87 Palaeont. afr., 14. 8 7 - 9 2 .

1972

A NEW SPECIES O F MACHAERODONT FROM MAKAPANSGAT

M

G . E . Collings
ABSTRACT The skull and associated mandible belonging to a new species of machaerodont, Megantereon problematicus Collings is described. The specimen came f r o m the lowermost horizon of banded and contaminated travertines at the Makapansgat Limeworks. A n estimate o f its relationship w i t h other sabre t o o t h cats is given.

CLASSIFICATION
Class Order Sub-order Infra-order Family Sub-family Genus Species Mammalia Carnivora Fissipedia Aeluroidea Felidae Machaerodontinae Megantereon Croizer and Jobert Megantereon problematicus nov.

DIAGNOSIS
A l a r g e Megantereon, w h o s e carnassials are longer a n d narrower t h a n any previously described species o f t h i s genus. T h e carnassial w i d t h t o l e n g t h r a t i o is 1 : 3 , as o p p o s e d t o a l l o t h e r d e s c r i b e d Megantereon species, w h i c h h a v e a carnassial w i d t h to length ratio n o t exceeding 1 : 2 .

2 , a n d 3 ) . I t was r e c o v e r e d b y M r . J . W . K i t c h i n g , i n 1958, f r o m the layer o f banded and cont a m i n a t e d t r a v e r t i n e s a t t h e site o f t h e s o - c a l l e d " A n c i e n t E n t r a n c e " at t h e Makapansgat L i m e w o r k s . T h e s e i m p u r e t r a v e r t i n e s are i m m e d i a t e l y o v e r l a i n b y t h e 'basal r e d b r e c c i a ' ( c a l c i f i e d r e d m u d o f B r a i n 1 9 5 8 ) w h i c h is, i n t u r n , s u c c e e d e d b y the relatively t h i n layer o f exceedingly boner i c h grey breccia. T h e s k u l l is c o n s i d e r a b l y distorted, having been subject t o lateral compression d u r i n g f o s s i l i z a t i o n , a n d a great deal o f t h e c r a n i u m is m i s s i n g . T h e t e e t h a n d m a n d i b l e h o w e v e r are p e r f e c t l y p r e s e r v e d , a n d e n a b l e a n a c c u r a t e d i a g n o s i s o f t h e genus a n d species. T h e p r e s e r v e d f e a t u r e s o f t h e s k u l l , s u c h as t h e small c o r o n o i d process a n d t h e specialisation o f the t e e t h , are i n a g r e e m e n t w i t h t h o s e o f t h e genus Megantereon. However, specimen M . 8 2 8 0 is d e f i n e d as a n e w species o f Megantereon o n the basis o f c e r t a i n d e n t a l c h a r a c t e r s , w h i c h h a v e n o t been observed i n other machaerodonts previously described. T h e general p r o p o r t i o n s o f t h e l o w e r carnassials i n a l l o t h e r species o f Megantereon are about twice the w i d t h to the length, whilst i n s p e c i m e n M . 8 2 8 0 , t h e l e n g t h is a p p r o x i m a t e l y three times the w i d t h . A c o m p a r i s o n o f its t e e t h measurements w i t h those o f some other Megantereon species a n d t h e S o u t h A f r i c a n sabre t o o t h cats is g i v e n i n T a b l e 1 . A brief description o f the dentition n o w f o l l o w s ( F i g s . 17). T h e u p p e r canines are m u c h flattened laterally, crenulated, moderately long, b u t o n l y s l i g h t l y c u r v e d ; P^ is m u c h r e d u c e d ; P"^ has a v e s t i g i a l p r o t o c c n e a n d a w e l l d e v e l o p e d p a r a s t y l e ; M ^ is s m a l l a n d r o u n d e d . T h e m a n d i b l e has a d e e p s y m p h y s i s , a m e n t a l crest a n d a m e n t a l p r o c e s s ; t h e c o r o n o i d p r o c e s s is m u c h r e d u c e d . T h e l o w e r canines are c o m p a r a t i v e l y s m a l l a n d r a i s e d a b o v e t h e l e v e l o f t h e c h e e k t e e t h ; P3 is m u c h r e d u c e d ; P4 has w e l l d e v e l o p e d accessory cusps; M l has a v e s t i g i a l m e t a c o n i d .

