Aureliachoerus (Suidae, Mammalia) from Agreda and other Miocene localities of Spain

A U R E L I A C H O E R US (SUIDAE, MAMMAI,IA) F R O M A G R E D A A N D O T H E R MIOCENE LOCAI,ITIES OF SPAIN JAN v a n d e r M A D E & JUAN M O R A L E S

MADE JAN van der & MORALES JoR~z 1999. Aureliachoerus (Suidae, Mammalia) from Agreda and other Miocene localities of Spain. [Aureliachoerus (Suidae, Mammalia) d'Agreda et autres gisements mioc~nes d'Espagne]. GEOBIOS, 32, 6: 901-914. Villeurbanne, le 31.12.1999. Manuscrit ddpos~ le 15.04.1998; accept~ d~finitivement le 16.11.1998. ABSTRACT - Fossils of the little pig Aureliachoerus GINSBURG,1974 from Agreda and other Spanish localities are described. A comparison of material from the type localities shows that A. aurelianensis and A. minus are different species. The small A. minus decreased further in size. The larger A. aurelianensis increased the length and complexity of the last molar. This is the oldest clear example of a masticatory adaptation that became spectacular and of great stratigraphic importance in later Suidae. A. minus ranges MN 3-5 (Neogene Mammal Units) or zones A-E of the Spanish local biozonation and A. aurelianensis ranges MN 3-4 or zones A-D. KEYWORDS: AURELIACHOERUS, SUIDAE, ARAGONIAN, MIOCENE. RI~SUMt~ - Des restes fossiles du petit suid6 Aureliachoerus provenant d'Agreda et d'autres gisements espagnols sont ddcrits. La comparaison entre le materiel des gisements types prouve qu'Aureliachoerus aurelianensis et Aureliachoerus minus sont des esp6ces distinctes. Le petit A. minus montre une tendance vers la diminution de la taille. La plus grande esp~ce A. aurelianensis montre un accroissement de la longueur et une complexit6 accrue de la derni~re molaire. C'est le plus vieil exemple clair d'une adaptation masticatrice qui est ult6rieurement devenue spectaculaire et d'un grand int~r~t stratigraphique chez les Suidae. L'extension stratigraphique de A. minus cotrespond ~ MN 3-5 (Unit~s Neog~nes des MammiFeres) ou encore aux zones A-E de la biozonation locale espagnole; A. aurelianensis s'6tend h MN 4-5 ou zones A-D. MOTS-CLI~S: AURELIACHOERUS, SUIDAE, ARAGONIEN, MIOCENE. INTRODUCTION

The recovery of new fossils of Aureliachoerus from Agreda m a d e us revise the S p a n i s h m a t e r i a l of this genus. This m a t e r i a l has been m e n t i o n e d before by Van der Made (1990a) and p a r t of it was described and/or m e n t i o n e d by Golpe Posse (1972, 1981), but most of it was not described in detail. Stehlin (1899-1900) i n t r o d u c e d the n a m e Palaeochoerus aurelianensis for a small suoid found in the sands of the Orl4anais. Golpe Posse (1972) described the small species Paleochoerus minus on the basis of m a t e r i a l from E1 C a n y e t and Can Canals. Ginsburg (1974) i n t r o d u c e d the generic n a m e AureIiachoerus for P. aurelianensis. Collier & Guex (1977) described a skull from La Romieu as Aureliachoerus aurelianensis, b u t noted the peculiarity of its small dimensions. Golpe Posse (1981) m a i n t a i n e d both P. aurelianensis and P. minus in the genus Palaeochoerus. Zapfe (1983) was of the opinion t h a t Aureliachoerus is a subgenus of Palaeochoerus. VAN DER MADE (1990a,b; 1994) included P. minus in Aureliachoerus and Van

der M a d e (1990b) assigned m a t e r i a l from Wintershof West and Pontlevoy to P. minus. H e l l m u n d (1992) noted t h a t P. minus has a size similar to, w h a t he believed to be, A. aurelianensis from W i n t e r s h o f West and P e t e r s b u c h 2 and t h e r e f o r e he considered b o t h species to be synonymous. H e l l m u n d did however, consider the genus Aureliachoerus to be valid. No c o m p a r i s o n b e t w e e n A. minus and A. aurelianensis from t h e i r type localities has been published. G i n s b u r g (1974) n a m e d the genus Aureliachoerus for the species "aurelianensis" because he placed t h a t species in the H y o t h e r i i n a e , Suidae, while he placed the genus Palaeochoerus in the Doliochoerinae, T a y a s s u i d a e (= "Old World tayassuids"). Some however, place Palaeochoerus in the Palaeochoerinae, Suidae ( M c K e n n a & Bell 1998). It has been suggested (Van der Made 1994), t h a t the classification of Palaeochoerus as a suid might be based on a classic p a p e r by P e a r s o n (1927), who d e s c r i b e d the b a s i c r a n i u m of some skulls of H yot heri um u n d e r t h e n a m e Palaeochoerus. P e a r s o n (1927) and G i n s b u r g (1974) m e n t i o n a

902 series of characters that separate the "Old World tayassuids" from the Suidae. In a series of papers one of us described additional characters that separate Suidae from the "Old World tayassuids", and arrived to placing this group in a subfamily of their own, the Palaeochoeridae, instead of in the Doliochoerinae, Dicotylidae (= Tayassuidae) (Van der Made 1994, 1996a, 1997). Though Ginsburg (1974) gave a diagnosis for Aureliachoerus that includes some of the important points, no detailed description of Aureliachoerus has been published, comparing it with Palaeochoerus and the other "Old World tayassuids". Ginsburg (1980) supposed that Aureliachoerus aurelianensis replaced Palaeochoerus typus in MN 2b and persisted till MN 5 (MN = Neogene Mammal Units; De Bruijn et al. 1992). Now that two species of Aureliachoerus are being recognized, it becomes necessary to reconsider the ranges of these species. In this paper we intend: 1) To describe the Spanish Aureliachoerus and in particular the new material from Agreda. 2) To discuss the differences between A. aurelianensis and A. minus, based on a comparison of material from the type localities. 3) To describe some of the differences between Aureliachoerus and Palaeochoerus, in particular those that have importance for the classification at the family level. 4) To discuss the stratigraphical range of the species of Aureliachoerus in Spain and Europe. Nomenclature Tooth nomenclature is after Van der Made (1996b). Abbreviations of measurements and indices Measurements are taken as in dicated by Van der Made (1996b). All measurements in mm. DAP: Antero-posterior diameter in cheek teeth or bones. DLL: Linguo-labial diameter in incisors. DLL': Linguo-labial diameter expressed as a percentage of the width (DT or, preferentially, DTp in lowers and DTa in uppers) of the first molar DMD' = (DLL Ix / DT M1) x 100 %. DMD: Meso-distal diameter in incisors. DMD': Meso-distal diameter expressed as a percentage of the width (DT or, preferentially, DTp in lowers and DTa in uppers; not DAP, as by Van der Made 1994 a, fig. 3) of the first molar DMD' = (DMD Ix / DT M1) x 100 %. DT: Transverse diameter ("width") ef a cheek tooth or bone. DTa: DT of the anterior lobe of a cheek tooth. DTp: Transverse diameter ("width") of the second lobe of a cheek tooth. DTpp: Width of third lobe of a M3. In M 3 with small talons, this measurement is difficult or impossible to take in a consistent way. Index: (DAP / DT) x 100% er (DMD / DLL) x 100%. L: Length of a bone in proximo-distal direction. Lext: External length of the astragalus. Lm: Length of the astragalus in the middle. Lint: Internal length of the astragalus. Lu: Length of the upper part of the calcanaeum.

