles 
P. 
hibbardi 
in 
several 
respects, 
includingthe 
heavy 
symphysis, 
large, 
deep 
ramus, 
swollen 
anteriorly 
for 
the 
enormous 
canine, 
and 
the 
steepangle 
of 
the 
anterior 
face 
of 
the 
jaw.The 
carnassial 
notch, 
in 
P. 
thinobates, 
is 
broadly 
open 
as 
it 
is 
in 
P. 
hibbardi. 
P. 
thinobates, 
however, 
is 
much 
largerthan 
P. 
hibbardi, 
being 
approximately 
the 
size 
of 
an 
African 
lion 
rather 
than 
the 
size 
of 
a 
large 
puma.

Kitts 
(1958) 
has 
placed 
P. 
thinobates 
in 
a 
new 


genus, 
Nimravides. 
The 
characters 
of 
P. 
hibbardi 


and 
P. 
pedionomus 
Macdonald 
bridge 
the 
gap 
be


tween 
P. 
thinobatesand 
typical 
Pseudaelurus, 
such 


as 
P. 
intrepidus 
(Leidy). 
Nimravides 
is 
best 
con


sidered 
a 
subgenus 
until 
the 
late 
Tertiary 
fclids 
are 


revised. 


Pseudaelurus 
pedionomus 
Macdonald 
was 
de


scribed 
from 
the 
lower 
Pliocene 
of 
Nebraska. 


Among 
described 
fossil 
cats, 
this 
form 
is 
most 
simi


lar 
to 
P. 
hibbardi. 
In 
both 
the 
ramus 
is 
deep 
and 


heavy, 
and 
the 
canine 
large. 
The 
two 
are 
of 
similar 


size. 
In 
P. 
hibbardi 
the 
jaw 
is 
more 
swollen 
anter


iorly, 
and 
the 
canine 
alveolus 
is 
larger. 
P 
t 
and 
P2 


are 
present 
in 
P. 
pedionomus, 
but 
only 
a 
tiny 
P2 


may 
occur 
in 
P. 
hibbardi. 
The 
diastema 
(C-P3) 
is 


much 
less 
in 
P. 
hibbardi 
than 
in 
P. 
pedionomus. 



The 
premolars 
of 
P. 
pedionomus 
are 
longer 
and 
wider 
than 
those 
of 
P. 
hibbardiand 
the 
molar 
is

, 


slightly 
longer 
and 
wider. 
The 
metaconid 
is 
better 
developed 
in 
P. 
hibbardi 
than 
in 
P. 
pedionomus. 
In 


P. 
pedionomus 
there 
is 
a 
distinct 
gap 
between 
P:( 
and 
P 
4, 
but 
no 
such 
gap 
occurs 
in 
P. 
hibbardi. 
There 
are 
two 
mental 
foramina 
in 
P. 
pedionomus, 
one 
beneath 
the 
center 
of 
the 
diastema 
and 
one 
beneath 
the 
anterior 
root 
of 
P3 
. 
In 
P. 
hibbardi 
there 
is 
a 
single 
mental 
foramenbeneath 
the 
anterior 
root 
of 
P3. 


Some 
of 
the 
differences 
listed 
may 
be 
subject 
to 
individual 
and 
sexual 
variation. 
The 
enormous 
canine 
alveolus 
and 
swollen 
anterior 
end 
of 
the 
ramus 
in 
P. 
hibbardi 
are 
present 
in 
two 
different 
specimens, 
however, 
and 
are 
probably 
valid 
speciescharacters. 
They 
readily 
serve 
to 
separate 
P. 
hibbardi 
from 
P. 
pedionomus. 
P. 
pedionomus, 
P. 
thinobates 
and 
P. 
hibbardi 
seem 
to 
represent 
a 
line 
of 
large, 
heavy-jawed 
cats 
(subgenus 
Nimravides)
with 
P. 
hibbardi 
the 
last 
known 
member 
of 
the 


group.

The 
postcranial 
elements 
referred 
to 
P. 
hibbardi 
are 
splendidly 
preserved 
and 
almost 
complete. 
The 
elements 
are 
so 
similar 
to 
the 
same 
bones 
of 
a 
large 


puma, 
even 
to 
details 
of 
shape 
of 
muscle 
scars 
and 
curvature 
of 
scapular 
blade, 
that 
one 
may 
suppose 
that 
the 
body 
shape 
was 
very 
much 
like 
that 
of 
the 
puma. 
Macdonald 
(1948a) 
thought 
that 
P. 
pedionomus 
was 
lynxlike, 
with 
some 
pumalike 
charac


ters. 
1 
would 
judge 
P. 
hibbardi 
to 
have 
been 
quite 
puma-like 
in 
its 
habits. 


Machairodus 
(Heterofelis) 
catocopis 
Cope(Plate 
1 
: 
Figs. 
7-11) 


Remains 
of 
a 
large 
saber-toothed 
cat 
from 
the 
ColTee 
Ranch 
quarry 
all 
seem 
to 
pertain 
to 
a 
singlespecies. 
Materials 
include 
a 
largely 
complete 
skeleton, 
several 
upper 
and 
lower 
jaws, 
and 
numerous 
isolated 
teeth 
and 
bones. 
Burt 
(1931) 
has 
brieflydescribed 
and 
figured 
a 
number 
of 
bones 
of 
Machairodus 
catocopis 
from 
the 
Coffee 
Ranch. 
The 
new 
material 
is 
much 
superior 
to 
that 
available 
to 
Burt, 
and 
it 
is 
possible 
to 
correct 
some 
errors 
in 
his 
work 
and 
greatly 
extend 
the 
knowledge 
of 
this 
interesting 


cat. 


DESCRIPTION.—The 
most 
important 
specimen 
is 
the 
skeleton 
TMM41261-8. 
It 
is 
of 
an 
old 
saber-tooth, 
possibly 
a 
female, 
tor 
some 
of 
the 
bones 


Fig. 
7. 
Restoration 
of 
skull 
of 
Machairodus 
catocopis 
based 
on 
the 
basicranium 
and 
right 
maxilla 
of 
TMM 
41261—8, 
and 
on 
associated 
premaxilla 
probably 
belonging 
to 
the 
same 
individual. 
Approx 
X 
.50 
natural 
size. 
The 
length 
of 
the 
gap 
in 
the 
center 
of 
the 
skull 
was 
determined 
by 
articulating 
the 
lower 
jaw 
of 
TMM 
41261—8 
in 
the 
glenoid 
fossa 
and 
placing 
upper 
and 
lower 
dentitions 
in 
occlusion. 



are 
only 
of 
moderate 
size 
when 
compared 
with 
other 
specimens 
from 
the 
quarry. 
The 
bones 
were 
found 
articulated 
or 
closely 
associated, 
and 
much 
of 
the 
skeleton 
was 
removed 
in 
a 
single 
large 
block 
of 
matrix 
(Plate 
1). 
The 
following 
descriptions, 
unless 
otherwise 
stated, 
are 
based 
on 
this 
specimen.

In 
the 
following 
accounts, 
comparisons 
are 
primarily 
with 
the 
bones 
of 
Smilodon 
californicus 
Bovard, 
the 
saber-toothfrom 
the 
Rancho 
La 
Brea, 
of 
California. 
Smilodon 
is 
not 
particularly 
close 
to 
Machairodus 
catocopis 
in 
morphological 
details, 
but 
it 
is 
the 
best-known 
of 
the 
saber-toothed 
cats. 
Nearly 
all 
of 
the 
bones 
of 
the 
skeleton 
have 
been 
figured 
in 
the 
classic 
work 
of 
Merriam 
and 
Stock 
(1932), 
and 
the 
tables 
in 
that 
publication 
show 
the 
expected 
range 
of 
variation 
in 
measurements 
of 
the 
skeleton 
elements. 
Bones 
from 
the 
Rancho 
La 
Brea 
are 
preserved 
in 
many 
museum 
collections, 
and 
most 
of 
the 
bones 
of 
the 
skeleton 
of 
Smilodon 
californicus 
were 
available 
for 
direct 
comparison.

The 
skeleton 
of 
Homotherium 
serum 
(Cope),
from 
the 
late 
Pleistocene 
deposits 
in 
Friesenhahn 
Cave, 
Texas, 
has 
been 
described 
in 
the 
importantwork 
by 
Meade 
(1961). 
(For 
use 
of 
Homotherium 
Fabrini 
rather 
than 
Dinohastis 
Cope, 
see 
Churcher, 
1968.) 
Machairodus 
catocopis 
resembles 
Homotherium 
serum 
more 
closely 
than 
it 
does 
Smilodon. 
However, 
few 
bones 
of 
Homotherium 
were 
available 
for 
direct 
comparison 
with 
Machairodus, 
and 
most 
of 
the 
stated 
differences 
between 
Machairodus 
catocopis 
and 
Homotherium 
serum 
are 
based 
on 


the 
figures 
and 
measurements 
given 
by 
Meade. 


Skull.—The 
skull 
had 
been 
stepped 
on, 
presum


ably 
by 
a 
large 
mammal 
such 
as 
a 
rhinoceros, 
and 


the 
anterior 
left 
portion 
destroyed. 
The 
top 
of 
the 


braincase, 
including 
all 
but 
the 
posterior 
inch 
of 
the 


sagittal 
crest, 
is 
missing. 
The 
glenoid 
processes 
were 


found 
in 
the 
matrix 
beneath 
their 
natural 
positions.

The 
right 
maxillary 
with 
canine 
and 
carnassial, 
and 


premaxillary 
with 
P3 
and 
alveoli 
of 
I 
1 
and 
12,I2 
were

, 


found 
nearby. 
The 
occipital 
and 
basicranial 
regions 


are 
almost 
completeand 
wellpreserved.

There 
is 
no 
point 
of 
contact 
between 
the 
maxil


lary 
and 
basicranium. 
In 
the 
restoration 
of 
the 
skull 


(Fig. 
7), 
the 
distance 
represented 
by 
the 
missing

bone 
was 
approximated 
by 
placing 
the 
right 
lower 


jaw 
ramus 
in 
occlusion 
with 
the 
upper 
carnassial, 


and 
then 
fitting 
the 
articular 
condyle 
of 
the 
man


dible 
in 
the 
glenoid 
fossa 
of 
the 
basicranium. 


The 
skull 
is 
long 
and 
slender, 
like 
the 
skull 
of 


Homotherium.The 
occipitalcrestprojects 
farback


ward, 
as 
it 
does 
in 
Homotherium, 
and 
is 
constricted 


anteriorly, 
behind 
the 
braincase, 
more 
than 
in 
Smilodon. 
Only 
the 
posterior 
part 
of 
the 
sagittal 
crest 
remains, 
but 
it 
is 
prominent 
as 
in 
other 
large 


cats. 
The 
edges 
of 
the 
occipital 
crest 
have 
been 
damaged 
but 
it 
is 
apparent 
that 
the 
crest, 
in 
posteroventral 
view, 
is 
broad 
and 
rounded, 
not 
triangular. 
The 
infraorbital 
foramen 
is 
large 
and 
oval, 
as 
in 
Homotherium.The 
premaxillary 
is 
strong 
and 
juts 
forward, 
more 
as 
In 
Homotherium 
than 
as 
in 
Smilodon. 


The 
braincase 
appears 
to 
have 
been 
more 
expanded, 
and 
rounded 
at 
the 
sides, 
than 
inSmilodon. 
The 
parts 
of 
the 
sides 
that 
remain 
are 
almost 
vertical, 
even 
slightly 
inclined 
outward 
above. 
In 
Smilodon 
the 
sides 
converge 
above, 
so 
that 
the 
braincase 


appears 
almost 
triangular 
in 
section. 


The 
basicranium 
is 
well 
preserved 
and 
important.
The 
ventral 
surfaces 
of 
the 
glenoids 
are 
separated 
from 
the 
top 
of 
the 
mastoid 
processes 
by 
a 
space 
of 
10 
mm. 
In 
a 
Smilodon 
of 
comparable 
size 
the 
separation 
is 
less, 
about 
6 
mm. 


The 
audital 
bulla 
of 
the 
left 
side 
is 
complete, 
and 
that 
of 
the 
right 
side 
partly 
so. 
The 
bullae 
are 
relatively 
larger 
than 
in 
any 
available 
specimen 
of 
Smilodon, 
but 
their 
size 
is 
known 
to 
vary 
widely 
in 
Smilodon. 


The 
opening 
to 
the 
eustachian 
canal 
is 
wide 
(12mm.) 
but 
narrows 
swiftly 
to 
a 
narrow 
slit, 
partiallydivided 
internally 
by 
a 
sharp 
ridge 
on 
the 
posteriorsurface. 


The 
external 
auditory 
meatus 
is 
almost 
round 


and 
is 
situated, 
as 
in 
Smilodon 
but 
not 
as 
in 
the 
true 


cats 
(Panthera, 
Felis), 
in 
the 
deep 
cavity 
between 


the 
zygomatic 
arch 
above 
and 
the 
mastoid 
process

below. 


The 
jugular 
foramen 
is 
in 
a 
triangular 
depression 


at 
the 
posterointernal 
corner 
of 
the 
audital 
bulla. 


This 
area 
is 
usually 
more 
rounded 
in 
Smilodon. 


The 
foramen 
ovale 
is 
a 
deep, 
round 
opening, 
6 


mm. 
wide, 
separated 
from 
the 
eustachian 
canal 
by 


abarofbone 
3.6mm.wide.Theforamenrotundum 


is 
similar 
to 
the 
foramen 
ovale 
in 
shape 
and 
size 
and 


is 
located 
10.7 
mm. 
anterior 
to 
it. 
There 
is 
a 
groove

in 
the 
alisphenoid 
between 
the 
two 
foramina, 
and 


the 
groove 
is 
partially 
spanned 
by 
a 
bar 
of 
bone. 


The 
bar 
of 
bone 
is 
present 
on 
both 
sides, 
and 
is 
thus 


a 
short 
alisphenoid 
canal. 
The 
canal 
is 
not 
as 
well 


developed 
as 
in 
dogs. 
Mawby 
(1965) 
has 
men


tioned 
the 
existence 
of 
this 
canal 
in 
M. 
catocopis. 
It 


appears 
to 
be 
absent 
in 
other 
sabertooths, 
and 
in 


Felis 
and 
Panthera 
as 
well. 
It 
may 
be 
simply 
a 
per


sistent 
primitive 
character 
in 
M. 
catocopis. 



Superior 
dentition.—The 
specimen 
best 
showingthe 
upper 
dentition 
is 
TMM41261-10, 
the 
upperjaw 
of 
a 
young-adult 
sabertooth 
with 
the 
alveolus 
of 
ll 
broken 
away, 
both 
P 
and 
P 
present 
canine 
alveolus 
and 
P3 
and 
P 
1 
present 
(Fig. 
8). 
An 
associatedcanineprobablybelongstothesame 
jaw.The 
following 
description 
is 
based 
primarily 
on 
this 
specimen, 
but 
is 
augmented 
by 
TMM41261-8 
and 
other 
specimenswhenhelpful.

I 
1is 
not 
preserved, 
but 
the 
alveoli 
in 
two 
premaxillaries 
suggest 
that 
it 
was 
a 
bit 
smaller 
and 
more 
compressed 
than 
12.I2. 
I 
2 
is 
a 
strong 
tooth 
with 
a 
heavy 
internal 
cingulum 
and 
large 
basal 
cusps 
at 
anterior 
and 
posterior 
margins. 
Itis 
a 
much 
strongerand 
more 
triangular 
tooth 
than 
the 
I 
2 
of 
Smilodon. 
The 
I 
2 
of 
Homotherium 
appears 
to 
be 
similarlylarge 
and 
triangular 
but 
has 
a 
basal 
cusp 
only 
on 
the 
posterior 
edge. 
This 
may 
be 
a 
minor 
and 
variable 
feature. 


I 
3 
is 
a 
large, 
strong 
triangular 
tooth, 
as 
it 
is 
in 
Homotherium. 
It 
has 
a 
strong 
basal 
cusp 
on 
the 
anterior 
margin, 
but 
posteriorly 
there 
is 
only 
a 
heavycingulum 
and 
a 
low 
swelling 
in 
place 
of 
a 
basal 
cusp. 
In 
Homotherium 
there 
is 
a 
strong 
posterior 


cusp. 
The 
anterior 
edge 
docs 
not 
seem 
to 
be 
serrate, 
as 
it 
is 
in 
Homotherium. 
The 
posterior 
edge 
is 
serrate 
for 
less 
than 
half 
of 
its 
length. 
Resemblance 
to 
Homotherium 
is 
strong; 
differences 
are 
minor 
and 
probablysubject 
toindividualvariation. 


The 
upper 
canine 
is 
compressed, 
relatively 
as 
much 
as 
in 
Smilodon. 
It 
is 
serrate 
anteriorly 
and 
posteriorly 
for 
the 
entire 
length 
of 
the 
enamel. 
It 
is 
relatively 
and 
actually 
a 
much 
smaller 
tooth 
than 
it 
is 
in 
Smilodonbut 
is 
a 
bit 
longer 
than 
the 
canine 
of 
Homotherium., 


Burt 
(1931:264) 
described 
the 
alveolus 
for 
a 
single-rooted 
P2 
in 
Machairodus 
and 
characterized 
the 
genus 
(p. 
271) 
as 
having 
P2 
“vestigial 
or 
absent.” 
None 
of 
the 
maxillaries 
on 
hand 
have 
an 
alveolus 
for 
P2 
and 
all 
are 
well 
preserved 
in 
the 
area 
such 
a 
tooth 
would 
occupy. 
The 
presence 
of 
the 
upper 
P2 
must 
have 
been 
a 
rare 
or 
abnormal 
occurence 
in 
Machairodus. 


