F R A G M E N T A M I N E R A L O G I C A et P A L A E O N T O L O G I C A 12. 1985 p. 59-66

Alces brevirostris Kretzoi from the Ördöglyuk Cave at Solymár (Hungary)

B y

I . VÖRÖS (Received November 30, 1984)

A b s t r a c t : The ske le ta l r emains of male Alces b r e v i r o s t r i s f r o m the Solymár-Ördöglyuk Cave are descr ibed i n th i s paper . Compar i son shows that A . b r e v i r o s t r i s i s the e a r l i e s t f o r m of the Alces l i n e .

T h e Ördöglyuk Cave i s s i tuated west of the v i l l age of Solymár on the NE slope o f the Z s i r o s -hegy at an a l t i tude of a p p r o x i m a t e l y 300 m above see l e v e l . In t h i s cave of compl ica ted s t ruc tu re L . VÉRTES and O. KADIÖ excavated and co l lec ted m a t e r i a l d u r i n g 1939 and 1942/43, w h i l e i n 1946 L . VÉRTES continued h is researches there by h i m s e l f .

In 1943 an almost comple te skeleton of an e l k came to l i g h t i n the so-ca l led "Kiskörút" sec­t ion of the cave, f r o m d a r k r e d c l a y . It was sent by L . VÉRTES together w i th other faunal remains to M . K R E T Z O I fo r i den t i f i c a t i on and analyses (VÉRTES 1950).

The m a m m a l i a n fauna of the Solymár-Ördöglyuk Cave y ie lded severa l new species and sub­species ( K R E T Z O I 1946, JÁNOSSY 1969, 1979). The e lk r emains were descr ibed i n manusc r ip t by M . K R E T Z O I as Alces b r e v i r o s t r i s new species .

The signif icance of the A lces b r e v i r o s t r i s ske le ta l r emains found i n the Solymár-Ördöglyuk Cave i s emphasized by the fact that the ve r tebrae , the e x t r e m i t y bones - as w e l l as the cast antfer , the s k u l l and the mandible (having been destroyed i n the meant ime) - belonged to a s ingle male i n ­d iv idua l and they are eas i ly measu red . T h e i r d imensions and morpho log i ca l c h a r a c t e r i s t i c s can be compared w i t h other Ple is tocene Alces r e m a i n s .

The Alces b r e v i r o s t r i s skeleton remains a re deposited i n the Paleontologica l Depar tment of the Hungar ian Natura l H i s t o r y Museum, under i n v . N o . : V . 8 2 . 1 4 .

I would l i k e to express m y s incere thanks here to P r o f . M . K R E T Z O I and P r o f . D.JÁNOSSY for t h e i r p e r m i s s i o n to study and to publ i sh t h i s m a t e r i a l .

Skeletal remains o f Alces b r e v i r o s t r i s

- r i g h t s ide cast ant ler ( F i g . 1, th is d rawing has been prepared on the basis of JÁNOSSY 1969, Ta f . I I , f i g . 3 ) . Des t royed . The photo and the d r awing shows v e r y w e l l the most c h a r a c t e r i s t i c features o f the ant ler , name ly the ex t r eme ly shor t beam, w i t h end i n a l a rge wide p a l m a t i o n . T h e r e are 7 t ines at the m a r g i n o f the pa lma t ion , the p o s t e r i o r m a r g i n o f the pa lmat ion is damaged, the t ines a re broken off . The I s ' t ine f r o m the a n t e r i o r m a r g i n of the pa lma t ion is r e l a t i v e l y s m a l l and separa ted . The pa lma t ion i s d iv ided in to two pa r t s along the med ian - sag i t t a l axis of the an t le r

by a wide and deep U-shaped fosse; there a re th ree t ines ( 2 n ^ , 3 r < ^, 4 t n ) i n the wide a n t e r i o r pa r t o f the pa lma t ion , w h i l e i n the~targe surface o f the pa lma t ion behind the fosse the 6 t n , 7*^ and x t n t ines are s i tua ted .