Collections BSPHGM: Bayerische S t a a t s s a m m l u n g ffir Palfiontologie und historische Geologie, Mtinchen. GML: Geolegical Museum of Lisbon. IPS: Instituto de Paleontolog/a, Sabadell. MGL: Museum Guimet, Lyon. MNCN: Museo Nacional de Ciencias Naturales, Madrid. MNHN: Mus6um National d'Histoire Naturelle, Paris. MPV: Museo Paleontol6gico de Valencia. MSNO: Museum des Sciences Naturelles, Orleans. NMB: Naturhistorisches Museum, Basel. NMW: Naturhistorisches Museum, Wien. SMNS: Staatliches Museum ftir Naturkunde, Stuttgart. SLJG: Steierm~irkisches Landes Museum, Graz. UCBL: Universit~ Claude Bernard, Lyon. UUU: Universiteitsmuseum, Universiteit Utrecht.

THE AURELIACHOER US LOCALITIES A N D T H E I R STRATIGRAPHY

AGREDA The locality of Agreda is situated near the village of Navarrete del Rio in the Calatayud-Teruel Basin (Arag6n; number I in Fig. 1). Daams & Freudenthal (1988) gave more information on the geographical and geological situation and gave the following list of rodents: Peridyromys murinus, Pseudodryomys ibericus, Pseudodryomys simplicidens-group, Armantomys / Praearmantomys sp., Microdyromys sp., Glirudinus modestus, Ligerimys fahlbuschi, Heteroxerus rubricati and Castoridae indet. This association places the locality in MN3 or biozone A of the upper Ramblian. The large mammal fauna is inedited and the following list should be considered as preliminary: Pseudailurus sp., Anchitherium sp., Aureliachoerus aurelianensis (described here), Palaeomeryx sp., Andegameryx sp. and Cervidae indet. The collection includes some otherwise indeterminate remains of turtles and birds. RUBIELOS DE MORA Crusafont et al. (1966) discovered fossil mammals in the small basin of Rubielos de Mora in a locality that is now known as Rubielos de Mora 1 (number 3 in Fig. 1) and dated the fauna as lower Vindobonian. De Bruijn & Moltzer (1974) described a collection of rodents from a locality nearby, Rubielos de Mora 2, and assigned the fauna to the "cricetid-vacuum", a period in the Early Miocene (MN 3b) without cricetids. De Bruijn et al. (1992) assigned the locality to the early part of MN 4 on the basis of Neocometes similis; the genus is not known in western Eruope before MN 4. Montoya et al. (1996) presented a new mammal locality (Alto Ballester) and synthesised the information of the basin. They assigned all localities to MN 3 or biozone A of Daams & Freudenthal (1990), giving more importance to the absence of modern cricetids. The Aureliachoerus molar and remains of Procervulus cf. dichotomus and Palaeomeryx sp. come

903

F~GUas 1 - G e o g r a p h i c a l p o s i t i o n s of t h e AureZiachoerus l oc a l i t i e s . 1 = A g r e d a , 2 = Bufiol, 3 = R u b i e l o s de Mora, 4 = E1 C a n y e t a n d C a n C a n a l s , 5 = St. A n d r e u de l a Barca, 6 = C o s t a B l a n c a , 7 = C a n J u l i a , 8 = Moll C a l opa . Situation gdographique des gisements d Aureliachoerus.

from the same bed as the locality Rubielos de Mora 2, but not from exactly the same locality. BUNOL The locality of Bufiol (Valencia) is situated in the Tertiary basin of Rambla de Bosna (number 2 in Fig. 1). Garcfa et al. (1975) gave an overview of the geology of the basin. An updated faunal list and an overview of the palaeontological research on the fossils from Bufiol is given by Van der Made et al. (1998). The locality was first cited in 1922 and a large number of papers deal with its fossil vertebrates and invertebrates. The rodents place the locality in MN 4, or in zone C of the Aragonian (Daams & Freudenthal 1981). The suids of the locality, including Aureliachoerus are described and figured by Van der Made et al. (1998). MOLI CALOPA The locality is situated in the Vall~s-Pened~s basin (Catalonia; number 9 in Fig. 1). Agustf et al. (1984) gave the faunal list, which includes Dorcatherium. They found the microfauna is not very helpfull in dating and placed the locality in MN 3b (at that time including the level of Artenay), but noted that the tragulid might indicate that the locality belonged to MN 4. At that time a number of entries was considered to mark the difference between MN 3 and MN 4, including that of Dorcatherium (MSIN, 1979). Now the level of Artenay is included in MN 4 (Mein 1990), and it is considered that the entry of Dorcatherium is within MN 4, not at the base, or in zone C of the Aragonian (De Bruijn et al. 1992; Van de r Made 1996). See also Fahlbusch

(1985).

EL CANYET AND CAN CANALS The localities of E1 Canyet and Can Canals are situated in the area of Rubf in the Vall~s PenedOs basin (Catalonia; number 4 in Fig. 1). Crusafont et al. (1955) described elements of these faunas. Agustf et al. (1985) described the geological context and placed the localties in MN 4, without giving the faunal list. The localities are in the "Complejo continental inferior", a continental unit that is overlain by the "Complejo marino y de transicidn', a marine unit with foraminifera that according to Anglada & Martfn (1971) belong to Blow zone N8. Golpe-Posse (1974) cited Dorcatherium from E1 Canyet and Deinotherium and Bunolistriodon from Can Canals, which is in accordance with the position in MN 4 or zone C. Golpe-Posse (1972) introduced the name Palaeochoerus minus for small pigs from E1 Canyet and Can Canals and later described them in more detail (Golpe-Posse 1981). CAN JULL~ Can Julia is situated in the Vall~s-Pened~s (Catalonia; number 7 in Fig. 1). Crusafont et al. (1955) described a collection of large mammals from Can Julia. At another place, but near, micromammals were found (Agusti & Cabrera 1980). The geological context and a composite faunal list are given by Agusti et al. (1984). The locality is overlain by marine sediments with a Blow zone N8 foraminiferal association (Anglada & Martfn 1971). The faunal list includes Megacricetodon and Gomphotherium, Golpe-Posse (1974) cited also Bunolistriodon; the locality seems to be MN 4, or zone C of the Aragonian.