INTRODUCTION
T h e P l i o c e n e / P l e i s t o c e n e sabre t o o t h cats are a n assemblage o f u n r e l a t e d genera, w h i c h are assumed t o have r a d i a t e d f r o m an earlier Oligocene f e l i d s t o c k , r e p r e s e n t e d b y Dinictis and Hoplophoneus ( M a t t h e w 1 9 1 0 ) . B o t h o f these s h o w e d m a c h a e r o d o n t tendencies, b u t i t was p r o b a b l y t h e c o m p a r a t i v e l y u n s p e c i a l i s e d Dinictis w h i c h led to t h e v a r i e d g r o u p o f l a t e r sabre t o o t h s , as w e l l as t h e m o d e r n f e l i d s . T h e P l e i s t o c e n e sabre t o o t h s c a n b e separated i n t o a n u m b e r o f groups, each f o l l o w i n g s l i g h t l y d i f f e r e n t lines o f m a c h a e r o d o n t m o d i f i c a t i o n . Megantereon a n d Smilodon are s u p e r f i c i a l l y v e r y s i m i l a r , a n d m a y b e said t o h a v e b e e n t h e m o s t s p e c i a l i s e d o f t h e P l e i s t o c e n e sabre t o o t h s . H o w e v e r , o n closer analysis, t h e y appear q u i t e unrelated, a n d c o u l d be examples o f parallel evolution. Megantereon, Machaerodus and Therailurus b e l o n g e d t o t h e O l d W o r l d , a n d seem t o h a v e c o - e x i s t e d i n t h e same h a b i t a t s , w h i l e Smilodon, so f a r , has o n l y b e e n f o u n d i n t h e N e w World.

DESCRIPTION
The specimen. N o . M . 8 2 8 0 o f the Bernard P r i c e I n s t i t u t e f o r P a l a e o n t o l o g i c a l R e s e a r c h , is t h e i n c o m p l e t e s k u l l a n d m a n d i b l e o f a large machaerod o n t b e l o n g i n g t o t h e g e n u s Megantereon (Figs. 1,

DISCUSSION
T h e inner surface o f b o t h upper and l o w e r t o o t h rows had been w o r n extensively to f o r m a c o n t i n u o u s o b l i q u e shearing plane and the i n d i v i d u a l cusps h a d b e e n o b l i t e r a t e d t o f o r m o n e