F i g . 1 . Alces b r e v i r o s t r i s , r i g h t cast an t le r (Drawn by I . Pisch)

- s k u l l f ragments (collapsed and p a r t l y des t royed) . Only f ragments of the f r o n t a l , ba s iocc ip i t a l , and palate bones are s t i l l p r e s e r v e d . A c c o r d i n g to the e a r l i e r d e s c r i p t i o n "the p r a e m a x i U a - , and f a -c ionasa l par t s are s t r i k i n g l y ve ry shor t , only 2/3 - 1/2 pa r t s o f recent A lces" ( K R E T Z O I 1946, JÁNOSSY 1969) . - 5 1. i n f . - a t las , ep is t ropheus , v e r t , c e r v i c a l e s I I I . - V u . Phys io log ica l length of v e r t , c e r v i c . ( I - V I I ) : 450 m m . - v e r t , t h o r a c a l i s I . - X I I I . - v e r t , l u m b a l i s I . - V . (Phys io logica l length: 330 m m ) .

GO

- 12 p r o x i m a l pa r t s of costae , - s t e r n u m ( P l . I I , F igs 1-4), I s t - I 5 * " s tenebra of m e s o s t e r n u m .

l S t 2 n d 3 r d 4 t h

Dimens ions ( in m m ) greates t length 78 70 73 68 smal l e s t m e d i a l breadth - 78 - 90

The r i g h t side of the 3 r c * mesostenebra i s b roken off, i t i s m i s s i n g ; there i s a bone p r o l i f e r a t i o n w i t h spondi los is and exostosis on the d e x t r a l l o w e r pa r t of the 4 t n and on the d e x t r a l upper pa r t of the ö̂ h mesostenebra; i . e . c i r c u l a r bone spondylar throses have s l i d in to each other f o r m i n g i n th i s way a "pseudo- jo in t" . The x i p h i s t e r n u m is in t ac t , i t s d imens ions : length 133, breadth of upper p a r t 76, m a x i m a l breadth o f l o w e r apex 41 m m , smal les t b read th of media l pa r t 28, i t s d iamete r 13 .5 m m . - scapula s i n . et dex t . d i s t . f r s . (both i n f r agmenta ry c o n d i t i o n ) . - humerus s i n . et dex t . ( P l . I , F igs 1 ,2 ) . T h e tube rcu lum med ia l e of the humerus d e x t . i s damaged. The diaphysis i s s t r i k i n g l y long, s lender , the epiphyses are r e l a t i v e l y s m a l l . The t roch l ea i s s t rong , t h i ck , i t s b read th 82 ( s in . ) - 81 (dex t . ) m m . The fossa supra t roch lea r i s i s deep. The c r i s t a i s sharp on the t r o c h l e a , on the upper m a r g i n of the t r o c h l e a on the median side there i s an i n c i s u r a . Breadth of caput h u m e r i 78 ( s in . ) - 80 (dext . ) m m . The epicondylus m e d i a l i s i s n a r r o w . - radius s i n . et dext . ( P l . I , F i g s 3 , 4 ) . The rad ius s i n . i s b roken in to several pieces whi le the d i s t a l epiphysis i s b roken off f r o m the dex t . one . The tubera l i g a m e n t i had not developed i n the l a t e r a l side of the p r o x i m a l ep iphys i s . - o lecranon s i n . et d e x t . T h e length of o lecranon i s 98 ( s in . ) - 100 (dex t . ) , i t s breadth 65 ( s in . ) -6 6 . 5 (dex t . ) m m . Bread th of the surface of the a r t i c l e : 47 ( s i n . ) - 44 .5 (dext . ) m m . - metacarpus dex t . ( P l . I , F i g . 5) - pe lv i s s i n . f r . ( P l . I I , F i g . 6 ) . Only pa r t of the corpus i l i i and the an te r io r p a r t of the aceta­bulum are s t i l l ava i l ab le . The f i ssure i n f ront o f the acetabulum i s deep. Height o f corpus i l i i 57, i t s th ickness is 30 m m . - f emur s i n . et dext . ( P l . I , F i g . 6 ) . The p r o x . and d i s t . epiphyses of the femur dex t . have b r o ­ken of f . The diaphysis i s l a t e r o - m e d i a l l y f la t tened, long and s l i m . D iame te r of caput f e m o r i 48, i t s breadth 60 m m . On the d i s t a l epiphysis the l a t e r a l condylus and epicondylus are s t rong ly developed. The n a r r o w (thin) w id th of t rochleae f emor i s i s 46 m m . - t i b i a s i n . et dex t . ( P l . I , F igs 7 , 8 ) . The med ia l m a r g i n o f the t i b i a dext . p r o x . epiphysis i s damaged. - m a l l e o l a r e s i n . et dex t . ( length 32 .5 ; breadth 40 m m ) . The m a l l e o l a r e dext . i s f r a g m e n t a r y . - as t ragalus s i n . et d e x t . ( P l . I I , F igs 7 , 8 ) . The astragalus s i n . i s f r agmenta ry . - calcaneus s i n . et dex t . ( P l . I I , F igs 7,8) - cen t ro ta r sa le ( T c + T 4 ) s i n . (breadth 69, length 57.5 mm) - cunéiforme i n t e r m e d i u m et l a te ra le (Tg-ß) s i n . - meta ta rsus s i n . et dex t . ( P l . I , F igs 9 ,10) - p h . I . dex t . pos t . l a t . - m e d . ( P l . n, F i g . 9) - p h . I . s i n . pos t . m e d . ( P l . I I , F i g . 10) - p h . I I . d e x t . pos t . l a t . - m e d . ( P l . n, F i g . 11) - p h . I I I . dex t . pos t , l a t - m e d . The inner m a r g i n of both p h . I I I . are damaged. - 2 os sesamoideum sup. - r u d i m e n t a r y metapodium 2 /5 (length 183, d i a m e t e r 15 m m ) . F o r the d imensions of the pos t c r an i a l skele ta l e lements see T a b l e 1 and 2 .