904 COSTA BLANCA The localities of Costa Blanca I and Costa Blanca II are situated in the Vall~s PenedSs basin (Catalonia; number 6 in Fig. 1). Golpe-Posse (1974) gave a faunal list for Costa Blanca II, which is not very usefull in dating the locality. Agusti et al. (1984) placed the locality in MN 3 (at that time including the level of Artenay, or zone B of the Aragonian) on the basis of Pseudodryomys ibericus. However, this species is also cited from Bufiol (Daams 1976), which is placed in zone C or MN 4.

Species Locality Collection and number

A. aurelianensis

A. aurelianensis

The locality of San Andreu de la Barca is situated in the Vall~s-Pened~s (Catalonia; number 5 in Fig. 1). Agustf et al. (1984) gave a faunal list: Pseudodryomys aff. ibericus, Peridyromys murinus, Ligerimys aff. antiquus, Melissiodon cf. dominans, Cainotherium miocaenicum and Aureliachoerus aurelianensis and placed the locality in MN 3. Later and more extensive collections of micromammals did not yield modern cricetids and the locality is placed in MN 3, zone A of the Aragonian (Agusti, pers. comm.).

A. aurelianensis

- Palaeochoerus

aurelianensis

STEHLIN, 1 8 9 9 /

D i a g n o s i s -"Petite taille. Canine sup~rieure m~le courte et bomb6e comme chez H. major, p4 muraille externe moins souvent divis~e que chez Hyotherium et toujours plus dissym~trique. Canine inf~rieure mille plus large que chez Hyotherium. Pr~molaires inf6rieures plus ~troites que chez H. major." (Diagnosis for the species and genus Aureliachoerus aurelianensis as given by Ginsburg 1974).

P3 DAP DTa DTp

1

9.5 4.5 5.3

P4 DAP DTa DTp

M1 DAP DTa DTp

i M2 DAP DTa DTp

M3 DAP DTa DTp DTpp

10.4 6.1 6.7

11.0 8.4 8.6

12.4 9.6 9.5

16.1 10.2 8.5 6.8

10.2 5.9 6.6

10.9

12.5 9.5 9.5

16.2 10.2 8.4 6.7

r

>10.5

1

>10.7 > 8.3 10.6

Costa Blanca lI IPS 1504 A. aurelianensis

A. aurelianensis

6.1 1

Costa Blanca II IPS 1575 A. aurelianensis

9.2 10.6

1

Costa Blanca II IPS 1576 A. aurelianensis

10.8 9.6 9.3

9.8

l

Costa Blanca II IPS 1560

> 8.8 7.7

6.9 1

Costa Blanca II IPS 1673 A. aurelianensis

Type species 1900.

P2 DAP DTa DTp

r

Agreda MNCN Ag 7

Aureliachoerus GINSBURG,1974

P1 DAP DTa DTp

Agreda MNCN Ag 1

SAN ANDREU DE LA BARCA

D E S C R I P T I O N OF T H E M A T E R I A L

r/1

6.2

8.7

9.3

2.9

4.3

6.5

r

18.6 10.2 8.3 8.1

Can Canals IPS 1418 A. aurelianensis

1

Moli Calopa IPS A. aurelianensis

5.7 3.1

1

San Andreu de ]a Barca IPS A. aurelianensis

6.5 2.9

9.5 4.2 4.8

10.2 6.4 6.8

10.4 7.6 8.1

12.0 10.5 9.2

16.7 9.1 7.6 6.4

7.2 3.1 3.1

10.4 4.3 4.5

10.2 6.4 7.4

11.0 8.9 9.3

12.6 10.2 10.3

18.5 10.3 8.8 7.8 ->16.4

1

Bufiol MPV BS23 A. minus

7.7 5.2 r

10.2 8.2 >9.3

11.3 .4 9.6

15.1 9.7 8.8 7.3

r

9.6 7.0 7.3

10.6 8.9 8.4

14.9 8.7 7.6 6.0

Costa Blanca I IPS 1954

Aureliachoerus aurelianensis (STEHLIN,1899) Lectotype - Stehlin (1899) did not indicate a holotype, nor a type locality, but mentioned material from Artenay, Fay-auxLoges, Chilleurs, Pontlevoy, Selles-sur-Cher, Brfittelen etc. He f i g u r e d a M 3 f r o m A r t e n a y (PI. 1, fig. 13), w h i c h t h e n w a s s t o r e d i n P a r i s a n d a P3-4 (P1-2 i n h i s t e r m i n o l o g y ) f r o m t h e " S a n d e n d e s O r l ~ a n a i s " , i n O r l e a n s (P1. 3, fig. 35). G i n s b u r g ( 1 9 7 4 ) i n d i c a t e d t h a t t h e t y p e w a s f i g u r e d b y S t e h l i n ( 1 8 9 9 , P1. 3, fig. 35). T h i s c a n b e t a k e n a s t h e i n d i c a t i o n o f a l e c t o t y p e . M a y e t ( 1 9 0 8 , p. 1 5 8 ) s t a t e d t h a t h e c o u l d n o t f i n d a n y m o r e t h e s p e c i m e n f i g u r e d b y S t e h l i n . S t e h l i n ( 1 8 9 9 , p. 1 3 2 ) m e n t i o n e d a specimen from Artenay in Orl6ans with number 207, but did n o t s t a t e t h a t i t is t h e s p e c i m e n h e f i g u r e d . A m a n d i b l e w i t h this number from Artenay and in Orleans has the P3-M2 and t h e M3 i n a l v e o l u s . T h e s p e c i m e n is f i g u r e d c o m p l e t e l y b y M a y e t ( 1 9 0 8 , P 1 . 5 , fig. 9). A s f a r a s c a n b e j u d g e d t h e s p e c i m e n in Orleans may be identical with the specimen figured by Stehlin, and thus be the lectotype.

A. minus

Can Canals IPS 1441 Aureliachoerus sp. 1

10.4

Can Julia IPS --

>6.9

TABLE I - D i m e n s i o n s o f t h e l o w e r c h e e k t e e t h o f A u r e l i a choerus, in mm. Dimensions des dents jugales infdrieures d'Aureliachoerus, en rnm. Type

locality

- Artenay.