88 l o n g i t u d i n a l ridge. This indicates that the animal was o l d a t t h e t i m e o f d e a t h . O b s e r v a t i o n s o n t h e t e e t h r e v e a l e d t h a t t h e a n i m a l also s u f f e r e d f r o m s o m e d e n t a l d e f e c t s . T h e r i g h t u p p e r c a n i n e has a g r e a t e r a n t e r o - p o s t e r i o r l e n g t h t h a n t h e l e f t orie a n d t h i s seems t o b e t h e r e s u l t o f a g e n e t i c a b n o r m a h t y . T h i s c a u s e d i t t o shear u n n a t u r a l l y against t h e l o w e r c a n i n e i n j a w c l o s u r e a n d m a y h a v e c o n t r i b u t e d t o d i f f e r e n t i a l w e a r o n b o t h sides of the cheek teeth. P3 a n d P4 o n t h e l e f t side h a d b e e n b r o k e n i n l i f e , a n d t h e l e f t P ^ , w h i c h o p p o s e s P 3 , is also b r o k e n ( c o m p a r e F i g s . 5 a n d 6 ) . T h i s m a y have i n f l u e n c e d t h e a n i m a l t o use t h e r i g h t side m o r e w h i l e m a s t i c a t i n g , as t h e r i g h t t e e t h a p p e a r m o r e w o r n than the left. T h e g e o l o g i c a l age o f s p e c i m e n M . 8 2 8 0 is e s t i m a t e d as L o w e r P l e i s t o c e n e . T h e f o s s i l f a u n a o f M a k a p a n s g a t shows a s i m i l a r i t y w i t h t h e o l d e s t h o r i z o n o f t h e O m o beds i n East A f r i c a . These have b e e n d a t e d b y t h e PotassiumArgon m e t h o d as 4 , 1 m i U i o n years o l d , w i t h t h e y o u n g e s t h o r i z o n b e i n g 1,4 m i l l i o n y e a r s o l d ( C o o k e 1 9 7 0 , pers. c o m m . ) . This makes the O m o beds older t h a n the E u r o p e a n V i l l a f r a n c h i a n w i t h w h i c h i t was p r e viously thought they corresponded. However, i n v i e w o f t h e e x t r e m e l y specialised d e n t i t i o n o f Megantereon problematicus i t is m o r e l i k e l y t h a t t h e M a k a p a n s g a t f o s s i l f a u n a a c c u m u l a t e d somew h a t l a t e r , a t a n age t h a t c o r r e s p o n d s t o t h e accepted Lower Pleistocene. Although the E u r o p e a n species o f Megantereon are r e g a r d e d as L o w e r P l e i s t o c e n e ( S t i r t o n 1 9 5 1 ) , t h e r e are des c r i p t i o n s o f o t h e r species f r o m t h e U p p e r P l i o c e n e of India (Pilgrim 1932, Colbert 1935), and f r o m the L o w e r PHocene of China (Young 1935, T e i l h a r d de C h a r d i n 1 9 3 9 , Chang 1957) w h i c h gives t h i s genus a l o n g r a n g e i n t i m e . I t is q u i t e p o s s i b l e t h a t a n A f r i c a n species c o u l d h a v e e v o l v e d independently f r o m a Pliocene group and reached a h i g h degree o f s p e c i a l i s a t i o n b y L o w e r P l e i s t o c e n e times. SUMMARY 1. T h e skull of a machaerodont from the M a k a p a n s g a t L i m e w o r k s is d e s c r i b e d . 2 . I t is a s c r i b e d t o a n e w species, Megantereon problematicus C o l l i n g s , o n t h e basis o f i t s unique dental pattern. 3 . A d e t e r m i n a t i o n o f t h e age o f t h e d e p o s i t is e s t i m a t e d as p r o b a b l y L o w e r P l e i s t o c e n e . ACKNOWLEDGEMENTS T h i s w o r k was c o m p l e t e d as p a r t o f t h e h o n o u r s course i n P a l a e o n t o l o g y at t h e U n i v e r s i t y of the Witwatersrand, and I am extremely grateful t o a l l t h o s e w h o gave t h e i r a d v i c e a n d e n c o u r a g e m e n t i n its p r o d u c t i o n ; M r . J . W . K i t c h i n g , w h o d i d most o f the preparation o n the skull; D r . A . R. I . Cruickshank, w h o read the manuscript; M r . B . Maguire a n d Miss J . Roets f o r discussion, a n d D r . C. K . B r a i n , w h o k i n d l y l e n t m e sabre t o o t h specimens f r o m t h e Transvaal M u s e u m collections. Explanations to text figures Fig. 1 Right lateral view of skull M.8280. Fig. 2 Palatal view of skull M.8280. Fig. 3 Outer view of right mandible M.8280. Fig. 4 Inner view o f l e f t mandible M.8280. Fig. 5 Inner view o f right cheek teeth M.8280. Fig. 6 Outer view of l e f t cheek teeth M.8280. Fig. 7 Dorsal view of lower jaw. Distortion corrected. M.8280. Key to Abbreviations alveolus al. ant. acc. csp. anterior accesory cusp cor. proc. coronoid process con. condyle external auditory meatus earn. frontal fr. glenoid giof. infra-orbital foramen jugal jm. maxilla mandibular foramen man. f o r . mandibular symphysis man. sym. mental crest men. cr. mental foramen men. f o r . mental process men. proc. meta. metacone orbit 0 . parietal Ppal. palatine para. paracone parad. paraconid parast. parastyle proto. protocone protod. protoconid pt. lac. f o r . posterior lacerate foramen posterior palatal foramen pt. pal. f o r . squamosal s. tb. typanic bulla REFERENCES B R A I N , C. K . , ( 1 9 5 8 ) . T h e T r a n s v a a l A p e - M a n b e a r i n g cave d e p o s i t s . Transv. Mus. Mem. N o . 11. C H A N G , H . C , ( 1 9 5 7 ) . O n n e w m a t e r i a l o f some m a c h a e r o d o n t s o f P o n t i a n age f r o m Shansi. Vertebr. Palasiat, 1, N o . 3, 1 9 3 - 1 9 8 . C O L B E R T , E. H . , (1935). Siwalik mammals i n the American Museum of Natural History. Trans. Am. Phil. Soc, 2 6 , 1 - 4 0 1 . M A T T H E W , W. D . , ( 1 9 1 0 ) . The Phylogeny o f the F e l i d a e . Bull. Am. Mus. nat. Hist., 28, 289-316. P I L G R I M , G. E., ( 1 9 3 2 ) . T h e fossil Carnivora o f India.. Palaeont. Indica., 1 8 , 1232. S T I R T O N , R. A . , ( 1 9 5 1 ) . Principles i n correlation and their apphcation to later Cenozoic H o l arctic c o n t i n e n t a l m a m m a l i a n fauna. Rept. 18th Internat. Geol. Congr. London., 11, 7 4 - 8 3 . T E I L H A R D D E C H A R D I N , P., ( 1 9 3 9 ) . O n tvyo s k u l l s o f Machairodus f r o m t h e l o w e r Pleist o c e n e b e d s o f C h o u k o u t e i n . Bull. Geol. Soc. China, 1 9 , 2 3 5 - 2 5 6 .

