1999 Contr. Tert. Quatern. Geol. 36(1-4) 25-39 9 figs, 1 tab. Leiden, December

Additional hominoid material from

the of Spain

and remarks on

hominoid dispersals into Europe

J. van der Made

Museo Nacional de Ciencias Naturales

Madrid, Spain

and

F. Ribot

Institut Paleontologic Dr M. Crusafont

Sabadell, Spain

from the Miocene of and remarks into — Made, J. van der & F. Ribot. Additional hominoid material Spain on hominoid dispersals Europe.

Contr. Tert. Quatern. Geol., 36(1-4): 25-39, 9 figs, 1 tab. Leiden,December 1999.

hominoid molar from in collections from 1920s housed at the Museu i Laboratori de A upper Hostalets, recently recognised the Geologia del Seminari is the this well be the first tooth to have been collected in the (Barcelona) assigned to genus ; may dryopithecine

hominoid into are discussed here as Vallès-Penedès. With Hostalets as one of the older EuropeanDryopithecus localities, dispersals Europe well. These coincided with dispersals of other , and such events were related to eustatic sea level changes and global climate. The remains of first have 15.5 Ma (beginning of MN 5); its affinities are not clear. Since the European hominoid may entered Europe ago different are to a different and even represent a Griphopithecus are at least 3 Ma younger, they likely represent , might genus.

than 14 Ma MN and later than 12.5 Ma MN it entered Griphopithecus entered Anatolia probably not later ago (late 5), not ago (late 6)

the The hominoids from Klein Hadersdorf and are these occurrences might represent same species. Remains Europe. ages of Çandir close;

much have also 12.5 Ma from Africa. from Neudorf-Sandbergare younger. Dryopithecus may entered Europe ago, coming directly

— hominoid Key words Dryopithecus, Hominoidea, Vallès-Penedès, stratigraphic age, dispersals.

E-28006 F. Institut J. Van der Made, Museo Nacional de Ciencias Naturales, CSIC, Jose Gutierrez Abascal 2, Madrid, Spain; Ribot,

Paleontologic Dr M. Crusafont, C. Escola Industrial 23, E-08201 Sabadell, Spain.

of Contents nari in Barcelona (MLGSB), one us (JvdM) recog-

suid fossils from nised the present specimen amongst

Hostalets. The newDryopithecus tooth is from a collec-

tion made by M. Guérin in the 1920s, which was later Introduction p. 25

transferred to the MLGSB. At that time, Hostalets was and of the new material 27 Description comparison . . p.

still assumed to be a 'locality', with fossils from a single The ages of Turkish and European 28 level only. To date, 'Hostalets' is considered to represent Miocene hominoids p.

an with two stratigraphical levels, a Late Aragonian Dispersal of dryopithecines into Europe p. 34 area, (Middle Miocene, MN 7 + 8; Neogene Units, Summary and conclusions p. 35 and Lower Vallesian de Bruijn et al., 1992), an (Upper Acknowledgements p. 37 considered to be older than the Miocene, MN 9) one, References p. 37 Can Ponsic and Can Llobateres localities (Agusti et al.,

1984). In view of the fact that the present Dryopithecus

Introduction tooth is from an old collection ('Hostalets oc'), its exact

stratigraphical provenance remains unknown.

Crusafont Pairo the When cataloguing and studying fossil suids in the col- Villalta Cornelia & (1941) were lections of the Museu i Laboratori de Geologia del Semi- first to record a dryopithecine from the Valles-Penedes. 26

- 1. MLGSB M² from - of occlusal middle and Fig. 48486, right of Dryopithecus Hostalets, x 4; upper figures stereo pair view, buccal

posterior views, lower - anterior and lingual views.

This fossil of The is of interest in (IPS 1), the type occidentalis present specimen special being

Villalta Cornelia & Crusafont Pairo, 1949, originally the oldest upper molar of Dryopithecus to have been consisted of two teeth united by a piece of mandible, now recorded to date from the area. Material from Can Ponsic lost. The isolated molars are now numbered IPS 1826 (= (= type material of D. crusafonti Begun, 1992a) and Can

M2) and IPS 1827 (= M3). The specimen was found by Llobateres (attributed to D. laietanus) is younger (Begun

Crusafont in the area of Hostalets, near a track leading et al., 1990). In addition, it represents one of the oldest from Can Vila to Can Mata. Villalta Cornelia & Crusa- remains of Dryopithecus. In light of this, the time seems font additional data the of Pairo (1941) considered it to be of Vindobonian (= right to present on (relative) ages

Late beds in the hominoid localities and discuss model of Aragonian) age, and noted that other European a

Can Mata-Ocata These hominoid into area had yielded Hipparion. dispersal Europe. authors were already aware of the fact that Hostalets did not represent a single locality. Six other mammal species Abbreviations — Numerous collections have been stud- were stated to have been collected from the beds that ied; to indicate the repositories of specimens examined yielded the mandible; none of these favours an Arago- the following abbreviations are used: nian or Vallesian age. Subsequent work has resulted in the the consensus that specimen is considered to be of GML Geological Museum, Lisbon

et 1984; Aragonian age (Agusti al., Begun et al., 1990). HGSB Hungarian Geological Survey, Budapest 27