Age - Early Miocene, Aragonian,

z o n e B, M N 4.

D i a g n o s i s - Large Aureliachoerus, with P4 with paracone and metacone not well separated and

905 Species Locality Collection number

r/l

A. aurelianensis Agreda MNCN Ag 1

1

P1 DAP DTa DTp

P2 DAP DTa DTp

P3 DAP DTa DTp

7.6 4.1 4.6

P4 DAP DTa DTp

M1 DAP DTa DTp

M2 DAP DTa DTp

8.4 10.2

10.5 10.7 10.6

+-12.4

8.3 10,2

10.5 10.6 10.6

11.9 12.3 11.3

9.1 11.i

10.8 11.0 12.1

M3 DAP DTa DTp

vidual. M N C N - Ag M N C N - Ag M N C N - Ag M N C N - Ag M N C N - Ag M N C N Ag M N C N Ag M N C N Ag M N C N Ag M N C N - Ag

2 Left 1 I. 3 R i g h t 12. 4 Left 13. 5 Left 12/3. 6 Left DI3? 7 F r a g m e n t of a r i g h t M 1. 8 F r a g m e n t of a r i g h t D 4. 9 Right astragalus. 10 R i g h t c a l c a n e u s . 11 F r a g m e n t of a r i g h t M T III.

-

-

r

A. aurelianensis Costa Blanca II P S 1502

6.3 3.3 3.4

10.0 6.0 7.8

1

13.2 11.7 9.4

>6.7 3.8 4.1

A. a u r e l i a n e n s i s Costa Blanca II IPS 1578

14.6 12.9 11.5

A. aurelianensis Costa Blanca II IPS 1670

13.1 12.8 11.0

A. aurelianensis Costa Blanca II IPS 1671

r

A. aurelianensis Costa Blanca II IPS 1673

r

A. aurelianensis Costa Blanca II IFS 1686

r

A. aurelianensis Bufiol MPV BS24

1

IPS IPS IPS IPS IPS IPS IPS IPS IPS IPS IPS IPS IPS IPS

1502 1504 1505 1507 1560 1575 1576 1577 1578 1670 1671 1673 1673 1686

- Left m a x i l l a w i t h p4 a n d M 1. - Left m a n d i b l e w i t h M 1 to M 3. - R e m a i n s of a premolar. - Left p 2 - Left P4. - Left m a n d i b l e w i t h M 2 a n d M 3. - Left m a n d i b l e w i t h P4 a n d M1. - R i g h t 11. - Left M 3. - R i g h t M 3. - R i g h t p3. - R i g h t p3. - Left m a n d i b l e w i t h P2 a n d P3. - P o s t e r i o r lobe of a r i g h t M 1 or M 2.

9.9

Can Canals:

6.8

Moll Calopa:

IPS 1418 - R i g h t M3 a n d f r a g m e n t of M 2. IPS 1413 - Left m a n d i b l e w i t h P1 to M 3.

_>8.9

San Andreu de la Barca: 6.8

IPS - Left m a n d i b l e w i t h P2 to M 3.

BuYtol: M P V BS23 - R i g h t M 3. M P V BS24 - Left M 3.

11.4 14.4 12.2 9.9

A, m i n u s El Canyet IPS 1416

1

8.6 9.6

10.4 10.3 9.8

10.8 11.1 10.0

12.5 9.8 8.7

A, m i n u s El Canyet IPS 1417

1

7.9 9.2

9.1 10.1 10.4

10.1 11.5 10.3

11.4 >_10.3 8.7

A. m i n u s El Canyet IPS 1420

r

<11.5 11.2 10.2

12.0 _>10.6 8.9

3_. m i n u s Can Canals IPS

r

4ureliachoerus sp. Rubielos de Mora MNCN

r

9.7

Aureliachoerus

minus

( G o L P E POSSE, 1 9 7 2 )

H o l o t y p e - IPS 1417, a left m a x i l l a w i t h p4 to M 2, a n d IPS 1422, a left M 3, from E1 C a n y e t . Golpe P o s s e (1972) i n d i c a t e d IPS 1417 as holotype a n d l a t e r (Golpe P o s s e 1981) s t a t e d t h a t also IPS 1422, a left M 3, IPS 1420, a r i g h t M 2, a n d IPS 1421, a r i g h t M 3, belong to t h e s a m e i n d i v i d u a l a n d t h u s to t h e holotype. H e l l m u n d (1992) i g n o r e d Golpe Posse's (1981) paper, a p p a r e n t l y c o n f u s e d IPS 1422 a n d IPS 1421, a n d i n d e p e n d e n t l y s t a t e d t h a t t h e left M 3 b e l o n g s to t h e holotype. T h e c o n t a c t facets of IPS 1422 a n d IPS 1417 fit v e r y well a n d t h e s p e c i m e n s h o u l d i n d e e d be c o n s i d e r e d as p a r t of t h e holotype. T h e s h a p e a n d s t a t e of w e a r s u g g e s t s t h a t t h e r i g h t M 2 a n d M 3 also belong to t h e s a m e individual. Paratypes Golpe P o s s e (1972) i n d i c a t e d IPS 1415 from C a n C a n a l s a n d IPS 1421 f r o m E1 C a n y e t as s y n t y p e s ; p r o b a b l y p a r a t y p e s w a s m e a n t . IPS 1415 w a s f i g u r e d (Golpe Posse 1972, P1.4, fig. 4b; 1981, P1. 1, fig. 3) a n d is identical to a spec i m e n s t u d i e d by us, w h i c h b e a r s t h e n u m b e r IPS 1441. -

12.5 >11.0 >10.3

TABLE D i m e n s i o n s of t h e u p p e r c h e e k t e e t h of Aureliachoerus, in ram. Dimensions des dents jugales supgrieures d ' A u r e l i a c h o e r u s , en ram. 2

-

Costa Blanca 11:

A, a u r e l i a n e n s i s Costa Blanca lI IPS 1505 A. a u r e l i a n e n s i s Costa Blanca II IPS 1507

-

-

T y p e l o c a l i t y - E1 C a n y e t . A g e - E a r l y Miocene, A r a g o n i a n , zone C, M N 4.

with

a tendency

for the

M3 increase

in length

by

adding distal cusps. M e a s u r e m e n t s o f t h e l e c t o t y p e - (DAP x DTa - DTp): P3 7.9 x 3.3 - 3.6; P4 9.9 x 5.2 - 5.7; M 1 10.5 x 7.6 - 8.4; M 211.8 x 9.8 - 9.0. Material

-

Agreda: M N C N Ag 1 R i g h t p1, ps-4, M1-3, P3-4, f r a g m e n t of M1, M2. 3 left p2,p4, M 1, f r a g m e n t of M 2, P4, M1-3; probably all of one indi-