T a k i n g in to cons ide ra t ion the dimensions o f the pos t c ran ia l ske le ta l elements of the few Pleis tocene Alces avai lable ( A Z Z A R O L I 1952, 1953, K A H L K E 1959, 1965, 1969, 1971, 1975) (Table 3) we can say that A l c e s b r e v i r o s t r i s i s a s m a l l - s i z e d A l c e s , w i t h the s lender s t y l o - , and zeugopodium and s t rong metapod ia . It has r e l a t i v e l y long humerus and f e m u r / t i b i a and shor t me ta -pod ia .

At present Alces b r e v i r o s t r i s is the sma l l e s t known fos s i l f o r m of the Alces l i n e . T h e a n t l e r -beam i s v e r y shor t and t h i c k ; the neck o f the an ima l i s shor t , w h i l e the dorsa l 1 and l u m b a l regions are r e l a t i v e l y l o n g . As r e g a r d s i t s e x t r e m i t i e s , the humerus and F e m u r / t i b i a are long compared to the metapodia, which suggests a t a l l body height (Table 1 and 3 ) . The diapysis of the metapodia of Alces b r e v i r o s t r i s are s t rong , wide and f l a t . L i b r a l c e s ga l l i cus i s charac te r ized by sho r t e r h u m e r u s / t i b i a and longer metapodia . The humerus and f e m u r / t i b i a of Alces b r e v i r o s t r i s are longer,

whi le i t s metacarpus and meta ta rsus are as long as, o r longer than the respect ive bones o f L i b r a l - ces g a l l i c u s . The diaphysis o f the metapodia o f Alces b r e v i r o s t r i s are s t rong, wide and f l a t . The re i s only a s m a l l difference between the breadth of the l a t e r a l and m e d i a l t rochleae on the d i s t a l epiphyses of the metacarpus and the me ta t a r sus .

Chronology of Alces b r e v i r o s t r i s

The s t r a t ig raphy of the sediment sy s t em of the Solymár-Ördöglyuk Cave is not ye t comple te ly known. L . VÉRTES publ ished only a shor t , p r e l i m i n a r y r e p o r t on his excavations i n the Ördöglyuk-Cave; the s t ra t ig raph ie a l d e s c r i p t i o n of the cave i s even less usable (VÉRTES 1950) . The d a r k - r e d c l ay which contained the r e m a i n s of Alces b r e v i r o s t r i s runs along the cave f r o m i t s uppermos t point (Dice ro rh inus k i rchbergens i s ) to i t s l o w e r m o s t p o i n t . T h i s l a y e r i s of Middle P le is tocene age.