P3 
is 
a 
much 
larger 
tooth 
than 
it 
is 
in 
either 
Homotherium 
or 
Smilodon. 
There 
are 
three 
major 
cusps 
and 
a 
well 
developed 
posterior 
basal 
cusp,
plus 
a 
strong 
anterior 
cingulum 
with 
several 
small 
cusps 
along 
its 
anterior 
edge. 


Fig. 
8. 
Right 
upper 
(TMM 
41261-10) 
and 
left 
lower 
(TMM 
41261-9) 
jaws 
of 
young 
adult 
Machairodus 
cotocopis, 
in 
lateral 
view. 
X 
.50 
natural 
size. 



P
P
4 
is 
very 
similar 
to 
the 
same 
tooth 
in 
Smilodon. 
The 
paracone 
averages 
shorter 
and 
the 
metacone 
longer, 
but 
both 
of 
these 
characters 
are 
variable 
in 
Smilodon. 
The 
P4 
appears 
to 
be 
larger 
in 
Machairodus 
than 
in 
Homotherium. 


Burt 
(1931:264) 
has 
termed 
the 
uper 
P 
4 
of 
Machairodus 
“relatively 
longer, 
narrower 
and 
more 
brachydont 
than 
the 
corresponding 
tooth 
in 
Smilodon.'' 
Measurements 
indicate 
that 
the 
carnassial 
of 
Machairodus 
falls 
into 
the 
upper 
range 
of 
measurements 
of 
Smilodon 
in 
length, 
and 
near 
the 
mean 
in 
width. 
The 
tooth 
does 
not 
seem 
to 
me 
to 
be 
more 
brachydont 
thanthatof 
Smilodon. 


The 
internal 
(protocone) 
root 
of 
P 
4 
is 
said 
byBurt 
to 
be 
better 
developed 
than 
it 
is 
in 
Smilodon. 
This 
is 
not 
always 
true, 
and 
in 
one 
tooth 
the 
internal 
root 
is 
firmly 
fused 
to 
the 
anterior 
root 
for 
the 
entire 
length. 
In 
Homotherium 
the 
upper 
P 
4 
lacks 
the 
internal 
root. 


The 
upper 
cheek 
teeth 
of 
Machairodus 
are 
primitive 
in 
having 
a 
larger, 
more 
complicated 
P3 
than 
either 
Smilodon 
or 
Homotherium, 
having 
the 
internal 
root 
present 
on 
P 
4 
and 
in 
being 
more 
up


, 


right, 
less 
backward-inclined. 
The 
upper 
dentition 
is 
not 
so 
primitive, 
however, 
as 
it 
appeared 
from 
the 
few 
specimensavailable 
toBurt. 


Mandible.—The 
mandible 
(Fig. 
8) 
is 
long 
and 
straight, 
as 
long 
as 
the 
longest 
mandibles 
of 
Smilodon 
and 
Homotherium. 
It 
is 
relatively 
deep, 
but 
thinner 
than 
in 
Smilodon. 
Mawby 
(1965:585) 
has 
characterized 
the 
mandible 
of 
Machairodus 
as 
relatively 
slender, 
but 
the 
specimens 
show 
it 
to 
have 
about 
the 
pz'oportions 
of 
Smilodon 
and 
Homotherium. 
The 
average 
length 
is 
about 
the 
same 
as 
in 
those 
genera, 
while 
the 
depth 
of 
the 
jaw 
averages, 
at 
most 
places, 
in 
the 
upper 
range 
of 
Homotherium 
measurements 
and 
in 
the 
upper 
range 
or 
greaterthanthe 
same 
measurementsofSmilodon. 


The 
lateral 
flanges 
extend 
below 
the 
symphysis, 
as 
in 
both 
Smilodon 
and 
Homotherium, 
but 
they 
are 
not 
well 
developed 
anteriorly. 
They 
form 
rounded 
ridges 
on 
the 
anterior 
edges 
of 
the 
jaw,
rather 
than 
keels 
of 
bone. 
The 
articular 
condyle 
is 
low, 
as 
in 
Homotherium, 
but 
not 
so 
low 
as 
in 
Smilodon. 
The 
angular 
process 
of 
M. 
catocopis 
is 
less 
inflected 
than 
that 
of 
Smilodon. 
The 
most 
strikingdifference 
separating 
the 
mandible 
of 
M. 
catocopisfrom 
that 
of 
the 
tw’o 
Pleistocene 
cats 
is 
the 
relativelyhigh, 
sloping, 
triangular, 
coronoid 
process. 
The 
coronoid 
process 
of 
Smilodon 
is 
much 
lower, 
and 
that 
ofHomotherium 
lower 
and 
flatteryet. 


There 
are 
two 
mental 
foramina 
in 
M. 
catocopis 
: 


P
P
a 
large 
anterior 
foramen 
located 
beneath 
or 
slightlyanterior 
to 
the 
center 
of 
the 
diastema, 
and 
directed 
internoposterioly, 
and 
a 
posterior 
foramen, 
oval 
in 
shape 
and 
located 
just 
beneath 
the 
anterior 
root 
of 
3. 
The 
foramina 
vary 
in 
size 
from 
specimen 
to 
specimen 
but 
are 
uniform 
in 
their 
positions. 
Both 
are 
near 
the 
lower 
margin 
of 
the 
jaw 
with 
the 
posterior 
foramen 
usually, 
perhaps 
always, 
lowest. 
Similar 
foramina 
are 
seen 
in 
Meade's 
figure 
of 
Homotherium 
(1961: 
Plate 
1), 
and 
are 
described 
in 
Ischyrosmilus 
by 
Mawby 
(1965). 
Smilodon,
however, 
has 
a 
single 
large 
mental 
foramen 
located 
near 
the 
center 
of 
the 
diastema, 
and 
sometimes 
an 
additional 
small 
foramen 
located 
farther 
forward. 


Inferior 
dentition.-—The 
lower 
incisors 
of 
Machairodus 
catocopis 
are 
not 
present 
in 
any 
of 
my 
specimens, 
nor 
in 
any 
of 
Burt’s 
specimens. 
The 
lower 
canines 
are 
present, 
though 
worn, 
in 
two 
rami. 
In 
both 
the 
canines 
are 
round 
in 
cross-section, 
both 
through 
the 
root 
and 
across 
the 
enamel 
of 
the 
crown. 
The 
alveolus 
is 
also 
round. 
These 
lower 
canines 
are 
not 
at 
all 
as 
described 
by 
Burt. 
The 
supposed 
lower 
canine 
figured 
by 
Burt 
(1931: 
Plate 
45) 
is 
flat 
and 
triangular, 
with 
a 
root 
longer 
than 
the 
depth 
of 
the 
lower 
jaw 
of 
Machairodus 
at 
the 
alveolus 
of 
the 
canine. 
The 
elongated 
alveoli 
shown 
in 
Burt’s 
Plate 
46 
may 
result 
from 
breakage. 
The 
canine 
of 
Machairodus 
is 
essentially 
as 
in 
Smilodon,
but 
the 
teeth 
are 
too 
worn 
to 
disclose 
details 
of 
basal 
cusps.

The 
cheek 
teeth 
of 
Machairodus 
catocopis 
are 
set 
nearly 
vertically 
in 
the 
jaw, 
thus 
differing 
markedlyfrom 
both 
Smilodon 
and 
Homotherium, 
in 
which 
the 
teeth 
are 
strongly 
inclined 
posteriorly. 
The 
teeth 
are 
thinner, 
less 
curved 
and 
folded 
on 
their 
lateral 
surfaces, 
than 
the 
teeth 
of 
Smilodon 
; 
they 
are 
not 
brachyodont 
as 
stated 
by 
Burt, 
but 
are 
relativelyhigher 
than 
the 
teeth 
of 
Smilodon, 
and 
perhaps 
of 


Homotheriumas 
well. 


P3 
is 
always 
present, 
larger 
than 
in 
Homotherium, 
and 
much 
larger 
than 
in 
the 
rare 
specimens 
of 
Smilodon 
that 
have 
this 
tooth. 
There 
arc 
three 
main 
cusps 
and 
a 
strong 
posterior 
cingulum. 
The 
tooth 
is 
placed 
on 
the 
outer 
edge 
of 
the 
jaw 
in 
Machairodus,

labial 
to 
the 
longitudinal 
axis 
of 
P., 
and 
Mt. 
It 
is 
separated 
from 
P, 
by 
a 
short 
distema. 
A 
similar 
position 
is 
figured 
for 
the 
jaw 
of 
Homotherium 
byMeade 
(1961: 
Plate 
1), 
but 
in 
Homotherium 
the 
P3 
is 
smaller 
and 
tipped 
backward 
rather 
than 
almost 
upright 
in 
the 
jaw.

P., 
is 
large, 
about 
as 
in 
Smilodon, 
but 
is 
simpler,
with 
three 
major 
cusps, 
a 
strong 
posterior 
cingulum, 



and 
small 
posterior 
cingular 
cusp. 
The 
P 
t 
of 
Machairodus 
is 
similar 
to 
that 
of 
Homotherium 
but 
not 
so 
strongly 
tipped 
backward 
in 
the 
jaw.

M, 
resembles 
the 
same 
tooth 
in 
Smilodon 
but 
is 
relatively 
longer, 
more 
slender, 
and 
less 
inclined 
posteriorly. 
The 
protoconid 
blade 
is 
longer 
than 
the 
paraconid 
blade, 
as 
in 
Smilodon. 
There 
is 
no 
metaconid 
and, 
as 
Burt 
concluded, 
M. 
catocopis 
probably 
does 
not 
have 
a 
metaconid 
on 
the 
M 
t. 


The 
lower 
dentition 
of 
Machairodus 
catocopis 
is 
of 
a 
simple 
and 
generalized 
marchairodont 
type.
There 
are 
no 
aberrant 
features. 
There 
is 
a 
greaterresemblance 
to 
teeth 
of 
Homotherium 
than 
to 
Smilodon. 


Vertebrae.—T 
he 
vertebralformula 
ol 
Machairo


dus 
catocopis 
appears 
to 
have 
been; 
cervicals, 
7; 
thoracics, 
13; 
lumbars, 
7; 
sacrals, 
3; 
caudals, 
2 
plus 
?. 


Burt 
(1931) 
has 
figured 
the 
damaged 
axis 
and 
atlas 
of 
Machairodus 
but 
the 
other 
vertebrae 
were 
hitherto 
unknown. 
The 
skeleton 
(TMM41261-8)
lacks 
only 
the 
fourth, 
ninth 
and 
tenth 
thoracic 
vertebrae 
and 
all 
caudals 
posterior 
to 
C2. 
Most 
of 
the 
vertebrae, 
especially 
the 
thoracics, 
have 
suffered 
some 
damage 
to 
projecting 
parts, 
such 
as 
neural 
spines 
and 
transverse 
processes. 
The 
damage 
is 
often 
confined 
to 
the 
side 
of 
the 
bone 
that 
lay 
upward 
in 
the 
matrix. 


The 
cervical 
vertebrae 
are 
approximately 
the 


size 
of 
those 
of 
Homotherium, 
but 
are 
much 
smaller 


than 
the 
cervicals 
of 
Smilodon. 
The 
latter 
are 
pow


erful, 
with 
high 
neural 
spines 
and 
broad, 
wide 


transverse 
processes. 
The 
vertebrae 
of 
Homother


iumaremorelike 
thoseofMachairodusbut,though 


no 
larger, 
appear 
more 
robust. 
'The 
sixth 
to 
seventh 


cervicals 
of 
Homotherium 
are 
unknown, 
but 
the 


thirdtofifth 
havebroader, 
stronger 
transverse 
proc


esses 
and 
apparently 
had 
longer, 
stronger 
neural 


spines. 
The 
neck 
and 
neck 
musculature 
of 
Mach


airodus 
was 
probably 
not 
as 
powerful 
as 
in 
Homo


theriurn 
andSmilodon. 


The 
atlas 
(Figs. 
9, 
10) 
of 
Machairodus 
differs 


markedly 
from 
that 
of 
both 
Smilodon 
and 
Homo


therium 
in 
having 
a 
much 
broader, 
more 
catlike, 


transverse 
process. 
As 
in 
the 
true 
cats, 
the 
anterior 


margin 
of 
the 
process 
extends 
outward 
and 
then 


sweeps 
backward 
in 
a 
smooth 
curve. 
In 
the 
Pleisto


cene 
saber-tooths 
the 
anterior 
margin 
of 
the 
trans


verse 
process 
is 
inclined 
steeply 
backward, 
forming 


a 


narrow 
blade. 
The 
process 
of 
Machairodus 
extends 
posteriorly 
to 
about 
the 
level 
of 
the 
middle 
of 
the 
axis, 
as 
it 
does 
in 
other 
saber-tooths. 
The 
dorsal 


surface 
of 
the 
neural 
arch 
is 
as 
in 
Smilodon, 
with 
a 
strong 
notch 
anteriorly. 
The 
ventral 
surface 
is 
narrow 
but 
has 
a 
strong 
median 
spine 
posteriorly. 
Ihe 
neural 
canal 
is 
marked 
internally 
by 
large 
shell-like 
spurs 
of 
bone 
at 
the 
upper 
level 
of 
the 
odontoid 
process. 
The 
spurs 
are 
relatively 
much 
more 
prominent 
than 
in 
the 
puma, 
and 
more 
prominent 
than 


figured 
in 
Smilodon. 
They 
have 
not 
been 
described 
in 
Homotherium. 


The 
axis 
(Figs. 
9, 
10) 
is 
well 
preserved. 
In 
lateral 
view 
the 
top 
of 
the 
neural 
spine 
is 
flatter 
and 
straighter 
than 
in 
Smilodon, 
and 
the 
posterior 
end 
is 
longer 
and 
more 
slender. 
The 
odontoid 
process 
is 
relatively 
larger 
and 
the 
transverse 
processes 
turn 
more 
downward, 
less 
backward, 
than 
in 
Smilodon. 
In 
posterior 
view 
the 
neural 
spine 
is 
slender, 
not 
broad 
as 
in 
Smilodon. 
It 
comes 
almost 
to 
a 
sharpedge 
dorsally, 
rather 
than 
being 
broad 
and 
flat 


above. 
The 
neural 
spine 
of 
Machairodus 
is 
catlike, 
and 
resembles 
that 
of 
the 
puma 
more 
than 
it 
does 
Smilodon. 
In 
Machairodus, 
however, 
the 
posterior 
part 
of 
the 
spine 
is 
deeply 
excavated 
ventrally,
above 
the 
posterior 
zygapophyses. 
There 
are 
no 
such 
depressions 
in 
the 
spine 
of 
the 
puma. 
The 
neural 
spine 
of 
the 
axis 
of 
Homotherium 
is 
not 
known. 


The 
third 
cervical 
vertebra 
(Fig. 
10) 
seems 
to 
resemble 
closely 
that 
of 
Homotherium 
except 
that 
the 
transverse 
process 
is 
narrower, 
and 
as 
seen 
from 
abovedoesnot 
appeartobebifurcateddistally.The 
transverse 
process 
is 
relatively 
longer 
and 
broader, 
more 
laterally 
inclined, 
than 
in 
Smilodon. 
"The 
presence 
of 
hyperapophyses 
cannot 
be 
determined 
for 
the 
part 
of 
the 
arch 
that 
would 
support 
them 
is 
missing. 
The 
centrum 
is 
keeled 
ventrally.

On 
the 
fourth 
cervical, 
there 
is 
a 
median 
longitudinal 
keel 
of 
bone 
on 
the 
dorsal 
surface 
of 
the 
neural 
arch, 
and 
a 
slightly 
thickened, 
raised 
area 
near 
its 
center 
represents 
the 
vestige 
of 
the 
neural 
spine. 
There 
are 
distinct 
neural 
spines 
on 
this 
vertebra 
in 
Smilodon 
and 
Homotherium. 
'The 
transverse 
process 
has 
an 
anterior 
extension 
that 
reaches 
pastthe 
anterior 
epiphysis 
of 
the 
centrum. 
There 
is 
no 
such 
extension 
in 
Smilodon. 
The 
lateral 
wall 
of 
the 
neural 
arch 
has 
a 
deep 
depression, 
such 
as 
is 
present 
in 
Panthera 
atrox, 
but 
not 
Smilodon. 
Such 
depressions 
are 
present 
on 
the 
third 
through 
sixth 
cervical 
vertebrae. 
The 
centrum 
is 
keeled 
ventrally.

The 
neural 
spine 
of 
the 
fifth 
cervical 
is 
broken 


off, 
but 
the 
remaining 
portion 
indicates 
that 
it 
re


sembled 
that 
of 
Smilodon 
but 
was 
smaller. 
The 
cen


trum 
is 
relatively 
broad 
with 
a 
strong 
median 



drawn 


is)

sequence
lumbar
Note 
size.natural
7.50
Approx.
missing.


are

10 


and 


9, 
4,

Thoracics
41216—8.