Small Aureliachoerus with p4 with well separated paracone and metacone, no tendency to enlarge the M3 is known. D

i

a

g

n

o

s

i

s

Material

-

-

El Canyet: IPS 1417 + IPS 1422- Left m a x i l l a w i t h p4 to M 3 I P S 1420 + IPS 1421- R i g h t m a x i l l a w i t h M 2 a n d M 3 IPS F r a g m e n t of 11or 12 -

906

Species Locality Collection and number

r/L

A. aurelianensis

1

11 12 DMD DMD

5.6

Agreda MNCN Ag 2,4,5 A. a u r e l i a n e n s i s

r

12/3 DMD

5.2 >3.8

6.2 3.8

DCx? DMD

r

D4 DAP DTa DTp

3.2 2.0

Agreda MNCN Ag 6 A. aurelianensis

D4 D3 DAP DAP DTa DTa DTm DTp DTp

5.1 >6.9

Agreda MNCN Ag 3 A. a u r e l i a n e n s i s

13 DMD

r

>6.9 r

5.1 6.9

Costa Blanca II IPS 1577 A. m i n u s

Can Canals IPS -A. m i n u s

l

9.4 4.1 5.8

Can Canals IPS -A. m i n u s

1

9.5

Can Canals IPS --

TABLE 3 - Dimensions of the incisors and milkteeth of Aureliachoerus, in mm. Can Canals:

IPS 1183 - Left D3 IPS 1415 (IPS 1441) - Right mandible with M1 to M3 IPS 1416 - Left maxilla with p4 to M3 IPS - Left D3 IPS - Fragment of a possible left D4 Costa Blanca I:

IPS 1954 - Right mandible with M1 to M3 Aureliachoerus Material -

Rubielos de Mora:

MNCN - Right M2 Can Julia:

IPS - Left M1/2

sp. indet.

of the

material

from

Spain.

In some cases, the u p p e r m o l a r s h a v e the protopreconule clearly fused to the c i n g u l u m a n d separate from the protocone (Figs 2.1, 3.1, 4.8, 4.9). In other cases this is not so clear; in p a r t i c u l a r w h e n the specimen is worn, t h e r e m a y seem to be a better connection w i t h the protocone (Fig. 4.1). In P a l a e o c h o e r u s t y p u s the protopreconule is always clearly connected to the protocone; m o s t Suidae, including H y o t h e r i u m h a v e the protopreconule fused to the cingulum. The lingual roots of the u p p e r m o l a r s are separate. W h e n this c h a r a c t e r can be o b s e r v e d in P a l a e o c h o e r u s , the roots are divergent, b u t connected by a thin bony s t r u c t u r e , like in a webbed foot. In Suidae the roots are s e p a r a t e and divergent. This c h a r a c t e r h a s b e e n discussed a l r e a d y by Stehlin (1899/1900) a n d Van der Made (1996a). One of the m o l a r s from Costa B l a n c a is relatively wide. Save for one M 1, the m o l a r s of A. m i n u s are far outside the r a n g e s of the A r t e n a y sample, in p a r t i c u l a r in the M~(Fig. 5). The W i n t e r s h o f West molars are on a v e r a g e s m a l l e r t h a n those from Artenay, and t e n d to be l a r g e r t h a n those from E1 C a n y e t a n d C a n Canals.

Agreda MNCN Ag 8

A. attrelianensis

Description

The talon of the M 3 is not extended distally in a n y of the specimens (Figs 2.1, 3.1-3, 4.8-9). This is also obvious in the bivariate plots, w h e r e the S p a n i s h specimens are s h o r t e r (relative to t h e i r width) t h a n those from A r t e n a y a n d W i n t e r s h o f W e s t . The roots of the l o w e r m o l a r s are b r o k e n or are in the m a n d i b l e a n d therefore c a n n o t be observed. I n suids t h e r e are four roots, two below each lobe, w h e r e a s in palaeochoerids the roots are fused r e s u l t i n g a single root below each lobe. The specim e n s from A g r e d a are s h o r t a n d wide c o m p a r e d to those from Artenay. The m o l a r s of A . m i n u s from Spain are outside the r a n g e s of A . a u r e l i a n e n s i s from type locality A r t e n a y (Fig. 6). The specimens from W i n t e r s h o f West t e n d to be small c o m p a r e d to those from A r t e n a y a n d the difference increases from M 1 to M 3. Most M 3 h a v e a simple t h i r d lobe w i t h only one m a j o r cusp (Figs 2.4-5, 3.4). Only a specimen from C a n Canals (Fig. 4.5) h a s two cusps in the t h i r d lobe. Its w i d t h is c o m p a r a b l e to the homologues in

FIGURE2 - Aureliachoerus aurelianensis from Agreda (1-10) and Aureliachoerus sp. from Rubelos de Mora (11). 1, MNCN Ag 1 right P3-M3; a) buccal view, b) occlusal view. 2, MNCN Ag 1 - left p2; a) lingual view, b) occlusal view, c) buccal view. 3, MNCN Ag 1 - left p4-M1, occlusal view. 4, MNCN Ag 1 - right M2.3; a) lingual view, b) occlusal view, c) buccal view. 5, MNCN Ag 1 - left M2_3; a) buccal view, b) occlusal view, c) linguall view 6, MNCN Ag i - right Ps; a) lingual view, b) occlusal view, c) buccal view. 7, MNCN 9 Ag 2 - left I1; a) lingual view, b) buccal view. 8, MNCN Ag 4 - left I3; a) lingual view, b) bucco-distal view. 9, MNCN Ag 1- left P4; a) buccal view, b) occlusal view, c) lingual view. 10, MNCN Ag 3 - right I2; a) labial view, b) lingual view. 11, MNCN Ag 1- right M1/2; occlusal view. 1, M N C N Ag 1- p3-M3 droites; a) rue buccale, b) rue occlusale. 2, M N C N A g 1- p2 gauche; a) rue linguale, b) rue occlusale, c) rue buccale. 3, M N C N Ag 1- p4-M1 gauche, rue occlusale. 4, M N C N Ag 1- M2. 3 droites; a) rue linguale, b) rue occlusale, c) rue buccale. 5, M N C N A g 1- M2_3 gauches; a) rue buccale, b) rue occlusale, c) rue linguale. 6, M N C N Ag 1 - P3 droite; a) rue linguale, b) vue occlusale, c) rue buccale. 7, M N C N Ag 2 - 11, gauche; a) rue linguale, b) rue buccale. 8, M N C N Ag 4 - I3 gauche; a) vue linguale, b) rue bucco-distale. 9, M N C N A g 1- P4 gauche; a) vue buccale, b) vue occlusale, c) rue linguale. 10, M N C N A g 3 - I2droite; a) vue labiale, b) rue linguale. 11, M N C N Ag 1- M 1/2 droite; rue occlusale.