F o r l a c k of the c h r o n o - and b i o s t r a t i g r a p h i c a l evaluat ion of the whole faunal assemblage the age of the d a r k - r e d c lay can be de te rmined only as a wide s p e c t r u m : thus the ch rono log i ca l pos i t ion of Alces b r e v i r o s t r i s is the f o l l o w i n g - f i r s t ha l f of the Ste inheim Faunal Wave, Minde l - R i s s (E l s te r -Saale) I n t e r g l a c i a l — R i s s (Saale) e a r l i e r phase ( K R E T Z O I & PÉCSI 1979).

Scheme of the evo lu t iona ry h i s to ry of the Pleis tocene E lks

L i b r a l c e s - l i n e A . l a t i f r o n s - l i n e A . a l ces - l ine

E a r l y Upper Pleis tocene ( R - W In te rg lac ia l )

Late M i d d l e Pleistocene (Riss)

M i d d l e Pleistocene ( M - R I n t e r g l a c i a l

E a r l y M i d d l e Pleistocene (Minde l )

Late L o w e r Pleistocene ( G - M In te rg lac ia l )

M i d d l e / E a r l y Lower P le is tocene

' L . c a r n u t o r u m '

' L . c a r n u t o r u m '

L . ga l l i cus L . m i n o r

A . l a t i f r o n s pos t remus A . alces

A . l a t i f r o n s

A . l a t i f r ons A . r e y n o l d s i

A . l a t i f r o n s A . r e y n o l d s i

A . alces

A . b r e v i r o s t r i s

R E F E R E N C E S

A Z Z A R O L I , A . (1952): L ' A l e e d i Sénéze. - Palaeontographia I t a l i c a , 47 (1950 - 51): 1 3 3 - 1 4 1 . A Z Z A R O L I , A . (1953): The deer of the W e y b o u r n c r a g and F o r e s t Bed of N o r f o l k . - B u l l . B r i t i s h

M u s . ( N a t . H i s t . ) G e o l . Ser . , 2 (1): 1-96. JÁNOSSY, D . (1969): S t ra t ig raphische Auswer tung der europäisch mittelpleistozänen W i r b e l t i e r ­

fauna. - B e r . deu tsch . Ges . g e o l . W i s s . , A . G e o l . Paläont., 14 (5): 573-643. JÁNOSSY, D . (1979): A magyarországi pleisztocén tagolása ger inces faunák alapján. ( S t r a t i g r a p h i ­

sche Gl iederung des ungar ischen Pleistozäns aufgrund Ver tebra ten) - Budapest, 207 p p . K A H L K E , H . - D . (1959): Die Cerv idenres te aus den altpleistozänen I lmk iesen von Süssenborn bei

W e i m a r . I I I . Die Pos tk ran ia l en Ske l e t t r e s t e . - A k a d . V e r l . B e r l i n , 44 p p . K A H L K E , H . - D . (1965): Die Cerv iden-Res te aus den Tonen von Voigts tedt i n Thüringen. - Paläont.

A b h . A b t . A . , 2 (2 /3 ) : 379-426. K A H L K E , H . - D . (1969): Die Cerv iden-Res te aus den Kiesen von Süssenborn bei W e i m a r . - Paläont.

A b h . A b t . A . , 3 (3 /4 ) : 547-610. K A H L K E , H . - D . (1971): C e r v i d a e . - I n . : N I K I F O R O V A , K . - V . a l . : Pleis tocene o f T i r a s p o l . -

K i sh inev , 137-156. ( in Russian) K A H L K E , H . - D . (1975): Die C e r v i d e n - R e s t e aus den T r a v e r t i n e n von W e i m a r - E h r i n g s d o r f . - A b h .

z e n t r . g e o l . I n s t . Paläont. A b h . , 23: 201-249 . K R E T Z O I , M . (1946): Die W i r b e l t i e r f a u n a aus d e m älteren P le i s tozan von Solymár. (Manuscr ip t ) K R E T Z O I , M . & PÉCSI, M . (1979): P l iocene and Pleistocene development and Chrono logy of the

Pannonian B a s i n . - A c t a Geo l . H u n g . , 22 (1 -4) : 3 -33 .