TMM


skeleton


from

catocopis


Machairodus


of

column
Vertebral


9. 
Fig. 


the

from

shown


are

vertebrae


Other

scavengers.


of

activities


the 


of

result

a 


as

probably


sideleft 
the 


on

missing


are

which


processes
transverse


show 


to

order

in 


side 
rightthe

from 


left. 



ventral 
keel. 
The 
vertebrarterial 
canal 
is 
rounded, 
not 
slitlike 
as 
in 
Smilodon. 
It 
resembles 
the 
canal 
of 
Homotherium. 
The 
inferior 
lamella 
of 
the 
transverse 
processes 
do 
not 
extend 
downward 
in 
a 
triangle, 
as 
in 
Smilodon, 
or 
a 
roundish 
plate 
as 
in 
P. 
atrox, 
or 
in 
a 
squarish 
plate 
as 
in 
Felis 
concolor. 
Instead 
the 
lamella 
forms 
a 
roundish 
oval 
plate. 
The 
anterior 
zygapophyses, 
as 
seen 
in 
lateral 
view, 
are 
rounder 
and 
more 
vertically 
placed 
than 
in 
Smilo


don. 


The 
spine 
of 
the 
sixth 
cervical, 
to 
judgefrom 
the 
part 
of 
the 
base 
remaining, 
was 
similar 
to 
that 
of 
Smilodon, 
but 
smaller. 
The 
anterior 
zygapophyses 
are 
more 
horizontal 
than 
those 
of 
Smilodon, 
appearing 
almost 
flat 
in 
front 
view. 
The 
inferior 
lamella 
is 
narrower 
than 
that 
of 
Smilodon, 
with 
the 
ventral 
surface 
sinuous 
in 
lateral 
view. 
Otherwise 
the 
vertebra 
is 
much 
like 
that 
of 
Smilodon. 
The 
sixth 
cervical 
of 
Homotherium 
is 
unknown. 
The 
seventh 
cervical 
vertebra 
is 
well 
preserved 
and 
the 
neural 
spine 
is 
present. 
It 
is 
much 
smaller 
and 
more 
slender 
than 
the 
spine 
of 
Smilodon, 
with 
a 
smaller 
distal 
swelling. 
The 
neural 
canal 
is 
rounder, 
less 
triangular, 
than 
the 
neural 
canal 
of 
Smilodon. 
The 
anterior 
zygapophyses 
are 
more 
horizontal 
and 
the 
transverse 
processes 
extend 
more 
ventrally, 
less 
laterally, 
than 
in 
Smilodon. 
The 
vertebra 
is 
more 
deeply 
indented 
between 
anterior 
and 
posteriorzygapophyses. 
The 
neural 
canal 
is 
shorter, 
so 
that 
more 
of 
the 
centrum 
is 
visible 
in 
dorsal 
view. 
Otherwise 
the 
vertebra 
is 
like 
that 
of 
Smilodon. 
The 
seventhcervicalofHomotherium 
isunknown. 


The 
first 
three 
thoracic 
vertebrae 
(Fig. 
10) 
were 
articulated 
with 
the 
last 
cervical. 
The 
fourth 
thoracic 
was 
not 
found. 
Vertebrae 
T5 
to 
T8 
were 
scattered 
but 
closely 
associated 
with 
the 
skeleton. 
T9 
and 
TlO 
were 
missing. 
Til 
had 
been 
dragged 
forward 
and 
lay 
partly 
beneath 
the 
last 
cervical. 
Tl2 
and 
Tl3 
were 
articulated 
and 
lay 
a 
few 
inches 
away 
from 
the 
articulated 
lumbar 
series. 


The 
thoracic 
series 
of 
Machairodus 
catocopis 
differs 
from 
the 
thoracic 
vertebrae 
of 
Smilodon 
and 
Homotherium 
in 
numerous 
ways. 
The 
most 
strikingdifferences 
are 
in 
the 
much 
smaller, 
weaker 
neural 
spines 
and 
more 
slender, 
weak 
transverse 
processesplaced 
lower 
on 
the 
neural 
arches. 
The 
back 
musculature 
of 
M. 
catocopis 
was 
not 
nearly 
so 
strong,
especially 
in 
the 
shoulder 
region, 
as 
that 
of 
Smilodon 
or 
Homotherium. 


The 
neural 
spine 
of 
T1 
is 
thin 
transversely 
but 
broad 
anteroposteriorly 
at 
the 
base. 
It 
narrows 
swiftly 
above 
to 
arch 
back 
in 
saberlike 
form 
to 
a 


narrow, 
scarcely 
swollen 
tip. 
In 
Smilodon 
and 
Homotherium 
the 
neural 
spine 
is 
thick 
and 
deep,
uniformly 
deep 
through 
its 
length, 
and 
broadly 
swollen 
at 
the 
tip. 
The 
centrum, 
in 
M. 
catocopis, 
is 
shallower 
than 
in 
Smilodon. 
'The 
anterior 
zygapophyses 
arc 
more 
vertical 
and 
the 
transverse 
processes 
extend 
almost 
horizontal 
rather 
than 
down


ward 
as 
in 
Smilodon. 
There 
is 
a 
shallow 
keel 
on 
the 
under 
side 
of 
the 
centrum, 
and 
a 
depression 
on 
each 
side 
of 
the 
keel, 
as 
in 
Homotherium. 
The 
ends 
of 
the 
transverse 
processes 
extend 
far 
forward, 
so 
that 
a 
line 
connecting 
their 
anterior 
tips 
lies 
10 
mm. 
in 
front 
of 
the 
centrum. 


T2 
bears 
the 
largest, 
heaviest 
neural 
spine 
in 
the 
series. 
The 
spine 
is 
still 
narrow 
and 
slender 
as 
compared 
with 
the 
neural 
spine 
of 
T2 
of 
Smilodon 
and 
Homotherium. 
There 
is 
a 
slight 
keel 
on 
the 
ventral 
side 
of 
the 
centrum, 
missing 
in 
Homotherium 
and 
Smilodon. 
A 
line 
connecting 
the 
anterior 
tips 
of 
the 
transverse 
processes 
lies 
2 
mm. 
ahead 
of 
the 
centrum, 
but 
in 
Smilodon 
and 
seemingly 
also 
in 
Homotherium, 
such 
a 
line 
would 
fall 
far 
behind 
the 
anterior 
epiphysis 
of 
the 
centrum. 


The 
neural 
spine 
of 
T3 
is 
slender, 
narrowingsmoothly 
and 
uniformly 
above 
to 
form 
a 
long,
slender 
triangle 
with 
a 
slight 
terminal 
swelling. 
It 
is 
smaller, 
thinner, 
more 
tapering, 
and 
has 
a 
smaller 
terminal 
button, 
than 
the 
corresponding 
spine 
of 
Smilodon, 
and 
is 
also 
more 
upright, 
with 
less 
backward 
inclination. 
There 
is 
a 
slight 
ventral 
keel 
on 
the 
centrum. 
The 
T3 
of 
Homotherium 
is 
unknown. 


The 
neural 
spine 
of 
T5 
is 
thicker 
at 
the 
base 
than 
in 
the 
more 
anterior 
vertebrae. 
It 
tapers 
uniformlyupward 
to 
form 
a 
long, 
slender 
triangle 
with 
scarcely 
any 
terminal 
button. 
The 
spine 
is 
much 
smaller 
and 
narrower, 
anteroposteriorly, 
than 
in 
Smilodon 
or 
Homotherium. 
The 
lateral 
depression 
on 
the 
spine 
of 
Homotherium, 
described 
by 
Meade, 
is 
lacking 
in 
M. 
catocopis. 
The 
centrum 
bears 
a 
scarcely 
perceptible 
keel. 


In 
T1 
to 
T5 
the 
neural 
spines 
are 
generally 
upright. 
From 
T6 
posteriorly 
they 
are 
strongly 
inclined 
backward. 
In 
Smilodon 
and 
Homotherium 
all 
of 
the 
thoracic 
neural 
spines 
are 
strongly 
inclined 
backward. 
The 
erect 
anterior 
spines 
are 
typical 
of 
the 
felines, 
and 
the 
thoracic 
vertebrae 
of 
M. 
catocopis 
resemble 
those 
of 
Felis 
concolor, 
the 
puma, 
in 
this 
respect.

The 
neural 
spine 
of 
T6 
is 
thicker 
than 
in 
the 
more 
anterior 
vertebrae, 
and 
rounder 
in 
cross 
section. 
It 
tapers 
rapidly, 
but 
not 
so 
rapidly 
as 
the 
spine 
of 
T5. 
It 
is 
smaller 
and 
lighter 
than 
the 
spines 
of 
T6 



of 
iSmilodon 
and 
Homotherium. 
The 
centrum 
is 
not 
keeled 
ventrally.

The 
part 
of 
the 
spine 
of 
T7 
remaining 
suggeststhat 
it 
was 
larger 
and 
more 
posteriorly 
inclined 
than 
the 
neural 
spine 
of 
T6. 
It 
was 
probably 
smaller 
than 


in 
Smilodon 
and 
Homotherium. 
The 
transverse 
processes 
are 
relatively 
longer 
and 
lighter 
than 
those 
of 
Smilodon. 
The 
centrum 
is 
not 
keeled, 
nor 
are 
anyofthecentraposteriortoT5 
keeled. 


The 
broken 
base 
of 
the 
spine 
of 
4’B 
indicates 
that 
it 
was 
strongly 
inclined 
backward. 
The 
vertebra 
throughout 
appears 
smaller 
and 
lighter 
in 
build 
than 
that 
of 
Smilodon. 
4’he 
posterior 
zygapophyses 
are 
almost 
horizontal. 


Tll 
has 
no 
neural 
spine 
but 
there 
is 
a 
low, 
sharp-
edged 
ridge 
or 
keel 
on 
the 
dorsal 
side 
of 
the 
neural 
arch. 
The 
posterior 
zygapophyses 
are 
elongated,
and 
almost 
vertical. 
In 
the 
known 
Homotherium 
and 
typical 
SmilodonTil 
bears 
a 
large, 
strong,

, 


posteriorly 
inclined 
neural 
spine. 
However, 
Merriam 
and 
Stock 
figure 
(1932: 
86) 
a 
“spinelesstype” 
of 
1 
1 
th 
thoracic 
vertebra 
that 
is 
a 
rare 
form 
in 
Smilodon. 
This 
closely 
resembles 
the 
presentspecimenofM.catocopis.ThecentrumofM. 
catocopis 
is 
relatively 
deeper 
than 
that 
of 
Smilodon, 
but 
the 
dorsal 
side 
(the 
ventral 
surface 
of 
the 
neural 
canal) 
is 
channeled 
and 
not 
flat 
or 
nearly 
flat 
as 
in 
Smilodon. 
As 
a 
result 
of 
the 
channeling, 
the 
centrum 


is 
heart-shaped 
in 
posterior 
view, 
and 
only 
slightlyless 
so 
in 
front 
view. 


The 
neural 
spine 
is 
undeveloped 
in 
Tl2, 
represented 
only 
by 
a 
low, 
knife-edged 
ridge 
22 
mm. 
longand 
4 
mm. 
high. 
There 
is 
a 
strong 
spine 
in 
4'12 
of 
Homotherium 
and 
Smilodon. 
The 
anapophyses 
are 
well-developed 
but 
there 
arc 
no 
metapophyses 
remaining 
on 
the 
damaged 
anterior 
edge 
of 
the 
anterior 
zygapophyses.

The 
neural 
spine 
of 
413 
is 
short, 
broad, 
and 
erect. 
Small 
metapophyses 
are 
present. 
The 
anapophyses 
are 
as 
in 
Smilodon. 
The 
centrum 
is 
channeled 
dorsally,
but 
istoobroadtoappearheart-shaped.There 
is 
no 
ventral 
ridge 
on 
the 
centrum, 
as 
in 
Homotherium. 


Relative 
to 
the 
thoracic 
vertebrae, 
the 
lumbars 
are 
stout 
and 
strong 
(Figs. 
9, 
10). 
They 
are 
smaller 
than 
the 
lumbars 
of 
Smilodon 
but 
comparable 
in 
strength, 
and 
were 
even 
more 
firmly 
united 
in 
articulation.
Thezygapophysesarealmost 
verticaland 
parallel, 
with 
the 
lower 
edges 
of 
the 
anterior 
zygapophyses 
curved 
inward, 
thus 
permitting 
almost 
no 
side-to-side 
movement. 
44ie 
neural 
arch 
is 
notched 
posteriorly, 
and 
the 
posterior 
face 
of 
the 
neural 
spine 
bears 
a 
deep 
groove, 
into 
which 
the 
anterior 
edge 
of 
the 
succeeding 
spine 
fits 
when 
the 
column 
is 
flexed. 
There 
was 
thus 
considerable 
vertical 
freedom 
of 
movement 
of 
the 
hip 
region. 
The 


Fig 
10. 
Vertebrae 
of 
Machairodus 
catocopis 
skeleton 
(TMM 
41261—8), 
viewed 
from 
above. 
A, 
atlas 
and 
axis; 
B, 
third 
lumbar; 
C, 
sacrum. 
Approx. 
X 
.50 
natural 
size. 
Note 
the 
anterior 
position 
of 
the 
neural 
spine 
and 
the 
deep 
groove 
in 
the 
posterior 
face 
of 
the 
spine 
for 
the 
reception 
of 
the 
neural 
spine 
of 
lumbar 
4 
when 
the 
vertebral 
column 
is 
deeply 
flexed. 



functional 
implications 
ol 
these 
conditions 
are 
discussed 
below. 
Anapophyses 
and 
metapophyses 
appear 
to 
have 
been 
less 
developed 
than 
in 
Smilodon. 
The 
most 
striking 
difference 
between 
the 
lumbar 
vertebrae 
of 
M. 
catocopis 
and 
those 
of 
Smilodon 
is 
the 
forward 
position 
of 
the 
neural 
spine 
in 
Ma


chairodus. 
There 
the 
neural 
spines 
slope 
anteriorly,
with 
the 
anterior 
edge 
upright 
and 
the 
posterioredge 
angled 
forward. 
In 
Smilodon 
the 
reverse 
is 
true, 
and 
the 
neural 
spines 
slope 
backward. 


Inlateralview,theneuralspineofLI 
isstrongly 
inclined 
forward, 
and 
the 
metapophysis 
is 
less 
prominent 
than 
in 
Smilodon. 
The 
anterior 
zygapophyses 
arc 
almost 
vertical. 
The 
transverse 
processes 
are 
strongly 
inclined 
downward; 
they 
are 
almost 
horizontal 
in 
Smilodon. 
The 
posterior 
zygapophyses 
are 
narrower 
and 
closer 
together 
than 
in 
Smilodon. 
The 
anapophyses, 
as 
a 
result, 
are 
closer 
together 
and 
almost 
parallel. 
Seen 
from 
above, 
the 
neural 
spine 
lies 
over 
the 
anterior 
edge 
of 
the 
vertebra, 
with 
the 
button 
on 
the 
top 
of 
the 
spine 
extending 
forward 
pastthe 
edge 
of 
the 
interzygapophyseal 
notch. 
There 
is 
a 
deep 
notch 
between 
the 
posterior 
zygapophyses, 
and 
the 
neural 
spine 
is 
deeply 
grooved 
for 
approximately 
half 
of 
the 
length 
of 
its 
posterior 
side, 
for 
the 
reception 
of 
the 
neural 
spine 
of 
L 
2 
when 
the 
vertebralcolumn 
isflexed. 


In 
L2, 
the 
neural 
spine 
is 
notched 
and 
grooved

posteriorly,for 
thereceptionof 
thespine 
ofL3,but 


the 
groove 
is 
not 
so 
deep 
as 
in 
the 
spine 
of 
LI. 
The 


anterior 
zygapophyses 
are 
almost 
vertical. 
The 
pos


terior 
zygapophyses 
are 
more 
widely 
separated 
than 


those 
of 
Smilodon. 
There 
is 
no 
ridge 
on 
the 
ventral 


surface 
of 
LI 
or 
on 
the 
anterior 
half 
of 
the 
centrum 


of 
L2, 
but 
a 
ridge 
begins 
on 
the 
posterior 
half 
of 


L2. 
In 
Smilodon 
and 
Homotherium 
the 
anterior 


lumbars 
are 
ridged 
ventrally.

In 
L 
3 
the 
neural 
spine 
is 
strong 
but 
so 
situated 


anteriorly 
that, 
when 
seen 
from 
directly 
above, 
more 


than 
half 
of 
the 
interzygapophyseal 
notch 
is 
ob


scured 
by 
the 
terminal 
button 
of 
the 
spine. 
The 


posterior 
zygapophyses 
are 
farther 
apart 
than 
those 


of 
L2, 
and 
much 
more 
so 
than 
those 
of 
LI. 
The 


notch 
and 
groove 
in 
the 
posterior 
face 
of 
the 
neural 


arch 
and 
spine 
are 
like 
those 
of 
L2. 
The 
centrum 
is 


heavily 
ridged 
ventrally.

The 
anterior 
zygapophyses 
of 
L 
4 
are 
not 
so 
erect 


as 
those 
of 
LI-3, 
and 
are 
like 
those 
of 
Smilodon. 