907

2a

, -~

4a

i~~

~v ~a

5b

~~:~ i 8 ?a

1

9a

O

3cm

i: ~ : ~ ~

~7b ~

l lob

908 Species Locality Collection and number

r/1

A. aurelianensis Agreda MNCN Ag 9

r

1

A. aurelianensis 35.4 Aga'eda MNCN Ag 10

Astragalus

Calcanaeum

r

metapodial

A. aurelianensis Agreda

r

TABLE 4 - D i m e n s i o n s

Lext Lm Lint

DTp DTd

R d

>21.6 18.8 20.7

>10.0 > 10.9

11.8 >3.1

head: DAP DT

neck: DAP DT

SF: DAP DT

9.3

9.0

11.8

8.6

5.1

8.9

The l o w e r i n c i s o r s (Figs 2.7, 2.10, 4.4) are both absolutely and relatively larger than those of P. typus.

Lu

DAPp DTp

MT III 8.6

of the bones of Aureliachoenus, in mm.

A. aurelianensis and superior to those inA. minus. M3 with additional cusps or even additional lobes occur in A. aurelianensis from Artenay. This is the explanation for the great variation in size in the M3 from Artenay, seen in Figure 6. The third lobe of the M 3 in A. minus from Can Canals, Costa Blanca I and Wintershof West is simple.

The p4 has the paracone and metacone very well separated in the type of A. minus (Fig. 3.1). In the specimens assigned to A. aurelianensis, the separation is not so clear (Figs 2.1, 4.1), particularly not at the buccal side, and resembles the state in the sample from Artenay. The p1, p2 and p3 are only represented inA. aurelianensis (Figs 2.1-2, 4.2). They are narrower than in P. typus and are not so high as is comon in the Palaeochoeridae. The P4 has the protoconid and metaconid very well separated (Figs 2.9, 4.3, 4.6). The protoprecristid is large and projects, but no separate paraconid is formed and the whole structure does not project as much foreward as in Palaeochoerus. Though described in other terms, this character was gaven great weight by Ginsburg (1974). The P1, P2 and P3 (Fig. 2.6) have cusps t h a t are not so high and pointed as in most Palaeochoeridae and Palaeochoerus in particular (as was noted already by Ginsburg, 1974), but are lower and blunt as in the Suidae.

A possible DC x is a very small tooth, with a high crown. There are but few fossil suoid DC x known, which makes a determination of this tooth problematic. The deciduous canines are changed at a very early stage in life of the suoids, therefore the teeth are very small and but few of them are preserved.

An 12/3has an elongate crown, which probably had a low main cusp. It has a long and low postcrista and lingual cingulum. Its low and elongate shape suggests it is a second incisor, rather than a third. The second and third upper incisors tend to have short, high and pointed crowns in Palaeochoeridae, whereas they are lower, more elongate and blunt in t h e Suidae. The m a n d i b l e is relatively deep as in Suidae; Palaeochoeridae tend to have shallower mandibles. The cheek teeth assigned to A. aurelianensis are larger than the homologues of A. minus and are close in size to the material from Artenay, type locality of A. aurelianensis (Figs 2-3). The upper premolars are narrower than the homologues in P. typus. Both A. minus and A. aurelianensis are smaller than Hyotherium meisneri, the smallest Hyotherium described.

DISCUSSION The Spanish material differs from the Palaeochoeridae, but resembles the Suidae in the following characters: 1) the morphology of the lingual roots of the upper molars, 2) the protopreconule morphology of the upper molars, 3) the shape of the P1-3, 4) the shape of the 12, 5) the depth of the mandible. The material differs in particular from Palaeochoerus in: 6) the paraconid of the P4, 7) the size of the I1.2, 8) the lesser width of the upper molars. In these characters, the m a t e r i a l resembles A. aurelianensis from its type locality Artenay. These characters, several of which were not mentioned by Ginsburg (1974), coroborate his observations t h a t Aureliachoerus is a suid and different from Palaeochoerus, which is no suid. Within the Spanish material there are two groups of different size and morphology. The first group is formed by the fossils from Agreda, Bufiol, St.

FIGURE 3 - A u r e l i a c h o e r u s m i n u s , f r o m E1 C a n y e t (1-2) a n d C a n C a n a l s (3-4). 1, I P S 1 4 1 7 - l e f t l~ 2 a n d I P S 1 4 2 2 - l e f t M3; a) l i n g u a l v i e w , b) o c c l u s a I v i e w , c) b u c c a I v i e w . 2 , I P S 1 4 2 0 - r i g h t M 2 d r o i t a n d I P S 1 4 2 1 - r i g h t M 3 d r o i t ; a) b u c c a ] v i e w , b) o c c l u s a ] v i e w , c) l i n g u a l view. 3, I P S 1 4 1 6 - a l e f t m a x i l l a w i t h p 4 to M 3, o c e l u s a l v i e w . 4, I P S 1 4 1 5 - a r i g h t m a n d i b l e w i t h M 1 to M3; a) o c c l u s a l view, b) l i n g u a l view. 1, I P 5 1 4 1 7 - p 4 - M 2 g a u c h e et I P S 1 4 2 2 - M 3 g a u c h e ; a) vue l i n g u a l e , b) r u e occlusale, c) v u e buccale. 2, I P S 1 4 2 0 - M 2 d r o i t e et I P S 1421- M 3 droite; a) r u e buccale, b) v u e occlusale, c) r u e l i n g u a l e . 3, I P S 1 4 1 6 - m a x i l l a i r e g a u c h e a v e c P4-M3, r u e occlusale. 4, I P S 1 4 1 5 - m a n d i b u l e d r o i t e a v e c M 1 et M3; a) v u e occlusale, b) vue l i n g u a l e .

909

4a

4b

3cm

910 1211-

111 v 10

vwV

p3

, . ",.~

109-

9 9

9

v

. i~o~

X

.,~1 ~"

Q

o

X

Artenay

9-

A. aureI-Spain

8-

A. minus-Spain Wintershof W

8-

Cetina ,

,

6

,

7 8

,

,

9 10

6

i

8

9

~

,

J

10 11 12

1'3

13-

10-

x St. Gerand

p4

16-

14-

15-

13-

14-

v

1211-

vv

=~'

(3

9

.xx

11-

~,

1213

13-

x

12-

<,

10-

8-

91011

%~

12-

9 . 1011

9

9

1514-

1O- Q a~ 9-

~

9

M3

a

~v

*

10-

6-

~x

9-

11-

o .