VÉRTES, L . (1950): A solymári-barlang rétegviszonyairól. (On the s t r a t i g raph ie al condi t ions of the Solymár-Cave) . - Földt. K ö z i . , 80: 199-203.

A u t h o r ' s address: D r . István VÖRÖS Archaeo log ica l Depar tment Hungar i an National Museum H-1088 Budapest Múzeum körút 14-16 . HUNGARY

Table 1

Alces b r e v i r o s t r i s , bone measurements ( in m m )

1 . 2 . 3. 4 . 5. 6. 7.

Humerus s i n . 398 102 41 83 124 45 78 dex t . 400 - 41 .5 83 .5 123 46 79

Radius s i n . - 79 - 72 46 - 53 dext . - 79 44 - 47 31 -

Metacarpus dex t . 315 57 39 67 39 28 37 F e m u r s i n . 452 122 40 104 62 48 126

dex t . - 120 40 - 61 47 -T i b i a s i n . 505 111 45 71.5 102 35 57

dex t . - - 45 .5 71.5 - 34.5 56.5 Meta ta rsus s i n . 373 55 38 68 53 .5 33 41 .5

dex t . 369 55 37 67 .5 53 .5 33 42 P h . I . dex t . pos t . l a t . 81 34 26 31 40 22 26

m e d . 80.5 34 27 32 40 23 26 s i n . pos t . m e d . 81 34 26 .5 31 .5 39 23 26

P h . I I . d e x t . pos t . l a t . 56 33 23 27 43 31 37 pos t . m e d . 57 33 23 26 41 32 38

1 . greatest length, 2 . p r o x . breadth, 3 . smal les t d i a m e t e r o f diaphys i s , 4 . d i s t . breadth, 5. p d i ame te r , 6 . sma l l e s t d i a m e t e r of d i aphys i s , 7. d i s t . d i a m e t e r .

As t raga lus s i n . dex t .

Calcaneus s i n . dex t .

Cen t ro ta r sa le s i n . P h . I I I . dex t . pos t . l a t .

m e d .

Length

77 158 157

57.5 79 83 .5

Bread th

54 54 54 54 69 28 .5 29

Height

46 .5 47 60 61

Scapula s i n . dex t .

sma l l e s t breadth of c o l l u m scapulae 59 breadth of angulus a r c t i c u l a r i s 82 .5 breadth of facies a r t i c u l a r i s 61 d i a m e t e r of facies a r t i c u l a r i s 55 54

E X P L A N A T I O N OF P L A T E S

P L A T E I

A lces b r e v i r o s t r i s

F i g . 1 . humerus dex t . F i g . 2 , humerus s i n . F i g . 3 . r ad ius dex t . F i g . 4 . r ad ius d i s t . s i n . F i g . 5. metacarpus d e x t . F i g . 8. f emur s i n . F i g . 7. t i b i a dex t . F i g . 8. t i b i a s i n . F i g . 9 . meta ta rsus dex t . F i g . 1 0 . meta ta rsus s i n .

P L A T E I I

Alces b r e v i r o s t r i s

F igs 1-4. s tenebra 1 s t - 5 ^ of mesos te rnum F i g . 5. x i p h i s t e r n u m F i g . 6 . p e l v i s s i n . f r . F i g . 7. calcaneus, as t ragalus s i n . F i g . 8. calcaneus, as t ragalus dext . F i g . 9 . p h . I . dex t . p o s t . l a t . - m e d . F i g . 1 0 . p h . I . s i n . pos t . m e d . F i g . 1 1 . p h . I I . d e x t . pos t . I a t . - m e d .