The 
notch 
and 
groove 
in 
the 
posterior 
face 
of 
the 


neural 
arch 
and 
spine 
are 
broader 
and 
shallower 


than 
in 
the 
more 
anterior 
vertebrae. 
The 
posterior

zygapophyses 
are 
more 
widely 
separated 
than 
those 


of 
Smilodon. 
The 
transverse 
process 
is 
like 
that 
ol 
Smilodon. 
The 
neural 
spine 
of 
L 
5 
is 
like 
that 
of 
L3, 
but 
the 
terminalbutton 
is 
smaller. 
The 
anterior 
zygapophyses 
are 
as 
in 
Smilodon. 
The 
posterior 
zygapophyses 


are 
similar 
to 
those 
of 
Smilodon, 
but 
the 
notch 
between(
hemisdeeperwhenseenindorsalview. 
1he 
grooveforthefrontoftheneuralspineofT6 
isonly 
aboutone-third 
of 
the 
lengthof 
thespineof 
vertebra 


L5.Thecentrum 
isverystronglykeeled. 


L 
6 
is 
much 
like 
that 
of 
Smilodon 
but 
the 
transverse 
process 
is 
longer, 
wider, 
and 
more 
downward-
turned. 
The 
anterior 
zygapophyses 
are 
cupped, 
like 
those 
of 
Smilodon, 
but 
the 
walls 
external 
to 
the 
zygapophyses 
are 
much 
thinner 
than 
in 
Smilodon. 
The 
posterior 
zygapophyses 
are 
smaller 
than 
those 
of 
Smilodon 
but 
have 
a 
similar 
notch 
between 
them. 
The 
neural 
spine 
bears 
a 
groove 
posteriorly 
for 
the 
spine 
of 
L7, 
as 
it 
does 
In 
Smilodon. 
Only 
the 
anterior 
half 
of 
the 
ventral 
surface 
of 
the 
centrum 
bears 
a 
ridge.

The 
metapophyses 
of 
L 
7 
do 
not 
extend 
so 
far 
forward 
as 
in 
Smilodon, 
and 
the 
centrum 
extends 
forward 
beyond 
them, 
as 
seen 
in 
dorsal 
view. 
In 
Smilodon 
the 
anterior 
margins 
of 
the 
metapophysesextend 
forward 
past 
the 
end 
ol 
the 
centrum, 
and 
there 
are 
excavations 
in 
the 
wall 
of 
the 
neural 
arch 
between 
the 
bases 
of 
the 
transverse 
processes 
and 
the 
metapophyses, 
lateral 
to 
the 
centrum. 
There 
are 
no 
such 
excavations 
in 
Machairodus. 
The 
postei'ior 
zygapophyses 
are 
more 
widely 
separated 
than 
those 
ol 
Smilodon. 
There 
is 
no 
ridge 
on 
the 
ventral 
surface 
of 
the 
centrum. 


The 
sacrum, 
in 
dorsal 
view 
(Fig. 
10), 
is 
less 


squarish, 
more 
tapering 
posteriorly, 
than 
the 
sacrum 


of 
Smilodon. 
It 
seems 
also 
to 
taper 
more 
than 
the 


sacrumofHomotherium. 
Thedorsalopeningsofthe 


sacral 
foramina 
are 
round-oval 
in 
shape, 
and 
are 


relatively 
larger 
and 
more 
prominent 
than 
in 
Smi


lodon. 
The 
transverse 
process 
of 
S3 
is 
longer, 
nar


rower, 
than 
in 
Smilodon. 
In 
anterior 
view 
the 
an


terior 
zygapophyses 
are 
flatter 
and 
more 
horizontal 


than 
those 
of 
Smilodon. 
The 
neural 
canal 
is 
shal


lower. 
The 
lateral 
rugosities, 
where 
the 
ilium 
was 


attached, 
arelargeandshowthattherewasapower


ful 
attachment 
between 
pelvis 
and 
sacrum. 


The 
sacrum 
of 
M. 
catocopis 
seems 
to 
resemble 


that 
of 
Homotherium 
more 
than 
Smilodon. 
How


ever, 
structural 
differences 
between 
the 
three 
genera 


are 
slight, 
and 
possibly 
within 
the 
range 
of 
indi


vidualvariationinany 
oneofthem. 


Only 
the 
proximal 
two 
caudal 
vertebrae 
were 



recovered, 
in 
articulation 
with 
the 
sacrum. 
The 
first 
caudal 
has 
a 
slender 
transverse 
process 
and 
a 
small 
but 
distinct 
neural 
spine. 
In 
Smilodon 
the 
transverse 
process 
is 
short 
and 
wide, 
and 
there 
is 
no 
neural 
spine. 
In 
C 
2 
there 
is 
no 
neural 
spine 
but 
the 
neural 
arch 
is 
complete.The 
transverse 
process 
is 
shorter 
and 
wider 
than 
in 
Cl. 
In 
Sinilodon 
and 


Homotherium 
the 
neural 
arch 
is 
incomplete 
in 
C2, 
and 
may 
be 
incomplete 
in 
the 
Cl 
of 
Homotherium. 
Both 
Pleistocene 
genera 
are 
known 
to 
have 
had 
short 
tails. 
The 
complete 
neural 
arch 
of 
the 
second 
caudal 
vertebra 
of 
Machairodus 
suggests 
a 
longerand 
more 
catlike 
tail. 


Ribs.—A 
few 
ribs 
of 
skeleton 
TMM41261-8 
were 
complete 
but 
most 
were 
damaged 
and 
many 
were 
missing. 
The 
ribs 
are 
generally 
catlike. 
They 
are 
relatively 
small 
and 
thin 
as 
compared 
with 
those 
of 


•Smilodon. 
Pectoral 
girdle 
and 
fore 
limb.—Both 
scapulae 
were 
present 
but 
most 
of 
the 
blades 
were 
crushed 
and 
shattered 
and 
could 
not 
be 
saved. 
The 
ventral 
parts 
were 
complete, 
along 
with 
parts 
of 
the 
scapularspines 
and 
some 
of 
the 
anterior 
and 
posterior 
margins 
ofboth 
specimens.

The 
suprascapular 
notch 
seems 
to 
be 
unusuallydeep, 
relatively 
deeper 
than 
in 
Homotherium,
Smilodon, 
or 
Felis 
concolor. 
The 
neck 
of 
the 
scapula, 
in 
consequence, 
is 
much 
constricted. 
Beyondthe 
scapular 
notch, 
the 
anterior 
border 
of 
the 
blade 
curves 
abruptly 
outward, 
as 
though 
to 
form 
a 
broad 
blade, 
as 
in 
Felis, 
but 
not 
enough 
remains 
to 
be 
sure 
of 
this. 
The 
posterior 
border 
is 
straight, 
as 
in 
Homotheriumand 
thepuma.

The 
left 
humerus 
is 
nearly 
complete 
but 
the 
rightlacks 
the 
proximal 
quarter. 
The 
humerus 
(Fig. 
11)
is 
vastly 
different 
from 
that 
of 
Smilodon. 
Althoughit 
is 
approximately 
the 
same 
length 
it 
is 
much 
more 
slender, 
with 
a 
mid 
shaft 
diameter 
approximatelyhalf 
that 
of 
a 
large 
Smilodon. 
At 
the 
distal 
end, 
the 
Smilodon 
humerus 
is 
about 
two 
inches 
wider. 
The 
humerus 
of 
Machairodus 
is 
also 
straighter 
in 
both 
anteroposterior 
and 
transverse 
view. 
There 
are 
numerous 
other 
differences 
but 
all 
are 
associated 
with 
the 
relative 
slenderness 
of 
the 
Machairodus 
bone. 
Someofthesedifferenceshavebeen 
notedbyBurt. 


The 
humerus 
of 
Homotherium, 
figured 
byMeade, 
is 
as 
long 
as 
thatof 
Machairodusbut 
stouter, 
and 
is 
more 
slender 
than 
that 
of 
Smilodon. 
In 
bulk 
and 
general 
appearance 
it 
is 
intermediate 
between 


M. 
catocopis 
andSmilodon. 
The 
ulna 
(Fig. 
11) 
is 
known 
from 
both 
sides 
of 
the 
skeleton 
(TMM41261-8) 
and 
an 
isolated 
speci


men. 
The 
bone 
is 
approximately 
the 
length 
of 
the 
ulna 
of 
Smilodon 
but 
much 
more 
slender. 
The 
ulna 
of 
Homotherium 
is 
slightly 
longer 
and 
slightly 
stouter 
than 
the 
ulna 
of 
Machairodus. 
In 
Smilodon 
the 
anterior 
dorsal 
surface 
of 
the 
olecranon 
bears 
a 
high, 
distinct 
crest, 
but 
in 
both 
Machairodus 
and 
Homotherium 
the 
crest 
is 
relatively 
and 
actuallymuch 
lower. 
M. 
catocopis 
differs 
from 
both 
Homotherium 
and 
Smilodon 
in 
lacking 
the 
deep, 
elon


gated 
depression 
on 
the 
shaft 
ventral 
to 
the 
facet 
for 
the 
articulation 
of 
the 
proximal 
end 
of 
the 
radius. 


The 
radius 
(Fig. 
11) 
is 
known 
from 
three 
specimens, 
two 
of 
which 
were 
isolated 
finds. 
The 
radius 
is 
very 
slender 
compared 
with 
that 
of 
Smilodon. 
It 
closely 
resembles 
the 
radius 
of 
Homotherium, 
but 
seems 
to 
be 
a 
bit 
more 
slender 
and 
curved. 
The 
neck 
of 
the 
proximal 
end 
appears 
more 
constricted, 
and 
the 
distal 
end 
of 
the 
shaft 
wider 
in 
relation 
to 
the 
rest 
of 
the 
shaft, 
than 
in 
Homotherium. 
There 
is 
considerable 
variation 
in 
all 
these 
proportions, 
however, 
as 
may 
be 
seen 
from 
the 
measurements. 


Manus.—The 
following 
elements 
are 
available: 
5 
scapholunars, 
1 
pisiform, 
1 
metacarpal 
I, 
1 
MC 
11, 
3 
MC 
Ill's, 
1 
MC 
IV, 
and 
numerous 
phalanges.

The 
scapholunars 
display 
some 
individual 
variation 
but 
are, 
in 
general, 
like 
the 
bone 
in 
Smilodon 
and 
Homotherium. 
However, 
they 
differ 
markedlyand 
uniformly 
from 
the 
scapholunar 
of 
Smilodon 
in 
that 
the 
“beak” 
is 
much 
narrower 
and 
more 
prominent.

The 
pisiform 
is 
much 
smaller 
than 
the 
pisiformof 
Smilodon, 
and 
appears 
stouter 
through 
the 
head, 
and 
less 
twisted. 
The 
cuniform 
articulation 
is 
relatively 
long, 
as 
compared 
with 
that 
of 
Smilodon, 
and 
the 
“heel” 
is 
therefore 
relatively 
longer 
but 
more 
slender. 
The 
pisiform 
resembles 
that 
of 
Felis 
co?icolor 
and 
Panther 
a 
atrox 
more 
than 
it 
does 
Smilodon. 


The 
first 
metacarpal 
is 
short 
and 
stout, 
like 
the 
MC 
I 
of 
Smilodon, 
and 
as 
in 
Smilodori, 
has 
an 
oblique 
ridge 
across 
the 
dorsal 
surface. 
This 
ridge 
is 
less 
developed 
than 
in 
Smilodoji. 
There 
are 
numerous 
other 
minor 
differences, 
but 
the 
general 
resemblance 
to 
Smilodon 
is 
close. 
On 
the 
other 
hand, 
the 
MC 
I 
of 
Panthera 
atrox, 
Felis 
concolor, 
and 
other 
true 
cats 
available, 
is 
quite 
different. 
The 
MC 
I 
of 
sabertoothsseems 
quite 
distinctive. 


Metacarpals 
11, 
111,andIV 
arelongandslender 
compared 
with 
the 
manus 
of 
Smilodon. 
They 
are 
also 
very 
straight, 
especially 
on 
the 
ventral 
surface,
compared 
with 
large 
true 
cats. 
The 
distal 
keel 
is 
not 



Fig. 
11. 
Limb 
bones 
of 
Machairodus 
calocopis, 
from 
left 
to 
right; 
humerus, 
ulna, 
radius, 
tibia, 
femur. 
The 
tibia 
(TMM 
41261—14) 
and 
femur 
(TMM 
41261 
13) 
are 
isolated 
discoveries. 
The 
other 
elements 
are 
from 
the 
skeleton 
(TMM 
41 
261—8). 
X 
.50 
natural 
size. 


“hooked"’ 
under, 
as 
in 
Smilodon. 
The 
proximal 
phalanges 
to 
their 
proper 
place. 
However, 
the 
singlearticularsurfaceofMCIIistriangularinshape,of 
ungualphalanxfoundwiththeskeletonappearsto 
MC 
111 
squarish 
with 
a 
deep, 
broad 
notch, 
and 
of 
have 
been 
from 
the 
fore 
foot. 
It 
is 
large 
and 
stronglyMG 
IV 
broad 
above 
and 
notched 
laterally. 
The 
hooded. 
It 
resembles 
the 
hooded 
terminal 
phalanxnotch 
lateral 
to 
the 
unciform 
facet 
is 
shallow, 
as 
of 
Smilodon 
in 
details 
of 
shape 
as 
well 
as 
size. 
Since 
inSmilodon,notdeepasinPantheraatrox. 
thelimbandfootbonesofM.catocopisare 
uni-
No 
effort 
has 
been 
made 
to 
assign 
the 
numerous 
formly 
smaller 
and 
more 
slender 
than 
those 
of 
Smi



lodon, 
the 
claws 
must 
have 
been 
relatively 
huge.

Pelvis 
andhindlimb.—Theinnominateresembles 
thatof 
Smilodonbut 
isthinnerandlighterinbuild. 
The 
distinct 
constriction 
anterior 
to 
the 
acetabulum 
in 
Smilodon 
is 
absent 
in 
M. 
catocopis. 
The 
ischium 
also 
appears 
to 
be 
more 
twisted. 


The 
distal 
ends 
of 
both 
femora 
of 
TMM41261-8 
were 
missing, 
as 
were 
all 
other 
bones 
of 
the 
hind 
legs, 
save 
for 
a 
few 
foot 
elements. 
The 
following 
description 
is 
based 
principally 
on 
TMM41261-13, 
a 


nearly 
complete 
femur, 
augmented 
by 
additional 
material. 


The 
femur 
of 
M. 
catocopis 
is 
long, 
straight 
and 
slender 
(Fig. 
11). 
It 
thus 
differs 
markedly 
from 
the 
femora 
of 
Smilodon, 
and 
is 
more 
like 
the 
femora 
of 
the 
true 
cats, 
such 
as 
Panthera 
and 
Felis. 
In 
Smilodon 
the 
lesser 
trochanter 
is 
more 
distinct 
from 
the 


head 
than 
it 
is 
in 
true 
cats, 
and 
in 
Homotherium 
and 
Machairodus 
also, 
where 
the 
lesser 
trochanter 
is 
placed 
closer 
to 
the 
head 
of 
the 
femur. 
The 
femur 
of 
Homotherium 
appears, 
in 
figures, 
to 
be 
stouter 
and 
more 
curved 
than 
that 
of 
Machairodus. 


The 
tibia 
of 
M. 
catocopis, 
known 
from 
two 
isolated 
specimens, 
is 
as 
long 
as 
in 
Smilodon 
and 
Homotherium, 
but 
straighter 
and 
more 
slender 
(Fig.
11). 
It 
resembles 
the 
tibia 
of 
Homotherium 
more 
than 
it 
does 
that 
of 
Srnilodon. 
The 
anterior 
notch 
in 
the 
distal 
end 
is 
much 
deeper 
than 
it 
is 
in 
either 
Smilodon 
or 
Homotherium. 
Tibiae 
tend 
to 
be 
featureless 
bones, 
but 
the 
difference 
in 
the 
anterior 
notch 
is 
striking. 
Burt 
(1931, 
Plate 
47) 
has 
figured 
a 
M. 
catocopis 
tibia 
with 
a 
similarly 
deep 
notch. 
The 
significance 
of 
this 
character 
is 
not 
clear. 


Pes.—'The 
following 
complete 
or 
nearly 
complete 
specimens 
are 
available: 
7 
astragali; 
3 
calcanea; 
1 
cuboid; 
1 
navicular; 
7 
metatarsal 
IFs; 
1 
MT 
III; 
2 
MT 
IV's; 
1 
MT 
V; 
and 
numerous 
unstudied 
phalanges.

Even 
the 
largest 
astragalus 
is 
smaller 
than 
the 
astragalus 
of 
a 
medium-sized 
Smilodon. 
The 
wall 
posterior 
to 
the 
deep 
groove 
between 
the 
calcaneal 
articulations, 
thick 
in 
Smilodon, 
is 
thin 
in 
M. 
catocopis. 
The 
bone 
appears 
more 
catlike 
than 
the 
astragalus 
of 
Smilodon, 
and 
probably 
of 
Homotherium 
also. 


The 
calcaneum, 
compared 
with 
that 
of 
Smilodon,
is 
shorter 
and 
not 
so 
broad. 
The 
distal 
end 
of 
the 
heel 
is 
more 
notched. 
The 
facet 
for 
the 
articulation 
of 
the 
navicular 
is 
smaller 
and 
less 
prominent. 
The 
major 
astragalar 
facet 
is 
smaller 
and 
more 
vertical. 
ThecalcaneumofM.catocopis 
issimilar,ingeneral, 
to 
that 
of 
Homotherium 
but 
the 
heel 
is 
distinctly 


longer 
and 
more 
notched 
distally. 
The 
heel 
is 
also 
less 
swollen 
at 
the 
distal 
end. 