10

7

14-

17]

M1

i

,

,

,

4

5

6

7

,54

15

9

i

i

i

~

5

6

7

8

r

10-

i

9

% v

21

v

161 V VvV

8

22

M2

20

v "v vV

19

v v v

18

9-

10. . . 121314

VVvV

11

ml~~ *

2 3 4 5 6

11x

w

vv v

12

8-

12+

V

v v

13-

." +

v

7-

ml V

*u

P4

13

V

5-

Horta das Tripas

1716-

V

14 v

P3

12 11-

4 15-

v

P2

1'0 1'1 1'2 1'3 1'4

FIGURE 5 - Bivariate plots of the upper cheek teeth. Abcis is DAP, ordinate is DT. Legend: 1, Palaeochoerus typus from St.G6rand-le-Puy (MGL, casts SMNS); 2, Aureliachoerus aurelianensis from Artenay; 3, Aureliachoerus aurelianensis from Spain; 4, Aureliachoerus m i n u s from Spain; 5, Aureliachoerus m i n u s from W i n t e r s h o f West; 6, Hyotheriurn rneisneri from Cetina de Aragon (Van der Made 1994); 7, H y o t h e r i u m cf. meisneri from Horta das Tripas (GML). D i a g r a m m e s de dispersion des dents j u g a l e s supgrieures.

Andreu de la Barca, Moli Calopa and a large M3 from Can Canals. These specimens resemble A. aurelianensis from Artenay in general size and morphology. Minor differences remain in that some of the Spanish cheek teeth are relatively wide and most of the M 3 are relatively short with simple third lobes. As stated above, the Artenay sample includes M 3 with additional lobes. From the Late Miocene onwards, the elongation of the third molar is a well known phenomenon in Tetraconodontinae (for instance, Nyanzachoerus, Notochoerus) and Suinae (Kolpochoerus, Hylochoerus, Metridiochoerus, Phacochoerus) and has been used in biostratigraphy (eg. Cooke 1978; Harris & White 1979). The palaeochoerid genus Taucanamo shows also a tendency towards elongation of the third molar (Van der Made 1997) and there are specimens from Sansan that have M3 with additional lobes (Ginsburg 1963). It seems

8-

16

7-

15 14

6 4

. . . . . 5

6

,oo] 7

8

9

8

. . . . 9

10 11 12

9 ~, . . . . . 8 9 16 11 12

~

FIGURE 6 - Bivariate plots of t h e lower cheek teeth. Legend as in figure 5. A u r e l i a c h o e r u s m i n u s is from Can Canals and Costa Blanca I. D i a g r a r n m e s de dispersion des d e n t s j u g a l e s infdrieures. Ldgende cornme p o u r la figure 5. Aureliachoerus m i n u s de C a n C a n a l s et de Costa B l a n c a L

likely that Aureliachoerus and Taucanamo are early examples of this adaptation. The relatively wide first and second molars and third molars with short and simple talon(id)s in Agreda and St. Andreu de la Barca (MN 3) simply reflect a more primitive state than in Artenay (MN 4) and Can Canals (MN 4). In suoids, elongation of the M3, by addition of posterior cusps or lobes, is usually accompanied by elongation of the M2, usually without the addition of posterior cusps, and even of the M1. When elongation of the M3 advances, frequently the hypsodonty of, in particular the posterior molars starts to increase. In a later stage this may be accompanied by decrease in enamel thickness and the formation of cementum. (Van der Made 1989, 1999). Larger third molars in bovids have been interpreted as an adaptation to the ingestion of larger quantities of food poorer in nutrients, and hypsodonty to the ingestion of more abrasive food; that

FIGURE 4 -Aureliachoerus aurelianesis from Costablanca II (1-4, 6-9) and from Can Canals (5). 1, IP51502 - left p4-M1; a) lingual view, b) occlusal view, c) buccal view. 2, IPS1671- right p3; a) buccal view, b) occlusal view, c) buccal view. 3, 1PS1576 - left mandible with P4M1; a) buccal view, b) occlusal view, c) lingual view. 4, IPS1577 - right I1; a) mesial view, b) lingual view, c) distal view. 5, IPS 1418 right M 3 and fragment of M2, occlusal view. 6, IPS1560 - left P4; a) buccal view, b) occlusal view, c) lingual view. 7, IPS1673 - left P2-3; a) buccal view, b) occlusal view, c) lingual view. 8, IPS 1578 - left M3; a) anterior view, b) lingual view, c) occlusal view, d) buccal view. 9, IP51670 - right M3; a) anterior view, b) lingual view, c) occlusal view, d) buccal view. 1, IP51502 - p4-M1 gauches; a) rue linguale, b) rue occlusale, c) rue buccale. 2, IPS1671- p3 droite; a) rue buccale, b) rue occlusale, c) rue buccale. 3, I P S 1 5 7 6 - m a n d i b u l e gauche avec P4-M1; a) rue buccale, b) rue occlusale, c) rue linguale. 4, I P S 1 5 7 7 - 11 droite; a) rue mgsiale, b) rue linguale, c) vue distale. 5, I P S 1 4 1 8 - M 3 droite et f r a g m e n t de M2, rue occlusale. 6, IP51560 - P4 gauche; a) rue buccale, b) rue occlusale, c) rue linguale. 7, I P S 1673 - P2-3 gauches; a) rue buccale, b) rue occlusale, c) rue linguale. 8, I P S 1 5 7 8 - M 3 gauche; a) rue antgrieure, b) rue linguale, c) rue occlusale, d) rue buccale. 9, I P S 1 6 7 0 - M 3 droite; a) rue antgrieure, b) rue linguale, c) rue occlusale, d) rue buccale.