P L A T E I L

Table 2

A lces b r e v i r o s t r i s . ver tebrae measurements (In m m )

Atlas

greatest length 118 greatest breadth 158 greatest height 76 length of body 50 breadth o f c r a n , a r t i c . surface 105 breadth of caud . a r t i c . surface 93

Epis t ropheus

greatest length 135 greatest height 130 length of body 110 length of a rch 67 breadth of caput c r a n i a l e 92 breadth of dens 40 breadth o f fossa caudal is 46 height of fossa caudal is 45

V e r t , c e r v i c a l I I I . - V I I .

1 . 2 . 3 . 4 . 5. 6 .

H I . 87 44 45 49 54 90 I V . 84 42 45 49 55 84 V . 72 - 44 48 54 92 V I . 78 36 43 49

V I I . 6 7 + 37 42 61 - 83

V e r t , thoraca l I . - X I I I .

I . 58 57 44 59 44 112 I I . 51 50 45 57 43 104 H I . 53 41 42 64 40 99 I V . 51 41 39 65 3 8 + 91 V . 54 45 38 66 38 89 V I . 56 44 39 - - 90 V I I . 55 42 37 52 36 V I I I . 58 44 38 56 36 I X . 56 42 38 - 34 8 4 +

X . 57+ - - 35 X I . 57 44 38 60 36 90 X I I . 5 9 + - 38 - - -X I I I . 57 44 38 58 39

1 . greatest length of body, 2 . breadth of caput ve r tebrae , 3 . height of cap . v e r t . , 4 . breadth of fossa ver tebrae , 5. height of fossa v e r t . , 6 . breadth across - p r o c . a r t i c . caud . by v e r t , c e r v i c a l , and - p r o c . t r ansve r sus by v e r t , t h o r a c .

V e r t , l umbar I . - V . 1 . 2 . 3 . 4 . 5. 6 .

I . 64 42 40 49 40 49 I I . 84 47 41 - 37 48 I I I . 6 5 45 41 50 38 50 I V . 67 46 40 51 39 49 V . 67 - 39 6 0 + 37 49

1 . - 5 . the same as those of the v e r t , c e r v i c / t h o r a c a l , 6 . l ength of a rch

T a b l e 3

Compara t ive measurements o f bones of foss i l e lk fin m m )

Humerus

A . b r e v i r o s t r i s

L . ga l l i cus ( A Z Z A R O L I 1952) A . r eyno ld s i ( A Z Z A R O L I 1953)

Radius

A . b r e v i r o s t r i s

L . ga l l i cus ( A Z Z A R O L I 1952) A . l a t i f rons Süssenborn ( K A H L K E 1959,1969)

Metacarpus

A . b r e v i r o s t r i s A . l a t i f r ons Süssenborn ( K A H L K E 1959,1969)

Voigts tedt ( K A H L K E 1965)

T i b i a

A . b r e v i r o s t r i s

L . ga l l i cus ( A Z Z A R O L I 1952) A . l a t i f r ons Süssenborn ( K A H L K E 1959,1969)

Meta tarsus

A . b r e v i r o s t r i s

L . ga l l i cus ( A Z Z A R O L I 1952) A . l a t i f rons Mosbach ( K A H L K E 1959) Süssenborn ( K A H L K E 1959,1969) T i r a s p o l ( K A H L K E 1971) A . l a t i f rons pos t remus E h r i n g s d o r f ( K A H L K E 1975)

1 .

398 400 355

445-450

360-388 492

31 5

50 5

440-470

370 373

350-370

457 .5 4 6 9 +

466

102

79

79

109

57

78 .1 80.2

m

55

55

63.8

74.2

3 .

41 41 .5

44

63 .2

39

45 45 .5

3 7

38

41 42 .5 45 .7

83 83 .5

72

9 5 +

07

83.5

87.8

71 .5 71 .5

92.6 92 .4

6 7 . 5 68

86.5 92 .7

79.4

5.

124 123

47

46

64

39

52.5 56.3

102

53.5 53.5

69 .5 72.0

45 46

31

28

3 9 . 1

35 34 .5

58 .5

33 33

39 .8

78

79

53

73 J

37

52 60 55.7

57 56.5

73 .1 74.2

42 41 .5

55.8 61.2

51.4

1 . -7 . the same as i n Table 1