The 
cuboid 
is 
generally 
similar 
to 
that 
of 
Smilodon, 
but 
there 
are 
numerous 
minor 
differences 
in 
addition 
to 
much 
smaller 
size. 
The 
facet 
for 
articulation 
with 
the 
navicular 
is 
relatively 
smaller 
and 
the 
facet 
for 
the 
ectocuniform 
is 
larger, 
with 
a 
median 
constriction 
not 
present 
in 
Smilodon. 
4'he 
facet 
for 
metatarsal 
IV 
is 
prominent, 
as 
in 
Smilodon, 
but 
the 
facet 
for 
metatarsal 
V 
is 
scarcely 
indicated. 
The 
groove 
for 
M. 
peroneus 
longus 
is 
relatively 
deeper 
and 
set 
closer 
to 
the 
distal 
border 
than 
it 
is 
in 
Smilodon. 


The 
navicular 
is 
much 
like 
that 
of 
Smilodon 
but 
smaller. 
The 
surface 
for 
the 
astragalar 
articulation 
is 
narrower 
and 
rounder 
than 
in 
Smilodon. 
The 
facet 
for 
articulation 
with 
the 
mesocuniform 
is 
low, 
not 
so 
prominent, 
as 
it 
is 
in 
Smilodon. 
The 
bone 
is 
also 
thinner 
dorsoventrally.

The 
metatarsals 
are 
markedly 
longer 
and 
more 
slender 
than 
those 
of 
Smilodon. 
They 
are 
also 
much 
straighter, 
and 
this 
is 
especially 
marked 
in 
MT 
111, 
the 
longest 
and 
straightest 
of 
the 
metatarsals. 
The 
ventral 
keel 
of 
the 
distal 
articulation 
of 
the 
metapodials 
of 
Smilodon 
is 
often 
“hooked” 
downward 
and 
backward. 
None 
of 
the 
metatarsals 
of 
M. 
catocopis 
are 
hooked. 


A 
number 
of 
minor 
differences 
are 
noted 
in 
the 
shape 
of 
the 
proximal 
articular 
surfaces 
of 
the 
metatarsals. 
Notable 
is 
the 
complete 
absence, 
on 
all 
seven 
specimens 
of 
MT 
11, 
of 
an 
articular 
surface 
for 
a 
rudimentary 
MT 
I. 
Instead 
there 
is 
a 
prominent 
crest 
on 
the 
lateral 
surface 
of 
the 
bone 
which 
would 
virtually 
preclude 
articulation 
with 
another 
metatarsal. 
There 
is 
an 
articular 
facet 
in 
Smilodon, 
and 
a 
vestigialmetatarsalI.SeeminglyM. 
catocopislacked 
MT 
I. 


REMARKS.—The 
skull, 
jaws 
and 
dentition 
of 


M. 
catocopis 
indicate 
a 
rather 
primitive, 
unspecialized 
machairodont. 
The 
skull 
is 
long 
and 
low, 
and 
cranial 
foramina 
are 
typically 
machairodont 
except 
that 
the 
alisphenoid 
canal 
is 
present. 
The 
mandible 
is 
deep, 
but 
the 
coronoid 
process 
is 
not 
as 
reduced 
as 
in 
the 
Pleistocene 
sabertooths, 
and 
the 
mandibular 
flange 
is 
not 
so 
well 
developed. 
The 
existence 
of 
the 
upper 
P2 
and 
triangular, 
flattened 
lower 
canine, 
noted 
by 
Burt 
as 
aberrant 
features, 
are 
not 
confirmed 
by 
the 
present 
material. 
The 
cheek 
teeth 
are 
high 
but 
compressed. 
The 
uppercarnassial 
still 
retains 
the 
lingual 
(protocone) 
root,
but 
reduction 
is 
evident 
and 
fusion 
of 
the 
root 
with 



the 
outer 
roots 
sometimes 
occurs. 
In 
the 
lower 
dentition, 
P;, 
is 
a 
large, 
broad 
tooth, 
whereas 
it 
is 
reduced 
or 
absent 
in 
the 
later 
sabertooths. 
The 
cheek 
teeth 
of 
both 
upper 
and 
lower 
jaws 
are 
set 
nearlyupright, 
rather 
than 
strongly 
inclined 
backward. 
Resemblance 
of 
skull, 
jaws 
and 
teeth 
of 
Machairodus 
catocopis 
to 
Homotherium 
is 
much 
closer 
than 
it 
is 
to 
Smilodon. 
The 
dentition 
of 
Machairodus 
is 
so 
generalized 
that 
the 
dentition 
of 
Homotherium 
could 
be 
derived 
from 
it. 


The 
body 
proportions 
are 
unusual. 
The 
head 
seems 
to 
have 
been 
disproportionately 
large, 
and 
the 
neck 
stout. 
The 
trunk 
was 
slender, 
and 
lacked 
the 
powerful 
muscles 
typical 
of 
the 
fore 
part 
of 
the 
body 
of 
Smilodonand 
to 
a 
more 
limited 
degree,
Homotherium. 
The, 
front 
legs 
were 
excessively 
slenderandif 
shorterthan 
thoseofHomotherium, 
were 
relatively 
long 
compared 
with 
their 
thickness. 
The 
feet 
were 
large, 
and 
the 
claws 
as 
large 
as 
those 
of 
Smilodon. 


The 
lumbar 
region 
was 
specialized 
for 
freedom 
of 
movement 
in 
the 
vertical 
plane, 
but 
almost 
rigidlaterally. 
The 
pelvis 
and 
sacrum 
are 
strongly 
built. 
The 
hind 
limbs 
are 
long, 
as 
long 
as 
in 
Homotherium, 
and 
the 
hind 
feet 
seem 
disproportionately 
long 
and 


slender. 


The 
slim 
body, 
long, 
slender 
limbs, 
and 
elongated,
slender 
feet, 
along 
with 
the 
curious 
adaptation 
of 
the 
lumbar 
region, 
may 
have 
been 
a 
specializationfor 
a 
swift, 
bounding 
gait. 
Certainly 
the 
ability 
of 
the 
hip 
region 
to 
flex 
vertically 
was 
highly 
developed. 
The 
lumbar 
vertebrae 
of 
the 
cheeta 
(Acinonyx 
jubatus) 
show 
almost 
the 
reverse 
of 
the 
kind 
of 
specialization 
noted 
in 
M. 
catocopis. 
In 
the 
cheeta 
the 
neural 
spines 
are 
small 
and 
well 
separated, 
and 
the 
zygapophyses 
are 
almost 
horizontal. 
The 
cheeta 
captures 
its 
prey 
in 
a 
swilt, 
dashing 
run. 
Agile 
twisting 
of 
the 
hip 
region 
is 
necessary 
to 
follow 
and 
inter


cept 
the 
dodging 
prey. 
The 
lumbar 
vertebrae 
of 
the 
puma 
(Felis 
concolor) 
have 
zygapophyses 
intermediate 
between 
the 
relatively 
vertical 
type 
oi 
Machairodus 
and 
the 
relatively 
horizontal 
type 
of 
Acinonyx. 
Perhaps 
the 
Pliocene 
cat 
overtook 
large

but 
clumsy 
prey 
in 
a 
series 
of 
long, 
bounding 
leaps,
fixed 
its 
huge 
claws 
in 
the 
skin 
of 
its 
victim, 
and 
drove 
the 
long 
canines 
into 
the 
flesh 
to 
Inflict 
gashesthrough 
which 
the 
prey 
quickly 
bled 
to 
death. 


Evans 
(1961: 
19) 
presents 
strong 
evidence 
that 
the 
principal 
prey 
of 
Homotherium 
at 
Friesenhahn 
Cave 
was 
the 
young 
of 
mammoths 
and 
mastodons. 
Adult 
elephants 
might 
have 
been 
killed 
as 
well, 
but 
their 
remains 
would 
have 
been 
left 
where 
killed, 
rather 
than 
dragged 
back 
into 
the 
cave. 
To 
judgefrom 
the 
few 
fossils 
recovered, 
mastodons 
were 
ex


tremely 
rare 
in 
the 
Coffee 
Ranch 
area 
when 
the 
deposits 
were 
forming. 
Rhinoceroses, 
especiallyAphelops, 
were 
common, 
and 
their 
remains 
are 
abundant. 
Fossils 
of 
young 
animals 
are 
no 
more 
common, 
relative 
to 
the 
remains 
of 
adult 
animals, 
than 
is 
usually 
the 
case 
of 
fossil 
deposits. 
Aphelops 
may 
have 
been 
the 
principal 
prey 
of 
Machairodus, 
and 
both 
young 
and 
adult 
animals 
were 
probably 
eaten. 


Some 
individual 
M. 
catocopis 
lived 
to 
a 
considerable 
age, 
and 
some 
teeth 
were 
found 
that 
had 
been 
worn 
nearly 
to 
the 
gums. 
An 
extreme 
case 
is 
represented 
by 
maxillary 
MU5634. 
This 
jaw 
has 
the 
teeth 
so 
worn 
as 
to 
be 
scarcely 
recognizable. 
Onlythe 
canine 
and 
the 
carnassial 
remain, 
and 
the 
other 
teeth 
seem 
to 
have 
been 
lost 
in 
life. 
The 
canine 
is 
worn 
until 
it 
is 
almost 
semicircular, 
nearly 
as 
wide 
as 
long. 
The 
carnassial 
is 
so 
worn 
that 
only 
a 
bit 
of 
enamel 
remains 
at 
the 
base, 
scarcely 
a 
square 
millimeter 
in 
area. 
This 
individual, 
in 
its 
old 
age, 
must 
have 
competed 
with 
the 
bone-eating 
dogs 
for 
the 
carrion 
left 
by 
more 
active 
predators. 



Speculation 
as 
to 
Faunal 
Relationships 
of 
the 
Carnivores 


Hemphill 
County 
was 
probably 
a 
grassland 
when 
the 
Coffee 
Ranch 
fauna 
lived. 
Grazing 
mammals 
are 
represented 
by 
abundant 
fossils, 
but 
browsingforms 
are 
rare. 
At 
least 
some 
bushes 
must 
have 
existed 
nearby, 
probably 
in 
canyons, 
for 
remains 
of 
mastodon, 
pigs, 
deer 
and 
ground 
sloths 
are 
occasionally 
found. 
The 
relative 
scarcity 
of 
browsers, 
however, 
precludes 
terming 
the 
area 
a 
savannah. 


The 
climate 
was 
probably 
warm 
and 
dry, 
not 
unlike 
the 
present 
climate 
of 
the 
area. 
The 
angularcaliche 
gravel, 
preserved 
in 
the 
greenish 
sand 
layer,
shows 
the 
presence 
of 
caliche 
caprock 
in 
Hemphillian 
times. 
Caliche 
forms 
and 
is 
preserved 
under 
semiarid 
conditions. 


The 
lake, 
where 
the 
bones 
collected, 
was 
seasonal 
and 
probably 
of 
the 
playa 
type. 
Permanent-water 
vertebrates, 
such 
as 
fishes, 
alligators, 
and 
aquaticturtles, 
are 
absent 
from 
the 
fauna. 
Of 
the 
numerous 
fossils 
found, 
only 
one 
was 
an 
articulated 
skeleton. 
Thousands 
of 
isolated 
complete 
and 
fragmentarybones 
were 
so 
scattered 
that 
they 
could 
only 
have 
been 
strewn 
over 
a 
dry 
surface. 
Tooth 
marks 
are 


present 
on 
many 
bones, 
and 
others 
show 
the 
evidenceofcrushinginthe 
jawsof 
thebone-eating 
dog.
This 
damage 
would 
scarcely 
have 
occurred 
if 
the 
bones 
were 
underwater. 


The 
Coffee 
Ranch 
local 
fauna 
includes 
speciesof 
fairly 
modern 
mammalian 
types, 
whose 
faunal 
relationships 
can 
be 
guessed 
on 
the 
basis 
of 
existingdescendent 
or 
related 
types, 
and 
extinct 
lineageswith 
no 
living 
relative. 
In 
the 
case 
of 
many 
small 
forms, 
the 
place 
they 
occupied 
in 
the 
middle 
Pliocene 
fauna 
seems 
clear. 
For 
example, 
Copemys 
is 
so 
similar, 
morphologically, 
to 
modern 
deer 
mice 
( 
Peromyscus) 
that 
one 
may 
feel 
confident 
that 
they 
were 
nocturnal 
seed-eaters 
that 
occupied 
burrows 
dug 
in 
sandy 
soil 
or 
nests 
placed 
in 
crevices 
in 
rocks. 
Similarly, 
Hypolagus 
was 
a 
rabbit 
and 
probably 
had 
much 
the 
same 
habits, 
and 
took 
the 
same 
place 
in 
the 
fauna, 
that 
the 
cottontail, 
Sylvilagus, 
occupiestoday. 
One 
cannot 
make 
similar 
assumptions 
for 
the 
hoofed 
mammals. 
Aphelops 
and 
Teleoceras 
are 
so 
distinct, 
morphologically, 
from 
living 
rhinoceroses, 


that 
one 
cannot 
judge 
the 
ecological 
relationshipsof 
the 
extinct 
forms 
from 
knowledge 
of 
the 
African 
and 
Asiatic 
species. 
This 
is 
equally 
true 
of 
the 
several 
species 
of 
camels 
and 
horses 
of 
the 
Coffee 
Ranch 
fauna. 


The 
carnivora 
of 
the 
Coffee 
Ranch 
include 
both 
modern 
and 
extinct 
lineages. 
Vulpes 
shermanensis 
was 
certainly 
much 
like 
a 
modern 
red 
fox 
in 
appearance 
and 
probably 
habits. 
It 
doubtless 
preyed 
on 
birds, 
rabbits, 
small 
rodents, 
and 
insects. 
It 
mayhave 
competed 
with 
the 
bone-eating 
dogs 
for 
carrion, 
in 
the 
manner 
that 
jackals 
compete 
with 
hyenas 
in 
the 
Old 
World 
today. 
The 
fox 
was 
probablymuch 
too 
swift, 
agile, 
and 
intelligent 
to 
be 
capturedby 
the 
thick-legged 
Osteoborus. 


The 
dentition 
and 
type 
of 
tooth 
wear 
attests 
to 
the 
hyenoid 
food 
habits 
of 
Osteoborus. 
Whether 
the 
dogs 
were 
strictly 
scavengers 
is 
doubtful, 
for 
some 
remains 
of 
small 
mammals 
were 
found 
in 
what, 
presumably, 
were 
scats. 
They 
may 
have 
been 
active 
predators 
as 
well 
as 
scavengers. 
They 
could 
scarcelyhave 
been 
swift 
runners, 
for 
their 
legs 
were 
curiouslyproportioned 
; 
long 
humerus 
and 
femur, 
short 
lower 
limbs, 
and 
stubby 
feet. 
Cursorial 
types 
usually 
show 
an 
elongation 
of 
the 
lower, 
not 
the 
upper, 
limbs. 
Perhaps 
they 
lived 
in 
packs, 
as 
some 
canids 
do 
today. 
Perhaps 
their 
activities 
were 
restricted 
to 
the 
immediate 
vicinity 
of 
water 
holes. 
This 
might 
account 
for 
the 
abundance 
of 
hyenoid 
dog 
remains 
in 
late 
Tertiary 
deposits.

Indarctos 
is 
a 
rare 
species, 
and 
perhaps 
was 
only 
an 
occasional 
visitor 
to 
the 
lake. 
This 
huge 
bear 
could 
have 
had 
little 
to 
fear 
from 
any 
other 
mammal 
of 
the 
fauna. 
Too 
little 
is 
known 
of 
its 
structure 
to 
guess 
at 
its 
habits 
or 
its 
place 
in 
the 
fauna. 


Pliotaxidae 
is 
so 
similar 
to 
the 
modern 
badger, 
except 
for 
smaller 
size, 
that 
it 
probably 
had 
similar 
habits. 
Doubtless 
it 
was 
a 
powerful 
burrower 
that 
fed 
on 
insects 
and 
small 
vertebrates, 
especially 
rodentsthatit 
dugfrom 
undergroundnests. 


Plesiogulo 
is 
the 
Pliocene 
equivalent 
of 
the 
modern 
wolverine, 
and 
may 
have 
had 
similar 
habits. 
Presumably 
it 
was 
a 
generalized 
but 
active 
carni



vore, 
preying 
on 
a 
variety 
of 
birds 
and 
small 
mammals, 
and 
even 
overcoming 
quite 
large 
prey 
when 
occasion 
permitted. 
It 
may 
have 
been 
swift 
and 
vicious 
enough 
to 
hold 
off 
carnivores 
such 
as 
Osteoborus, 
as 
the 
modern 
wolverine 
defies 
wolves 
in 
the 
treeless 
subarctic. 
It 
may, 
like 
the 
wolverine, 
also 
have 
employed 
skunk-like 
musk. 


Pseudaelurus 
was 
a 
pumalike 
cat 
and 
probablyhad 
pumalike 
habits. 
If 
so 
it 
probably 
inhabited 
the 
cliffs 
and 
canyons, 
coming 
to 
the 
water 
only 
rarely.
Prey 
of 
suitable 
size 
included 
the 
abundant 
horses 
and 
smaller 
camels, 
as 
well 
as 
peccaries 
and 
deer. 
Possible 
habits 
of 
Machairodus 
have 
been 
discussed. 
Probably 
this 
cat 
crept 
to 
a 
point 
near 
its 
prey, 
mainly 
rhinoceroses, 
and 
captured 
it 
in 
a 
bounding 
charge. 
Doubtless 
vegetation 
near 
the 


lake’s 
edge 
was 
taller 
and 
more 
suitable 
for 
cover 
than 
better 
grazed 
grassland. 
Yet, 
it 
is 
doubtful 
that 
the 
sabertooth 
was 
restricted 
to 
the 
vicinity 
of 
the 
lake 
or 
fossils 
would 
have 
been 
more 
numerous. 