911

2a

2b

4a

i!!~i~ii~~ii~ i~! 6a

9

~i!: 84 ::~

EJ

912 is to say to grazing (Van der Made in press). Elongation, but no increase in hypsodonty, of the third molars Aureliachoerus might thus be related to the quantity of food ingested. Aureliachoerus tends to have relatively thick enamel, much thicker t h a n in bovids, and thicker than in most other snoids (Van der Made 1996). Bovidae are folivoFous and include browsers, grazers and mixed feeders. The recent suids Phachochoerus and Hylochoerus have relatively thin enamel and are grazers and browsers, respectively. Hunter and Fortelius (1994) interpreted the fossil suid Listriodon as a folivore on the basis of macro and micro wear of the cheek teeth. This suid has very thin enamel. Many recent suids feed on various kinds of hard food items, including acorns and the like, and root. They tend to have thicker enamel than the predominantly herbivouros suids (data still from Van der Made 1996). Frugivorous primates are reported to have thicker enamel t h a n folivorous primates (Kay 1981). Within the same taxon, enamel thickness is greater in the species feeding on hard objects (Dumont 1995). In bovids, the hypsodonty tends to increase from the first to the third molar. This is not the case in suids with thick enamel, but in these suids enamel thickness increases from the first to the third molar. (Van der Made 1996.) The observations on M3 size and enamel thickness suggest that: 1) Aureliachoerus was no typical grazer or browser, but consumed hard rather than abrasive food items, and 2) it may have consumed large ammounts of food. Alternatively, the thick enamel might be a compensation for the lack of hypsodonty. A study of the microwear might help in the interpretation of the Aureliachoerus dentition. The second group of Spanish Aureliachoerus is forreed by fossils from Can Canals and E1 Canyet. Nearly all specimens are far outside the metrical ranges for Artenay and in addition have P4 with much better separated paracone and metacone and relatively short and wide molars, in particular the M3. The Wintershof West sample seems to be intermediate between Artenay on the one hand and Can Canals and E1 Canyet on the other (Figs 5-6). We did not have the opportunity to study material from Les Beilleaux t h a t Ginsburg et al. (1988) assigned to A. aurelianensis. This material has wide and large M1 and relatively simple M3. The material from Agreda and Les Beilleaux shows that a large form occurred in MN 3 and was roughly contemporaneous to the small form from Wintershof West. The population from Wintershof West (MN 3) is here interpreted as ancestral to those from Can Canals and E1 Canyet (both MN 4) and a very small A. minus from Seegraben (MN 5; Van tier Made 1998).

There seem thus to be two lineages in the genus Aureliachoerus: the large A. aurelianensis with p4 with not well separated paracone and metacone and with M3 with a tendency to become more elongate with more distal cusps and A. minus with a p4 with well separated buccal cusps, with small and short M3 and with a tendency to become smaller. Hellmund (1992) stated that A. minus is identical in size to material from Wintershof West and Petersbuch 2 and therefore should be included in A. aurelianensis. We agree that the material from Wintershof West is very similar to the type material of A. minus, but the comparison of both samples to the one from Artenay, the type locality of A. aurelianensis, revealed important diffeFences. Ginsburg (1980) indicated t h a t A. aurelianensis ranged from MN 2b to MN 5. The first occurrence of Aureliachoerus in France is based on material from Selles-sur-Cher. The description of this material (Ginsburg & Hugueney 1980) suggests that it indeed belongs to a suid, but, on the basis of this material, or at least this description, it is impossible for us to asses whether the material belonged to A. minus, A. aurelianensis or to some small Hyotherium. Aureliachoerus from Pontlevoy (Stehlin 1925; MNHNP, UUU) seems to be small and thus belongs probably to A. minus. Agrada and St. Andreu de la Barca are MN 3 or zone A, the remaining Catalonian localities, Bufiol and Rubielos de Mora are MN 4 or zone C. Thus the range ofA. aurelianensis in Spain is MN 3-4 or zones A-C and A. minus is found only in MN 4 or zone C.

Material from Horta das Tripas (GML) has been assigned to A. aurelianensis (ANTUNES, 1984). One of us suggested that the morphology and size of the remains fit better Hyotherium meisneri (VAN DER MADE, 1994). Antunes (1996) states that such a determination is impossible since that species does not occur in MN 3. Small hyotheriines occur in a number of MN 3 localities, such as Tuchorzice, Budenheim, Haslach, etc., though they are not exactly identical to H. meisneri. The size of the remains from Horta das Tripas indicates a larger animal than A. aurelianensis and coincides with H. meisneri. However, important elements, such as the M3, are lacking and untill the small Hyotherium species are revised, it cannot be affirmed whether H. meisneri is present in Horta das Tripas. Remains fFom Seegraben (SLJG; MN 5) and Oberdorf (SLJG, NMW; MN 4) belong to A. minus (VAN r)ER MADE, 1998). A. minus is found also in Wintershof West (BSPHGM; MN 3) and in Petersbuch 2 (BSPHGM; MN 4). Collier & Guex (1977) described a skull from La Romieu (MN 4)

913 MN

A. aurelianensis

A. minus Seegt-aben Pontlevoy

5 Baigneaux-en-Beauee? Fay-aux-Loges Rubielosde Mora? Bufiol Can Canals Moll Calopa ?Costa Blanca II Artenay

La Romieu El Canyet Can Canals ?Oberdorf4 ?Oberdorfoc

St. Andreude la Barea Agreda Chilleurs-aux-Bois Les Beiileaux? Chitenay?

WintershofWest

Petersbuch 2

FIGURE 7 - The stratigraphic distribution of the Aureliachoerus

species. A question mark in front of a locality name indicates incertainly about its stratigTaphic position. A question mark behind indicates uncertainly about the taxonomic assignation. Distribution stratigraphique des esp&es d'Aureliachoerus. Le

point d'interrogation devunt le hombre du gisement indique l'incertitude sur sa position stratigraphique. Le point d'interrogation derriere indique l'incertitude sur la ddtermination taxonomique.

and assigned it to A. aurelianensis. However, its small size suggests that it belongs to A. minus. Material from Chilleurs-aux-Bois (MSNO; MN 3), Fay-aux-Loges (MSNO, Mayet 1908, P1. 5.12; MN 4), and possibly Chitenay (NMB; MN 3) and Baigneaux-en-Beauce (NMB; MN 4) belongs to A. aurelianensis. Ginsburg et al. (1988) described material from Les Beilleaux (MN 3) as A. aurelianensis.

The stratigraphic position of the Aureliachoerus localities is given in Figure 7. On the basis of the localities mentioned above, the stratigraphical range in Europe is for A. minus MN 3-5 or zones AE and for A. aurelianensis MN 3-4 or zones A-C/D. A c k n o w l e d g e m e n t s - We t h a n k Dr. Beatriz Azanza (Universidad de Zaragoza) and Dr. Remmert Daams for their help in the study of the large mammals of the CalatayudTeruel basin. The fossils from Agreda were obtained by the Ramblian Project during the field campaign of 1982 and were collected by the team lead by Dr. R. Daams. We t h a n k the following persons and institutions for access to material and help of any other kind: J. Agustf, M. Belinchdn, F. Ch~vrier, B. Engesser, L. Ginsburg, W. Grfif, C. Gudrin, K. Heissig, E. Heizmann, L. Hellemans, G. HSck, M. Hugueney, H. Mayr, S. Moy~t Sol,t, R. Niederl, M. Nieto, M. Philippe, R. Sgmehez, P. Sondaar, M. Telles Antunes. The present study was realized within the projects PB-95-0113 and PB-93-0114 of the DGICYT and PB96-1026-C03-02 of the DGES, and the "Unidades Asoeiadas" program of the CSIC.

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