IGGML Institut fiir Geowissenschaften/Geologie der Montan-

universitat Mesio-distal diametre 11.3 IPS Institut Paleontologic Dr M. Crusafont, Sabadell

IPUW Institut fur Palaontologie, Universitat Wien Bucco-lingual width of the first lobe 11.5

Universiteit IVAU Instituut voor Aardwetenschappen, (= maximum width)

Utrecht Bucco-lingual width between the lobes 10.4 MGB Museo Geologico, Barcelona

MGL Museum Guimet, Lyon Bucco-lingual width of the posterior lobe 9.7

MLGSB Museu i Laboratori de Geologia del Seminari, Barce- of Maximum diagonal the crown, D1 12.5 lona

Nacional de MNCN Museo Ciencias Naturales, Madrid the dl 11.5 Minimum diagonal of crown,

MNHN Museum national d'Histoire naturelle, Paris Maximum diagonal of the occlusal surface, 10.2 MPZ Museo Paleontologico de la Universidad de Zaragoza D2 MTA Maden Tetkik ve Arama, Ankara

NMB Naturhistorisches Museum, Basel Minimum diagonal of the occlusal surface, 7.2

NMM Naturhistorisches Museum, Mainz d2

NMW Naturhistorisches Museum, Wien Mesio-distal length of the anterior fovea 4.6 PIMUZ Palaontologisches Institut und Museum, Universitat

Zurich Mesio-distal length of the posterior fovea 3.5

PDTFAU Paleoantropoloii, Dil ve Tarih Cografya Facultesi,

Ankara Universitesi, Ankara

SLJG Steiermarkisches Landesmuseum Joanneum,Graz Table 1. Measurements (in mm) of MLGSB 48486, right M²

SMNS Staatliches Museum fiirNaturkunde, Stuttgart ofDryopithecus from Hostalets.

UCBL Universite Claude Bernard, Lyon

There is no indication of a metaconule in this crest. The

DESCRIPTION direction and the tri- AND COMPARISON OF THE NEW crest is in a diagonal separates large

MATERIAL gonid basin from the smaller talon. The trigonal crest of

the hypoconc is mesio-buccally directed and ends near the

is bunodont is end of the crista On the distal side of the The fact that the present tooth (Fig. 1) lingual obliqua.

the for its confounded with that of there is small probably reason being metacone a accessory cusp, a 'postmetacone'

The is a suid. 'Bunodont molars' occur in suids, ursids and pri- or 'postero-external accessory cusp'. hypocone

united to the the transcrista mates, though this descriptive term includes very differ- 'postmetacone' by posterior

and the crest. There is much variation in ent morphologies. The low cusps and other morphologi- marginal-distal

features indicate that the the of the Carabelli in the molars cal present specimen belongs to development cusp upper

Hostalets does not have a a . The trigon and hypocone are recognisable, of D. laietanus; the tooth from

Carabelli all. indicating that it is an upper molar. Aragonian and Valle- cusp at

sian European include pliopithecids, Dryopith- The four sides of the tooth are smooth. There is no

unlike in which have ecus and Griphopithecus. lingual cingulum, pliopithecids, a

The occlusal surface has a mesio-distally elongated, wide cingulum along the hypocone. There is a deep furrow

the wall between the and rectangular outline. The base of the crown has a nearly on lingual protocone hypocone,

and similar furrow the buccal wall. square outline, being slightly rounded bucco-distally, un- a on

The molars of darwini like the M' which have a reduced or even absent metacone, upper Griphopithecus are

outline. The tooth molar from resulting in a trianglular or trapezoidal relatively wide; an upper Neudorf-Sandberg

assumed be NT from Hostalets is therefore either an M' or an M. Meas- (Dvinska Nova Ves), to an (Steininger,

has a of 11.6 mm and a width of 13.5 mm urements are given in Table 1. 1967), length

It is much and, in particular, much The mesial crista of the paracone is short and adds to (IPUW, cast). larger

the formationof the mesio-marginal crest, which is longer wider than the Dryopithecus upper molars (compare Fig. and with the mesial of the The tooth is than the Ml of Dryopithecus from merges crest protocone (proto- 2). larger

the Valles-Penedes and but enters into the crista). The mesio-lingual crest of the paracone forms part Rudabanya, of the anterior transcrista, which ends abruptly at the base ranges for the M2 (Fig. 2), and is similar in morphology to 2 1 of the protocrista. The mesio-marginal crest, protocrista the M and M of D. laietanus (see Golpe Posse, 1993 for

anterior in small detailed The size of MGSB 48486 exceeds and transcrista converge one point, a cus- a description).

small that of of the M of from the Vallès- pule, reminiscent of a protoconule. A disto-lingual any 1 Dryopithecus 2 crest is directed from the to the buccal crest of Pencdès and Rudabänya, but is within the of the M paracone ranges We believe the tooth is M' of the protocone. These two crests do not meet and no second (Fig. 2). an Dryopithecus.