Lower Canines

Upper Canines

Diastema Length 30,4 1 1

ength Diastema 1

73 03

ant. post. trans.

ant. post. trans.

02 Ol O i b

ant. post, trans.

ant. post. trans. 12,9 6,7

ant. post, trans.

ant. post, trans.

crown height ant. post, trans.

tsoCTic n Oi^-J Ol

crown height ant. post, trans.

o 3 xs
3 O

-o
Ol 00 b

36,9 14,5

12,8 10,3 1 1 I 1

18,4 8,8
00 t - i

1' K 5 O l O l 4i>. ^ b o o b

Therailurus barlowi (Ewer)

s.
O
1-b

1 1

1 1

Ol ^Oi

4^
b

I' IsD K 3 O OO b b

f^
b

1+

Therailurus piveteaui Ewer

40,0 12,0 1

E3

1 1

1 1

1 1

1 1

1 1

1' K D C D K D OO O "-J b b

Machaerodus transvaalensis Broom Megantereon whitei Broom

1
1

1 1 1 1 1 1 1 1
1 1 1 1 1 I 12,4 6,8

1 1 1 1 1 1 1 1 1
1 1 1 1 1 1 1 1 1

19,6 9,4
h-i "bilso

Megantereon gracilis Broom

cn 13

1'
O

os
O

H - l OO O l Ol ^ O 00

45,5 1 1 34,5 1 1

12,8

22,6 9,9 17,0

53,2 8,9 j

13,0 7,0

a
^ C7>

Ol

4^ 4^
O

Ol

OO 0 0

Megantereon problematicus Collings

y k o B >

1
0 0 KD

h-i KD KD O l OO O l

4^
O

30,2 10,6

12,1 8,2

12,0 5,4

K)

00

OO

Megantereon eurynodon Ewer

21,0 12,5

33,0 12,3

15,5

12,5 6,3

26,0 13,0
1 K ) O O b b

CT> 0 0 o o b

00 b

4^
Ol

Megantereon falconeri Pommel

1
b Ol

1 1

i 1

1 I

Megantereon palaeindicus (Bose) Megantereon praecox Pilgrim

sr

1 1

1 1

1 1

1 1

1 1

1 1

>
s 3 o
r
% cu O

13,0 13,5

1 1 1 1 1 1 1 1 1 1 1 1
21,5
O) Ol t - i NO \^ b b

35,0 13,0

25,0 10,0

O O

Ol O

Megantereon inexpectatus Teilhard

16,0 r 7,0

1 1
1

1 1

1 1 1
h-i OO O l OO h - i b b b

25,0 11,0

18,0

Megantereon nihowanensts Teilhard & Piveteau

1 1 1 1 1 1 1 1 1 1
1 1 1 1 1 1 1 22,0

1 1 11
1 1
H - ' ND 00 t o 01 Ol

Isolated canine associated w i t h M. nihowanensts Megantereon megantereon Croizet & Jobert

1 1

KD 0 0 KD O K o b

68

91

Fig. 3

men.
for.

c m s

Fig. 4

m a n . for.

c m s

5 cms