The 
general 
aspect 
of 
the 
carnivore 
fauna 
of 
the 
Coffee 
Ranch 
fauna 
was, 
except 
for 
the 
bone-eatingdog 
and 
saber-toothed 
cat, 
quite 
modern. 
Foxes, 
coyotes 
and 
badgers 
occur 
in 
the 
region 
today, 
and 
bears 
and 
pumas 
lived 
nearby 
in 
historic 
times. 
The 
wolverine 
now 
occurs 
only 
far 
to 
the 
north. 
The 
hoofed-mammal 
fauna, 
in 
contrast, 
bears 
little 
resemblance 
to 
modern 
North 
American 
forms. 
The 
two 
rhinoceroses, 
four 
horses, 
four 
camels, 
dimunitive 
antilocaprid, 
deer, 
and 
mastodon, 
have 
no 
close 
relatives 
in 
our 
fauna. 
Only 
the 
peccary 
is 
similar 


to 
existing 
species. 



TABLES 



TABLE 
I 
Measurements 
of 
Osteoborus 
skulls 
MU5044 
MU3229 
MU5686 
MU7408 


Occipital 
condyles 
to 
alveoli 
of 
incisors 
Zygomatic 
breadth 
Breadth 
across 
fourth 
premolarsBreadth 
across 
second 
premolars 
Breadth 
across 
canines 
...... 
192.0 
143.0a* 
83.3 
51.0 
141.0a 
89.3 
51.3 
51.2 
79.0 
45.9 
49.8 
Breadth 
across 
incisor 
row 
31.2 
32.0 
Least 
interorbital 
breadth 
50.9 
Least 
postorbital 
constriction 
Breadth 
across 
infraorbital 
foramina, 
anteriorly 
Breadth 
across 
occipital 
condylesLength, 
incisors 
to 
back 
of 
M2 
Length, 
canines 
to 
back 
of 
M3 
...... 
... 
30.2 
47.6 
37.7 
103.8 
85.4 
41.5 
96.0 
80.0 
35.8 
* 
Approximate. 


TABLE 
2 


MeasurementsofOsteoborusupperdentitions.Figuresinparenthesesare 
measurements 
of 
18randomly-chosen,young-adultcoyotes 
{Canislatrans) 
ofunknown 
sex, 
trapped 
in 
Wichita 
County, 
Texas. 


Mean 
and 
Standard 
Coefficient 
N 
standard 
error 
deviation 
of 
variation 


Length 
PLM 
2 
8 
57.51 
± 
2.15 
5.26 
9.14 


(18) 
(59.84 
± 
0.48) 
(1.98) 
(3.30)
Length 
P--M 
1 
8 
53.81 
± 
1.39 
3.40 
6.32 
(18) 
(54.21 
± 
0.49) 
(2.00) 
(3.69)
Length 
P4-~M2 
9 
45.07 
± 
0.98 
2.76 
6.13 
(18) 
(36.84 
± 
0.32) 
(1.31) 
(3.54)
Length 
P4-Ml 
12 
38.39± 
0.54 
1.78 
4.63 
(18) 
(30.56 
± 
0.32) 
(1.30) 
(4.25)
Length 
P2 
11 
8.99 
± 
0.21 
0.67 
7.46 
(18) 
(10.96 
± 
0.16) 
(0.68) 
(6.17)
Breadth 
P2 
11 
5.20 
± 
0.11 
0.34 
6.35 
(18) 
( 
3.93 
± 
0.06) 
(0.26) 
(6.68)
Length 
P 
3 
10 
11.45 
± 
0.25 
0.76 
6.63 
(18) 
(12.42 
± 
0.19) 
(0.78) 
(6.27)
Breadth 
P 
3 
10 
5.92 
± 
0.15 
0.44 
7.39 
(18) 
( 
4.29 
± 
0.08) 
(0.32) 
(7.39)
LengthP4 
16 
24.94±0.36 
1.40 
5.61 
(18) 
(19.23 
± 
0.21) 
(0.88) 
(4.60)
Breadth 
P4 
16 
12.54 
± 
0.20 
0.79 
6.30 
(18) 
( 
9.29 
± 
0.16) 
(0.68) 
(7.29)
Length 
M1 
15 
15.91 
± 
0.34 
1.27 
7.96 
(18) 
(13.41 
± 
0.16) 
(0.67) 
(4.97)
Breadth 
M 
1 
15 
19.39 
± 
0.30 
1.12 
5.76 
(18) 
(15.18 
± 
0.15) 
(0.62) 
(4.10) 

TABLE 
3 


Measurements 
of 
Osteoborus 
lower 
dentitions. 
Figures 
in 
parentheses 
are 
measurements 
of18randomly-chosen,young-adultcoyotes(Canislatrans) 
ofunknown 
sex, 
trapped 
in 
Wichita 
County, 
Texas 


Mean 
and 
Standard 
CoefficientN 
standard 
error 
deviation 
of 
variation 


Length 
C-M: 
6 
85.50 
± 
1.85 
4.14 
4.84 


(18) 
(86.66 
± 
0 
55) 
(2.27) 
(2.62)
Length 
P 
2-M2 
8 
69.75 
± 
0 
90 
2.38 
3.42 
(18) 
(65.65 
± 
0.39) 
(1.61) 
(2.46)
Length 
P4-M2 
15 
53.91 
± 
0.48 
1.79 
3.32 
(18) 
(41.74 
± 
0.32) 
(1.31) 
(3.14)
Length 
P4—Mi 
17 
42.50 
± 
0.34 
1.33 
3.13 
(18) 
(32.13 
± 
0.28) 
(1.16) 
(3.62)
Length 
M4—M 
2 
13 
38.75 
± 
0.54 
1.86 
4.81 
(18) 
(30.62 
± 
0.23) 
(0.93) 
(3.04)
Length 
P 
2 
8 
6.81 
± 
0 
20 
0.54 
7.92 
(18) 
( 
9.71 
± 
0.13) 
(0 
55) 
(5.62)
Breadth 
P 
2 
8 
4.84 
± 
0.14 
0.38 
7.87 
(18) 
( 
4.15 
± 
0.06) 
(0.24) 
(5.86)
Length 
P:i 
12 
8.63 
± 
0 
19 
0.64 
7.39 
(18) 
(10.82 
± 
0.15) 
(0.62) 
(5.76)
Breadth 
Pi 
12 
5.74 
± 
0.11 
0.38 
6.63 
(18) 
( 
4.52 
± 
0.06) 
(0.26) 
(5.79)
LengthP4 
18 
16.66± 
0.21 
0.88 
5.30 
(18) 
(12.19 
± 
0.11) 
(0.46) 
(3.81)
Breadth 
P4 
17 
10.69 
±0.13 
0 
53 
4 
93 
(18) 
( 
5.25 
± 
0.07) 
(0.29) 
(5.51)
Length 
Mi 
18 
27.58 
± 
0.29 
1.22 
4.41 
(18) 
(21.05 
± 
0.22) 
(0.89) 
(4.24)
Breadth 
Mi 
18 
11.77 
± 
0.13 
0.52 
4 
44 
(18) 
( 
8.15 
± 
0.08) 
(0.33) 
(4.07)
Length 
M2 
12 
12.63± 
0.21 
0 
70 
5 
52 
(18) 
(10.06 
± 
0.15) 
(0.61) 
(6.08)
Breadth 
M2 
11 
8.65 
± 
0.16 
0.51 
5.89 
(18) 
( 
6.72 
± 
0.11) 
(0.44) 
(6.50) 
TABLE 
4 


Measurements 
of 
Osteoborus 
vertebrae 


Cervicals 
Thoracics 


3 
5 
7 
111213 
Length 
of 
centrum 
29.4 
18.8 
18.6 
22.3 
24.4 
27.5 
Breadth 
anterior 
zygapophyses 
32.0 
37.2 
37.7 
17.5 
18.7 
Breadth 
posterior 
zygapophyses 
27.8 
38.3 
33.2 
16.0 
15.9 
Height 
of 
centrum 
anteriorly 
12.2 
8.8 
12.3 
13.5 
14.7 
15.0 
Breadth 
of 
centrum 
posteriorly 
20.0 
20.1 
27.0 
22.0 
23.5 
23.5 



TABLE 
5 
Measurements 
of 
Osteoborus 
limb 
bones 


Number 
Mean 
Minimum 
Maximum 
Humerus 
Anteroposterior 
diameter, 
midshaft 
Transverse 
diameter, 
midshaft 
Transverse 
breadth 
of 
distal 
end 
Least 
anteroposterior 
diameter, 
distal 
end 
5 
5 
9 
9 
20.9 
15.9 
49.6 
14.3 
19.3 
14.7 
45.1 
13.3 
23.3 
16.6 
52.0 
15.5 
Radius 
Greatest 
lengthTransverse 
breadth 
of 
proximal 
end 
Anteroposteriordiameter 
of 
proximal 
end 
Transverse 
diameter, 
midshaft 
Anteroposterior 
diameter, 
midshaft 
Transverse 
breadth 
of 
distal 
end 
Anteroposterior 
diameter 
of 
distal 
end 
1 
5 
5 
4 
4 
2 
2 
160.0 
21.5 
15.2 
14.4 
10.7 
27.7 
18.8 
20.9 
14.8 
13.1 
9.3 
26.6 
18.5 
22.6 
16.0 
17.5 
12.8 
28.8 
19.0 
Ulna 
Greatest 
lengthAnteroposteriorlength 
of 
top 
of 
olecranon 
Greatest 
breadth 
of 
top 
of 
olecranon 
Anteroposterior 
lengthat 
sigmoid 
cavity 
Transverse 
diameter 
at 
tendon 
scar 
Anteroposterior 
diameter 
of 
distal 
end 
Transverse 
diameter 
of 
distal 
end 
1 
1 
1 
1 
1 
1 
1 
190 
34.9 
19.8 
22.6 
11.5 
16.8 
11.0 
Femur 
Length 
from 
top 
of 
head 
to 
distal 
end 
Least 
vertical 
constriction 
distal 
to 
head 
Anterposterior 
constriction 
at 
head 
Anteroposterior 
diameter 
at 
midshaft 
Transverse 
diameter 
at 
midshaft 
Anteroposterior 
diameter 
of 
distal 
end 
Transverse 
diameter 
of 
distal 
end 
1 
1 
1 
1 
1 
1 
1 
198.0 
18.0 
11.9 
15.0 
17.9 
36.8 
36.0 
Tibia 
Greatest 
lengthAnteroposteriordiameter 
of 
proximal 
end 
Transverse 
diameter 
of 
proximal 
end 
Anteroposterior 
diameter 
at 
midshaft 
Transverse 
diameter 
at 
midshaft 
Anteroposterior 
diameter 
of 
distal 
end 
Transverse 
diameter 
of 
distal 
end 
2 
2 
1 
3 
3 
4 
4 
174.0 
30.9 
41.1 
17.1 
14.2 
18.5 
27.0 
172.0 
30.5 
15.0 
13.1 
17.5 
25.3 
176.0 
31.3 
18.6 
15.1 
19.2 
28.1 
AstragalusAnteroposterior 
length 
Transverse 
breadth 
of 
tibial 
surface 
Least 
dorsoventral 
height 
Least 
transverse 
constriction 
at 
neck 
6 
6 
5 
6 
29.8 
18.8 
14.3 
11.9 
28.2 
17.1 
13.7 
11.3 
31.7 
20.3 
15.0 
12.8 
Calcaneum 
Anteroposterior 
length 
Greatest 
breadth 
of 
astragalar 
surfaces 
Transverse 
diameter 
of 
distal 
end 
Dorsoventral 
height 
of 
distal 
end 
1 
1 
1 
1 
55.8 
24.6 
15.0 
20.0 
Metacarpal 
11 
Greatest 
lengthGreatest 
height 
of 
proximal 
end 
Greatest 
height 
of 
distal 
end 
2 
2 
2 
47.2 
12.0 
9.7 
46.8 
11.4 
9.5 
47.5 
12.5 
9.9 



TABLE 
5—(Continued) 


Number 
Mean 
Minimum 
Maximum 
MetacarpalIII 
Greatest 
lengthGreatest 
height 
of 
proximal 
end 
Greatest 
height 
of 
distal 
end 
2 
2 
2 
61.8 
13.3 
11.4 
59.3 
13.1 
11.1 
64.2 
13.5 
11.7 
Metacarpal 
IV 
Greatest 
lengthGreatest 
height 
of 
proximal 
end 
Greatest 
height 
of 
distal 
end 
1 
1 
1 
62.1 
12.1 
8.5 
Metatarsal 
II 
Greatest 
lengthGreatest 
height 
of 
proximal 
end 
Greatest 
height 
of 
distal 
end 
1 
1 
1 
52.7 
12.7 
8.8 
Metatarsal 
III 
Greatest 
length 
1 
56.0 


TABLE 
6 


Measurements 
of 
undetermined 
canid 
skull 
(TMM41261-4) 


Length 
from 
end 
of 
nasals 
to 
end 
of 
occipital 
crest 
124.2 
Breadth 
across 
interorbital 
region 
39.0 
Breadth 
across 
postorbital 
processes 
46.9 
Postorbital 
constriction 
27.2 
Breadth 
across 
parietals 
55.0 
Breadth 
across 
occipital 
crest 
32.1 


TABLE 
7 


Measurements 
of 
teeth 
of 
Indarctos 
nevadensis 


Type 
(Macdonald, 


TMM41261-1 
MU3350 
1959) 
P4 
anteroposterior 
length 
transverse 
breadth 
25.6 
14.2 
23.5 
14.7 
anteroposterior 
length 
transverse 
breadth 
of 
trigonid 
transverse 
breadth 
of 
talonid 
45.0 
19.2 
23.7 
43.1 
19.1 
22.6 
40.5 
17.8 
20.6 
M 
2 
, 
anteroposterior 
length 
transverse 
breadth 
34.2 
25.5 
M.{, 
anteroposterior 
length 
transverse 
breadth 
19.7 
approximate20.1 



TABLE 
8 
Measurements 
of 
Pseudaelurus 
hibbardi 
Lower 
Jaws 
TMM41261-3 
TMM41261-5 


Anteroposterior 
diameter 
of 
canine 
alveolus 
..... 
17.0 
Diastema, 
C-P3 
14.8 


14.7 
P3, 
greatest 
anteroposterior 
diameter 
11.1 
P3. 
greatest 
transverse 
diameter 
5.5 
P4, 
greatest 
anteroposterior 
diameter 
15.5 
15.2 
P4, 
greatest 
transverse 
diameter 
.. 
7.4 
7.4 
Ml, 
greatest 
anteroposterior 
diameter 
-21.5 
Ml, 
greatest 
transverse 
diameter 
8.9 
Paraconid, 
maximum 
anteroposterior 
diameter 
.... 
8.2 
Protoconid, 
maximum 
anteroposterior 
diameter 
8.5 
Length, 
P3-M1 
inclusive 
47.4 
Width 
of 
mandible 
below 
center 
of 
P3 
15.2 
Width 
of 
mandible 
below 
center 
of 
Ml 
12.8 
Upper 
Carnassial, 
P4 
MU6389 
Anteroposterior 
diameter 
24.6 
Greatest 
transverse 
diameter 
12.1 


Scapula 
TMM41261-15 


Length, 
coracoid 
process 
to 
blade 
at 
end 
of 
spine 
185.0 
approximateLength 
of 
spine 
155 
0 
approximateHeight 
of 
glenoid 
fossa 
37.7 
Breadth 
of 
glenoid 
fossa 
26.7

— 


Least 
vertical 
diameter 
of 
neck 
33.7 


Humerus 
TMM41261-16 
Greatest 
length 
232 
approximateGreatest 
proximal 
breadth 
51.0 
Greatest 
anteroposterior 
diameter 
of 
head 
41.2 
Greatest 
breadth 
of 
distal 
end 
54.8 
Greatestanteroposterior 
diameter 
ofarticulatingsurface 


of 
distal 
end 
16.9 


Anteroposterior 
diameter 
of 
trochlea 
31.6 


Anteroposterior 
diameter 
of 
capitulum 
29.1 


Radius 
TMM41261-17 


Greatest 
length 
202.0 
Greatest 
proximal 
breadth 
26.3 
Greatest 
anteroposterior 
diameter, 
proximal 
18.5 
Midshaft 
breadth 
18.8 
Anteroposterior 
diameter 
at 
midshaft 
11.6 
Greatest 
distal 
breadth 
34.9 
Anteroposterior 
diameter 
at 
distal 
end 
21.0 


Astragalus 
TMM41261-19 
Greatest 
length 
39.0 
Least 
dorsopalmar 
height 
15.2 
Least 
transverse 
breadth 
of 
neck 
19.1 


Calcanea 
TMM41261-21 
TMM41261-22 
Greatest 
length 
74.6 
75.7 
Width 
at 
sustentaculum 
29.8 
29.6 


Navicular 
MU6389 
Greatest 
length 
28.4 
Greatest 
breadth 
22.0 
Least 
height 
measured 
across 
at 
notch 
11.3 



TABLE 
9 
Measurements 
of 
Machairodus 
mandibles 
TMM 
41261-8R 
TMM 
41261-8L 
TMM 
41261-9 
MU 
3527 


Length 
from 
anterior 
end 
of 
symphysis 
to 
posterior 
end 
of 
condyle.
Length 
from 
anterior 
end 
of 
outer 
flange 
to 
posterior 
end 
of 
condyle.
Length 
of 
symphysis 
measured 
alonganterior 
border. 
Least 
depth 
of 
ramus 
below 
diastema. 
Depth 
oframus 
below 
posterior 
end 
of 
M/l.
Thickness 
of 
ramus 
below 
M/l.
Height 
from 
inferior 
border 
of 
angle 
to 
summit 
of 
condyle.
Height 
from 
inferior 
border 
of 
angle 
to 
summit 
of 
coronoid 
process.
Transverse 
width 
of 
condyle.
Greatest 
depth 
of 
condyle. 
225.0 
202.0 
37.6 
45.5 
18.5 
41.2 
90.0 
43.8 
16.2 
55.5 
38.4 
46.2 
18.5 
63.1 
38.2 
44.0 
15.6 
36.7 
17.2 
TABLE 
10 
Measurements 
of 
Machairodus 
superior 
dentitions 


TMM 
TMM 
MU 
MU 
MU 
41261-8 
41261-10 
5670 
1867 
5373 


Length 
from 
anterior 
end 
of 
canine 
alveolus 
to 
posterior 
end 
of 
P4/. 
109a 
Length 
fromanterior 
end 
ofP3/ 
to 
posterior 
end 
of 
P4/. 
66.0 
73.4 


Length 
of 
diastema 
from 
posterior 
end 
ofalveolusforCtoanteriorend 
of 
alveolus 
for 
P3/. 
10.4 
9.8 
12/, 
greatest 
transverse 
diameter. 
11.9 
13/, 
greatest 
transverse 
diameter. 
12.3 
11.5 
C/, 
anteroposterior 
diameter 
at 
alveolus. 
32.4 
28.9 
C/, 
transverse 
diameter 
at 
alveolus. 
13.4 
12.7 
Length 
of 
C 
from 
alveolar 
border 
to 
tip 
of 
tooth. 
88.0a 
78.0a 
P3/, 
anteroposterior 
diameter. 
26.1 
P3/, 
greatest 
transverse 
diameter. 
10.8 
P4/, 
anteroposterior 
diameter. 
47.0 
46.5 
45.1 
P4/, 
greatest 
transverse 
diameter 
across 
protocone. 
17.6 
16.4 
16.4 
18.8 
P4/, 
greatest 
anteroposterior 
diameter 
of 
base 
of 
paracone. 
15.2 
15.4 
14.6 
P4/, 
length 
of 
metacone 
blade. 
19.5 
20.5 
20.0 



TABLE 
11 
Measurements 
of 
Marchairodus 
inferior 
dentitions 


TMMTMM 
MU 
MU 
MU 
MU 
MU 
MU 
MU 
41261-8 
41261-9 
3527 
4360 
6399 
6397 
5597 
4544 
4545 


Length 
from 
anterior 
end 
of 
C 
toposterior 
end 
ofM/l. 
128.0 
133.0 
Length 
fromanterior 
end 
of 
P/3 
toposterior 
end 
ofM/l. 
81.2a 
83.1 
81.0a 
Length 
from 
anterior 
end 
of 
P/3 
to 
posterior 
end 
ofM/l. 
59.8a 
62.5 
61.0 
Length 
of 
diastema 
measured 
between 
alveoli 
for 
C 
and 
P/3. 
34.5 
36.1 
Length 
of 
diastema 
measured 
between 
alveoli 
for 
P/3 
and 
P/4. 
3.7 
3.8 
3.4 
Length 
measured 
from 
posteriorborder 
of 
alveolus 
for 
C 
to 
an


teriorborder 
of 
alveolus 
for 
P/4. 
53.9 
57.3 
/C, 
greatest 
transverse 
diameter. 
11.4 
/C, 
greatest 
anteroposteriordiameteratbase 
ofenamel. 
12.5 
P/3, 
anteroposterior 
diameter. 
19.4 
21.7 


P/3, 
greatest 
transverse 
diameter. 
8.3 
9.5 
P/4, 
anteroposterior 
diameter. 
28.6 
28.7 
28.9 
27.9 
P/4, 
greatest 
transverse 
diameter. 
10.9 
12.0 
12.0 
12.0 
P/4, 
basal 
length 
of 
principal 
cusp 
11.1 
M/l, 
anteroposterior 
diameter. 
32.1 
35.8 
32.6 
32.3 
32 
0 
M/l, 
greatest 
transverse 
diameter. 
13.8 
12.4 
13.7 
12.4 
13.6 


TABLE 
12 


Measurements 
of 
Machairodus 
cervical 
vertebrae, 
TMM41261—B 


ATLAS 
Greatest 
width 
across 
transverse 
processes 
147.0 
Greatest 
width 
of 
anterior 
end 
across 
articulation 
for 
condyles 
68.6 
Greatest 
width 
across 
outer 
borders 
of 
articulation 
for 
axis 
62.0 
Lengthfromanteriorendofarticulationforcondylestoposteriorend 
ofarticulation 


64.7 
Length 
of 
neural 
arch 
along 
median 
line 
27.9 
Greatest 
length 
of 
transverse 
process 
taken 
oblique 
to 
axis 
of 
vertebra 
... 
74.4 
Greatest 
height 
from 
ventral 
surface 
of 
inferior 
arch 
to 
dorsal 
surface 
of 
neural 
arch 
.. 
41.0 
AXIS 
Length 
of 
centrum 
to 
tip 
of 
odontoid 
process 
82.3

... 


Depth 
of 
centrum 
measured 
normal 
to 
floor 
of 
neural 
canal 
and 
across 


20.0 
Greatest 
transverse 
width 
across 
posterior 
epiphysis 
of 
centrum 
41.0 
Width 
across 
articulating 
surface 
for 
atlas 
59.7 
3RD. 
4TH. 
5TH. 
6TH. 
7TH. 
Length 
between 
ends 
of 
zygapophyses 
56.7 
48.8 
45.6 
44.5 
Length 
of 
centrum 
normal 
to 
posterior 
face 
37.7 
36.4 
32.5 
29.3 
28.8 
Width 
across 
anterior 
zygapophyses 
58.0 
60.0 
58.2 
Width 
across 
posterior 
zygapophyses 
56.5 
57.3 
54.3 
Width 
of 
neural 
canal 
at 
anterior 
end 
20.2 
22.1 
23.1 
24.6 
26.1 
Greatesttransversewidth 
ofposteriorepiphysis 


of 
centrum 
30.3 
34.6 
32.1 
29.3 
30.0 



Greatest 
width 
across 
ends 
of 
transverse 
processes 
96.0 
94.0 
89.8 


Greatest 
length 
of 
transverse 
process 
from 
outer 
end 
to 
end 
of 
anterointernal 
projection 
of 
inferior 
lamella 
44.0 
42.5 
34.4 


Heightfromventralborder 
ofposteriorepiphysisof 
centrum 
to 
top 
of 
neural 
spine 
84.0 
Depth 
of 
centrum 
measured 
normal 
to 
floor 
of 
neural 
canal 
and 
across 
posterior 
epiphysis 
21.7 
21.3 
17.5 
20.0 
20.4 


TABLE 
13 


Measurements 
of 
Machairodus 
thoracic 
vertebrae 


facets. 
normal 
processes

face. 


outer

posterior 
posterior 
end. 
capitularend 
centrum

of

posterioranterior 
measuredtransversespine.

toof

of 
of

at

anteriornormalfacetsfacetsanterioracrossposteriorly,endsdiameterborderneural

number 
tocentrumcanal. 
of 


fromacrossacrosscanalof 
acrosstop

anteroposteriorprocess. 
bottom

to

vertebralengthcentrumwidthwidthneuralwidthcentrumneuralwidthfrom

of 


— 
of 
zygapophyses. 
zygapophyses. 
of 
of 
of 


to 
of

ThoracicGreatestzygapophysis. 
LengthGreatestGreatestHeightGreatestDepthfloorGreatestGreatesttransverseDistanceposteriorly

4-bro 
COb 
Ln 
034-Obp05b 
03 
"-4 
ro 
b 
COcoLnp 
24.7 
o 
b 
ro 
4^ 
b 
03ob 
4-ro 
'co 
034* 
03 
rob 
ro 
b 
'O4^bp 
19.3 
-rCO 
03 
4*4-Ln 
rocob 
034^Ln 
03ob 
rob 
-p 
Ln 
ro 
fo 
03 
16.3 
OOib 
Ln 
4-03 
ro 
¦*•405 
03 
*05 
roVi05 
-o 
03COo 
rob 
05 
-P03Ln 
ro 
•"4b 
ro 
*05 
ro 
"4L 
ro 
cc 
03 
Ln 
roro 
Ln 
ro 
03 
14.7 


-

'O 
ro 
"4b 
roLnLn 
Ln 
03-o 
ro 
03 
CO 
4^ 
05 
ro 
05 
ro 
-o 
roLn*05 
ro'co 
03'Ob 
ro 
ro 
-4oroP 
15.1 


4^ 
03robo 
rol 
c£) 
P 
ro 
ro 
ro 
CO 
ro 
ro 
Ln 
LnbP 
K3 
LPb 
0305 
co 
o 
roLn 
05 
03 
03 
03 
ro 
ro 
03 
4-b 
4^COLn 
03 
05 
4^O 
ro 
4^ 
05 
to 
-p 
oo 
03Ln 
03 
K3 
Ln 
4^03bp 
-pb 



TABLE 
14 
Measurements 
of 
Machairodus 
lumbar 
vertebrae 


of

faceof 


line. 
zygapophysis. 
floorepiphysis. 
anterior 
spine.

of

of

facezygapophysis. 
centrum. 
neural

normalmediannormalborderof 


to 
alonganteriorposteriormetapophysis. 
posteriorepiphysisto 
posteriorventraltop

numbermeasuredfromendof 
measuredacross

andofto

acrossacross


to

vertebracentrumepiphysislengthwidthwidthposteriorcentrumand 
middlecentrum

of 


— 
of 
canalfrom

ofof

LumbarLengthposteriorGreatestmetapophysisGreatestGreatestWidthDepthneuralHeightepipnysis

.44*1—105 
o4^Onbo 
NO 
OO 
05 
ooOn 
NOlo 
NObNO 
4^05S 
05Ons 
4>-4>-bo 
NOCO 
NO 
OOOnbn 
NO 
OO 
4^ 
NO 
oo 
4^CO 
OO 
0505 
05 
4^c_n 
05 
NOCO 
OO 
OOOnbo 
NO 
OO 
*-4lo 
4^ 
Onoo 
*“* 
05CO4-4^On 
CO 
NOCO 
o 
OO•^J 
OO 
NO 
o 


k 


“

On 
On4^ 
05 
o 
05 
4*4^ 
NO 
oo 
HooCO 
NO 
NO 
OO 
COCO 
lo 
05 
C_Ji 
05bn 
05lo 
*-* 
4^4^ 
4* 
ooo 
ooCObn 
NO 
05 
CO 
CO 
C_nO 
o>IObn 
4^CO 
4* 
CnLnlo 
OO 
SCbo 
no 
l>0 
'-4CT)lo 


TABLE 
15 


Measurements 
of 
Machairodus 
sacrum, 
TMM41261—8 


Greatest 
length 
measured 
parallel 
to 
median 
line. 
112.0 
Greatest 
width 
at 
anterior 
end 
and 
at 
outer 
surfaces 
for 
ilia. 
85.2 
Greatest 
width 
of 
third 
sacral 
vertebra 
across 
transverse 
processes. 
57.0 
Greatest 
width 
between 
dorsal 
surfaces 
of 
anterior 
zygapophyses. 
56.4a 
Greatest 
depthofcentrumoffirstsacralvertebrameasurednormaltofloor 


of 
neural 
canal 
and 
across 
anterior 
epiphysis. 
22.8 
Greatest 
distance 
from 
dorsal 
margin 
of 
anterior 
zygapophysis 
to 
ventral 
border 
of 
surface 
joining 
with 
ilium. 
59.0 


TABLE 
16 


Measurements 
of 
Machairodus 
caudal 
vertebrae, 
TMM41261—8 


1. 
2. 
Greatest 
length 
of 
centrum 
measured 
parallel 
to 
central 
axis. 
26.5 
26.8 
Width 
across 
anterior 
zygapophyses. 
19.1 
23.9 
Width 
across 
posterior 
zygapophyses. 
14.4 
24.0 
Height 
from 
middle 
of 
ventral 
border 
of 
centrum 
to 
top
of 
neural 
spine 
or 
neural 
arch. 
31.0 
34.1 



TABLE 
17 
Measurements 
of 
Machairodus 
scapula, 
TMM41261-8 
Greatest 
width 
of 
articulating 
end 
measured 
across 
glenoid 
cavity.
Greatest 
transverse 
diameter 
across 
glenoid 
cavity.
Least 
width 
of 
neck 
across 
articulating 
end. 
right 
62.1 
35.0 
52.3 
left 
61.1 
35.1 
52.4 
TABLE 
18 
Measurements 
of 
Machairodus 
humerus 


TMM 
TMM 
MU 
MU 
41261-8R 
41261-8L 
6865 
6869 


Greatest 
length 
measured 
parallel 
to 
longitudinal 
axis. 
Greatest 
transverse 
diameter 
of 
proximal 
extremity. 
Greatest 
anteroposterior 
diameter 
of 
proximal 
extremity.
Transverse 
diameter 
at 
middle 
of 
shaft. 
23.4 
314.0 
21.5 
63.3 
86.4 
Anteroposterior 
diameter 
at 
middle 
of 
shaft. 
Greatest 
width 
of 
distal 
extremity. 
Least 
anteroposterior 
diameter 
of 
articulating 
surface 
for 
ulna. 
38.7 
71.0 
24.8 
37.1 
73.8 
24.8 
74.3 
24.0 
74.6 
23.8 
TABLE 
19 
Measurements 
of 
Machairodus 
ulna 
TMM 
TMM 
MU 
41261-8R 
41261-8L 
6870 
Greatest 
length 
measured 
parallel 
to 
longitudinal 
axis 
of 
ulna. 
Greatest 
width 
of 
posterior 
surface 
of 
olecranon 
process. 
Greatest 
transverse 
width 
of 
greater 
sigmoid 
cavity.
Anteroposterior 
diameter 
of 
shaft 
at 
proximal 
end 
of 
tendon 
scar. 
Transverse 
diameter 
of 
shaft 
at 
proximal 
end 
of 
tendon 
scar. 
Greatest 
anteroposterior 
diameter 
of 
distal 
extremity. 
Greatest 
width 
of 
distal 
extremity. 
316.0 
35.0 
38.1 
29.7 
18.0 
27.0 
37.4 
35.6 
26.9 
353a 
44.5 
30.0a 
20.7 
TABLE 
20 
Measurements 
of 
Machairodus 
radius 
TMM 
MU 
MU 
41261-8 
5533 
5271 
Length 
measured 
along 
internal 
border. 
Long 
diameter 
of 
proximal 
end. 
Greatest 
diameter 
taken 
at 
right 
angles 
to 
long 
diameter 
of 
proximal 
end. 
Greatest 
diameter 
of 
shaft 
at 
middle. 
267.0 
33.0 
25.7 
25.1 
247.5 
29.8 
24.0 
23.6 
285.0 
36.0 
26.4 
23.2 
Least 
diameter 
of 
shaft 
at 
middle. 
15.2 
14.8 
16.3 
Greatest 
width 
at 
distal 
end 
taken 
normal 
to 
internal 
face. 
50.4 
43.6 
61.7 
Greatest 
thickness 
of 
distal 
end. 
33.5 
28.7 
30.9 



TABLE 
21 


Measurements 
of 
Machairodus 
scapholunar 


MUMU 
MU 
MU 
5417 
7911 
7975 
7941 


Greatest 
transverse 
diameter 
measured 
normal 
to 
external 
border 
of 
proximal 
surface. 
56.0 
51.1 
47.4 
48.1 


Greatest 
dorsopalmar 
length. 
35.6 
38.1 
31.3 
31.6 


Greatest 
proximal 
distal 
diameter. 
28.8 
25.6 
24.7 
24.4 


TABLE 
22 


Measurements 
of 
Machairodus 
pisiform, 
MU7867 


Greatest 
length 
from 
and 
measured 
normal 
to 
contact 


edge 
of 
ulnar 
and 
unciform 
facets 
to 
the 
head. 
35.5 
Long 
diameter 
of 
articulating 
end. 
23.0 
Long 
diameter 
of 
head. 
19.0 
Greatest 
proximal 
diameter 
of 
head 


normal 
to 
long 
diameter. 
13.2 


TABLE 
23 


Measurements 
of 
Machairodus 
metacarpal 
I, 
MU5929 


Greatest 
length 
normal 
to 
proximal 
end. 
34.0 
Greatest 
width 
of 
proxima 
1 
end. 
22.7 
Transverse 
width 
of 
distal 
articulating 
surface. 
17.5 


TABLE 
24 


Measurements 
of 
Machairodus 
metacarpals 


at

at

diameter 
diameter 
diameter 


end 
end 
of

length 
transversedorsoventraldiametershaft 
diametershaft 
transverseshaft 


of 
of

proximalproximalend

distal

GreatestGreatestGreatestTransversemiddleDorsoventralmiddleGreatest

ofofat 


sn>Pop 
*-s 
p 
M 


Hss4“-roO') 
CO 
CO 
roo 
CD 
roCO 
'O 
p 
P 
pCD 
d4*-CO 
ro 
o 
o 
ropbo 
COLn 
COLn 
roPro 
dLnCO2 
opo 
ro 
Ln 
rop 
CD 
-Ln 
pKo 
roo 
Ln 


CDPOP 
P 


p

Hs4* 
CD 
CO 
ro 
CD4-rocoo 
CO 
CO 
ro 
o 
sccnCO 
OJ 
ro 
CD 
ro 
ro 
roro 
CD 
Cjn 
4* 
ro 
CO 
roro 
CO 
SdC-nCO 
ro 
'O 
CD 
CDbo 
ro 
CD 
p 
CD 
cofo 
ro 
CO 


sro 
Pop 
op 
sdO')CO 
C-n 
O 
Lo 
pLo 
roo 
p 
roLn 
CDro 


<dPo 
*“5 
cL 


Hss•4^ 
O? 
CO 
COCO 
CO 
p4* 
pLo 
o4* 
roo 
oo 



TABLE 
25 
Measurements 
of 
Machairodus 
femur 


TMM 
TMM 
TMM 
41261-8R 
41261-8L 
41261-13 


Greatest 
length 
from 
top 
of 
greater 
trochanter 
to 
distal 
condyles, 
measured 
parallel 
to 
long 
axis. 
Transverse 
diameter 
of 
proximal 
end, 
outer 
face 
of 
greater 
trochanter 
to 
inner 
side 
of 
head, 
taken 
normal 
to 
median 
longitudinal 
plane.
Greatest 
anteroposterior 
diameter 
of 
head. 
Transverse 
diameter 
of 
shaft 
at 
middle. 
74.1 
35.3 
29.0 
73.3 
35.1 
28.4 
342.0 
79.4 
35.6 
30.9 
Anteroposterior 
diameter 
of 
shaft 
at 
middle. 
25.1 
28.0 
26.6 
Greatest 
width 
of 
distal 
extremity.
Greatest 
anteroposterior 
diameter 
of 
distal 
extremity 
at 
right 
angles 
to 
longitudinalaxis 
of 
femur. 
Greatest 
width 
of 
intercondylar 
notch. 
Greatest 
width 
of 
articular 
surface 
of 
inner 
condyle. 
65.9 
63.2 
14.5 
26.0 


TABLE 
26 


Measurements 
of 
Machairodus 
tibia. 


TMM41261-14 
MU3671 


Greatest 
length 
measured 
parallel 
to 
long 
axis. 
284.0 
293.0 
Greatest 
transverse 
diameter 
ofproximal 
end. 
64.5 
68.4 
Transverse 
diameter 
of 
shaft 
at 
middle. 
26.2 
30.8 
Greatest 
transverse 
diameter 
of 
distal 
end. 
50.7 
52.5a 
Greatest 
anteroposterior 
diameter 
of 
distal 
end. 
31.5 
31.0a 


TABLE 
27 


Measurements 
of 
Machairodus 
astragalus. 


TMMMU 
MU 
MU 
41261-8 
5743 
5744 
5764 


Greatest 
length. 
Greatest 
width. 
47.1 
48.5 
51.3 
51.3 
49.2 
50.5 
47.3 
48.7 
Least 
distance 
across 
neck. 
19.1 
21.0 
19.7 
20.9 
Greatest 
diameter 
of 
head. 
28.8 
30.7 
32.5 
TABLE 
28 
Measurements 
of 
Machairodus 
calcaneum. 
MU 
MU 
MU 
7562 
7644 
7536 
Greatest 
length.
Greatest 
width 
measured 
across 
90.7 
83.5 
94.4 
astragalar 
facets 
As1 
and 
As2 
. 
Greatest 
width 
across 
cuboid 
surface 
36.7 
40.5 
measured 
from 
astragalar 
facet 
As3 
to 
outer 
side. 
29.3 
29.1 
31.1 



TABLE 
29 
Measurements 
of 
Machairodus 
cuboid, 
MU7949 
Greatest 
proximodistal 
diameter. 
Greatest 
transverse 
diameter. 
Greatest 
dorsoplantar 
length. 
28.7 
22.7 
29.1 
TABLE 
30 
Measurements 
of 
Machairodus 
navicular, 
MU5963 
Dorsoplantar 
length.
Transverse 
diameter. 
35.6 
26.1 
TABLE 
31 
Measurements 
of 
Machairodus 
metatarsals 
s 
s 
S 
s 
S 
s 
H 
H 
d 
d 
d 
d 
d 
5 
d 
d 
d 
s 
cd 
CD 
CO 
O') 
CO 
CD 
CO 
CO 
cr, 
co 
CD 
CO 
oo 
CO 
oo 
oo 
sp 
P 
Metatarsalsnumber 
oo 
OO 
O 
oo 
cr 
oo 
p 
CO 
CO 
P 
p 
oo 
ro 
CD 
ro 
CD 
00r 
CO 
?0 
o 
£ 
ro 
o 
o 
O 
o 
o 
£ 
Greatestlength. 
oo 
OI 
Cn 
oo 
p 
p 
ro 
Ln 
oo 
CD 
fo 
00 
fo 
bn 
br 
Ln 
P 
Lb 
bj 
b 
p 
P 
ro 
ro 
ro 
Greatesttransversediameter 
COLn 
O') 
P 
lo 
bi 
PLn 
P 
P 
P 
P 
b 
OO 
p 
b 
ofproximalend. 
0) 
sro 
Kro 
s(T> 
P 
ro 
ro 
P 
P 
ro 
P 
P 
ro 
ro 
ro 
o 
ro 
ro 
oo 
o 
P 
P 
Greatestdorsoventraldiameter 
OO 
P 
< 
bo 
fo 
*¦* 
P^ 
HH 
< 
fo 
£-HH 
CD 
b 
'co 
P 
*1 
ofproximalend. 
CO 
b 
l. 
l 
. 
,. 
, 
. 
H_4 
Transversediameterat 
roLn 
robo 
Ln 
CD 
b 
bo 
CD 
b 
br 
b 
b 
middleof 
shaft. 
(ji 
oo 
oo 
oo 
Dorsoventraldiameterat 
b 
br 
bo 
L>0 
Ln 
Ln 
p 
Ln 
b 
b 
middleof 
shaft. 
ro 
o 
ro 
ro 
(3 
ro 
o 
Greatesttransversediameter 
bo 
cd 
fo 
cd 
P 
b 
bo 
CD 
bo 
b 
b 
bo 
bj 
at 
distalendof 
shaft. 



Literature 
Cited 


Burt, 
W. 
H. 
1931. 
Machairodus 
catocopis 
Cope 
from 
the 
Pliocene 
of 
Texas. 
Univ. 
California 
Pubis. 
Geol. 
Sci. 


20: 
261-292. 
Churcher, 
C. 
S. 
1968. 
The 
affinities 
of 
Dinobastis 
serus 
Cope 
1893. 
Quaternaria, 
8: 
263-275. 


Evans, 
Glen 
L. 
1961. 
The 
Friesenhahn 
Cave. 
Bull. 
Texas 
Memorial 
Mus., 
2 
: 
5-22. 


Evernden, 
J. 
F., 
D. 
E. 
Savage, 
G. 
H. 
Curtis 
and 
G. 
T. 
James. 
1964. 
Potassium-argon 
dates 
and 
the 
cenozoic 
mammalian 
chronology 
of 
North 
America. 
Amer. 
Jour. 
Sci., 
262: 
145-198. 


Frick, 
C. 
1921. 
Extinct 
vertebrate 
faunas 
of 
the 
badlands 
of 
Bautista 
Creek 
and 
San 
Timoteo 
Canyon,
southern 
California. 
Univ. 
California 
Pubis. 
Geol. 
Sci.,

12: 
277-424. 
Flall, 
E. 
R. 
1944. 
A 
new 
genus 
of 
American 
Pliocene 
badger 
with 
remarks 
on 
the 
relationship 
of 
badgers 
of 
the 
northern 
hemisphere. 
Carnegie 
Inst. 
Wash. 
Publ.,

551: 
9-23. 
Hibbard, 
C. 
W. 
1934. 
Two 
new 
genera 
of 
felidae 
from 
the 
middle 
Pliocene 
of 
Kansas. 
Trans. 
Kansas 
Acad. 
Sci., 
37: 
239-255. 


. 
1937. 
Additional 
fauna 
of 
Edson 
quarry 
of 
the 
middle 
Pliocene 
of 
Kansas. 
Amer. 
Mid. 
Nat., 
18: 
460-464. 


Hesse, 
C. 
J. 
1936. 
A 
Pliocene 
vertebrate 
fauna 
from 
Optima, 
Oklahoma. 
Univ. 
California 
Pubis. 
Geol. 
Sci.,

24: 
57-70. 
Kitts, 
D. 
B. 
1958. 
Nimravides, 
a 
new 
genus 
of 
felidae 
from 
the 
Pliocene 
of 
California, 
Texas 
and 
Oklahoma. 
Jour. 
Mamm., 
39: 
368-375. 


Kurten, 
B. 
1963. 
Fossil 
bears 
from 
Texas. 
Texas 
Memorial 
Mus., 
Pearce-Sellards 
Ser. 
1: 
1-15. 


Long, 
C. 
A. 
1965. 
Comparison 
of 
juvenile 
skulls 
of 
the 
mustelid 
genera 
Taxidae 
and 
Meles, 
with 
comments 
on 
the 
subfamily 
Taxidiinae 
Pocock. 
Amer. 
Mid. 
Nat., 
74; 
225-232. 


Macdonald, 
J. 
R. 
1948a. 
A 
new 
species 
of 
Pseudaelurus 
from 
the 
lower 
Pliocene 
of 
Nebraska. 
Univ. 
California 
Pubis. 
Geol. 
Sci., 
28: 
45-52. 


. 
1948b. 
The 
Pliocene 
carnivores 
of 
the 
Black 
Hawk 
Ranch. 
Univ. 
California 
Pubis. 
Geol. 
Sci., 
28: 
53-80. 


1959. 
The 
middle 
Pliocene 
mammalian 
fauna 
from 
Smiths 
Valley, 
Nevada. 
Jour. 
Paleo., 
33: 
872-887. 


Martin, 
H. 
T. 
1928. 
Two 
new 
carnivores 
from 
the 
Pliocene 
of 
Kansas. 
Jour. 
Mamm., 
9; 
233-236. 


Matthew, 
W. 
D. 
1932. 
A 
review 
of 
the 
rhinoceroses 


with 
a 
description 
of 
Aphelops 
material 
from 
the 
Pliocene 
of 
Texas. 
Univ. 
California 
Pubis., 
Geol. 
Sci., 
20: 
411-480. 


Matthew, 
W. 
D., 
and 
R. 
A. 
Stirton. 
1930a. 
Osteologyand 
affinities 
of 
Borophagus. 
Univ. 
California 
Pubis. 
Geol. 
Sci., 
19: 
171-216. 


—. 
1930b. 
Equidae 
from 
the 
Pliocene 
of 
Texas. 
Univ. 
California 
Pubis. 
Geol. 
Sci., 
19: 
349-369. 


Mawby, 
J. 
E. 
1965. 
Machairodonts 
from 
the 
late 
Cenozoic 
of 
the 
Panhandle 
of 
Texas. 
Jour. 
Mamm., 
46: 
573


587. 
Meade, 
G. 
E. 
1961. 
The 
Saber-toothed 
Cat, 
Dinobastis 
serus. 
Bull. 
Texas 
Memorial 
Mus., 
2 
:23—60. 


Merriam, 
J. 
C., 
C. 
Stock 
and 
C. 
L. 
Moody. 
1916. 
An 
American 
Pliocene 
bear. 
Univ. 
California 
Pubis. 
Geol. 
Sci., 
10: 
87-109. 


Merriam, 
J. 
C. 
and 
C. 
Stock. 
1932. 
The 
felidae 
of 
the 
Rancho 
la 
Brea. 
Carnegie 
Inst. 
Wash. 
Publ., 
422: 
1— 


231. 
Reed, 
L. 
C. 
and 
O. 
M. 
Longnecker, 
Jr. 
1932. 
The 
geology 
of 
Hemphill 
County, 
Texas. 
Univ. 
Texas 
Bull.,
3231: 
1-98. 


Savage, 
D. 
E. 
1941. 
Two 
new 
middle 
Pliocene 
carnivores 
from 
Oklahoma 
with 
notes 
on 
the 
Optima 
fauna. 
Amer. 
Mid. 
Nat., 
25: 
692-710. 


Sellards, 
E. 
H. 
1916. 
Fossil 
vertebrates 
from 
Florida: 
a 
new 
Miocene 
fauna; 
new 
Pliocene 
species; 
the 
Pleistocene 
fauna. 
Ann. 
Rept. 
Florida 
Geol. 
Surv., 
8:77-119. 


VanderHoof, 
V. 
L. 
and 
J. 
Greggory. 
1940. 
A 
review 
of 
the 
genus 
Aelurodon. 
Univ. 
California 
Pubis. 
Geol. 
Sci., 
25: 
143-164. 


Webb, 
D. 
S. 
1965. 
The 
osteology 
of 
Camelops. 
Bull. 
Los 
Angeles 
CountyMus., 
Science, 
1 
: 
1-54. 


Wood, 
H. 
E. 
(Chairman), 
et 
al. 
1941. 
Nomenclature 
and 
correlation 
of 
the 
North 
American 
continental 
Tertiary. 
Geol. 
Soc. 
Amer. 
Bull., 
52: 
1-48. 



Bulletins 
of 
the 
Texas 
Memorial 
Museum 


Ann 
Suhm 
and 
Edward 
B. 
Jelks. 
Published 
jointly 
by 
the 
Texas 
Me-

No. 
1. 
Mylohyus 
nasutus, 
Long-nosed 
Peccary 
of 
the 
Texas 
Pleistocene, 
byE. 
L. 
Lundelius, 
Jr.. 
1960 
... 
$l.OO 
No. 
2. 
The 
Friesenhahn 
Cave 
(Part 
I), 
by 
Glen 
L. 
Evans; 
The 
Saber-
toothed 
cat, 
Dinobastis 
serus 
(Part 
II), 
by 
Grayson 
E. 
Meade, 
1960 
1.00 
No. 
3. 
ABibliography 
of 
Recent 
Texas 
Mammals, 
By 
Gerald 
G. 
Raun, 
1962 
1.00 
No. 
4. 
Handbook 
of 
Texas 
Archeology: 
Type 
Descriptions, 
edited 
by 
Dee 


morial 
Museum 
and 
the 
Texas 
Archeological 
Society 
Out 
of 
print* 


No. 
5. 
Salvage 
Archeology 
of 
Canyon 
Reservoir: 
The 
Wunderlich, 
Footbridge, 
and 
Oblate 
Sites 
by 
Leroy 
Johnson, 
Jr., 
Dee 
Ann 
Suhm, 
and 
Curtis 
Tunnell, 
1962 
-2.00 


No. 
6. 
The 
Ethnography 
and 
Ethnology 
of 
Franz 
Boaz, 
by 
Leslie 
A. 
White, 
1963 
--2.00 


No. 
7. 
Fossil 
Vertebrates 
from 
Miller's 
Cave, 
Llano 
County, 
Texas, 
byThomas 
Patton, 
1963 
-2.00 


No. 
8. 
Interactions 
Between 
a 
Bisexual 
Fish 
Species 
and 
its 
Gynogenetic 
Sexual 
Parasite, 
by 
Clark 
Hubbs, 
1963 
2.00 


No. 
9. 
Oedaleops 
campi 
(Reptilia: 
Pelycosauria) 
A 
new 
genus 
and 
species 
from 
the 
Lower 
Permian 
of 
New 
Mexico, 
and 
the 
family 
Eothyrididae,
by 
Wann 
Langston, 
Jr., 
1965 
LOO 


No. 
10. 
Blancan 
Mammalian 
Fauna 
and 
Pleistocene 
Formations, 
HudspethCounty, 
Texas, 
by 
William 
Samuel 
Strain, 
1966 
2.00 


No. 
11. 
A 
Population 
of 
Woodrats 
(Neotoma 
micropus) 
by 
G. 
G. 
Raun, 
1966 
2.00 


No. 
12. 
Toward 
a 
Statistical 
Overview 
of 
the 
Archaic 
Cultures 
of 
Central 
& 
Southwestern 
Texas, 
by 
Leßoy 
Johnson, 
Jr., 
1967 
2.00 
No. 
13. 
GeographicVariationsinSurvivalofHybridsBetweenEtheostomatine 
Fishes, 
by 
C. 
L. 
Hubbs, 
1967 
2.00 
No. 
14. 
A 
Lipan 
Apache 
Mission. 
San 
Lorenzo 
De 
La 
Santa 
Cruz, 
1762-1771, 
by 
Curtis 
1). 
Tunnell 
and 
W. 
W. 
Newcomb, 
Jr., 
1969 
3.00 


* 
Reprints 
available 
at 
Texas 
Memorial 
Museum 
9.00