Ribot also Ribot et all anterior transcrista is formed. The crista obliqua is con- (1993; see al., 1996) assigned

material from the Valles-Penedes D. laietanus. tinuous and connects proto- and metacone. to 28

Fig. 2. Bivariate plots of meso-distal diameter (DAP) and transverse diameter (DT) of the tooth from Hostalets (triangles) compared to the

M¹ and M² ofDryopithecus fromRudabánya and central Europe (dots) and the Vallès-Penedès (crosses); measurements in mm.

THE AGES OF TURKISH AND EUROPEAN MIOCENE problems with individual localities and with the estima-

tion ofthe ofthe used Mammal HOMINOIDS age commonly Neogene

Units (MN units; de Bruijn et al., 1992). For instance,

et without For several reasons it is important to have a good strati- de Bruijn al. (1992), explanation, placed graphic control of the hominoid localities. Hominoid Klein Hadersdorf and Neudorf-Sandberg in MN 6,

Mein in of the of type material is often incomplete, and for that reason whereas (1986), a study ages European material hominoid forward the from several localities which are geographi- localities, put arguments to place cally and stratigraphically close, frequently is united. localities in MN 8. There was a tremendous difference

This is lim- in the estimate of the of the lower boundary of MN obviously a subjective element, but may be age discussion the of MN 5. In ited by precise stratigraphic (and palaeobiogeographic) 5. There has been on content

information. The reconstruction of hominoid history is Spain and France, where the reference localities of the

MN units tend be included another reason. Before arriving in Europe, hominoids to situated, palaeontologists passed through Anatolia; correlations between these two faunas of Aragonian zone D (Daams & Freudenthal,

in MN but in central similar faunas areas are obviously important. 1988) 5, Europe

There still the of in MN4. are many problems in correlations were placed the important European hominoid localities; there are

- of the second Fig. 3. Correlation scheme; left column eustatic sea level cycles ofHaq et al. (1987), and ages (in Ma) of onset each cycle,

indication in this the corrected Miller al. Each starts with a which is age column being age by et (1996). cycle sea level low,

assumed to have allowed for intercontinental dispersals of terrestrial . The thick black horizontal lines indicate these

dispersal events. The next columns give the zonation of the Aragonian and Vallesian (Daams & Freudenthal, 1988) and the MN

units with dates of the lower limits according to Krijgsman et al. (1994, 1996). The last column gives the Faunal Sets of Pickford

other references Van The (1981) and African localities with their radiometric ages (Pickford, 1986; given by der Made, 1996a).

central columns show suoid and bovid lineages and their European and Turkish localities. Double lines in a column indicate that

in that particular lineage, the samples above the double lines are more advanced than those below; generally this is reflected in a

taxonomic change. The lineages are described in detail elsewhere; Bunolistriodon lockharti lineage (Van der Made, 1996a),

Taucanamo lineage (Van der Made, 1997a, 1998),Bunolistriodon latidens lineage (Van der Made, 1996a), Listriodon splendens

lineage (Van der Made, 1996a), lineage (Van der Made, 1989, 1990a, 1999b), Schizochoerus lineage (Van der Made,

1997a, 1998),Tethytragus langai lineage (Fig. 5; Van der Made, 1994), Tethytragus koehlerae lineage (Van der Made, 1994),

Parachleuastochoeruslineage (Van der Made, 1990a, 1999b). 29 30

As a compromise, zone D was included in MN 5 (Krijgs- Engelswies is in the transition of the lower to the

there is difference be- middle unit of the Sweet Water Molasse and the man et al., 1996). Still, a great Upper

middle unit tween the estimates of the ages of the MN units based on locality of Ravensburg is in the (Heizmann, correlations with marine strata in the western Paratethys 1992). Both localities have yielded the suid Bunolistri-

mammal odon loackharti. The from is ad- and Spanish sections with superimposed locali- one Ravensburg an

and Units and vanced form, of the end of MN5, and that from ties long palaeomagnetic sequences. MN 7 typical is MN 8 have been united into MN 7+8 (de Bruijn et al., Engelswies has an evolutionary stage which more

1992). In the present paper, MN 7 and MN 8 are used in primitive than zone E of the Aragonian (Van der Made,

should be an informal manner to indicate early or late MN 7+8, 1996a). The Engelswies locality probably

the later of low in MN which respectively. placed in part zone D, or 5,

based mi- The stratigraphy and dispersal events are synthesised corresponds to Ziegler's (1995) opinion, on in Fig. 3. The MN units, the Aragonian and Vallesian cromammals. The age is between some 16 and 14 Ma;

and the estimated of their lower boundaries closer to the latter date and therefore 1.5 to 3 zones ages probably localities. are given in columns on the left, while Faunal Sets (Pick- Ma older than the accepted Griphopithecus ford, 1981), selected African and Arabian localities and Age estimates for Klein Hadersdorf and Neudorf-

varied Klein Hadersdorf their ages are given in the right column. The central col- Sandberg have widely. was

suoid bovid and in MN 8 because of the of ‘Gazella umns present a selection of and lineages placed presence their localities. The evolution in these lineages is grad- stehlini’ (Mein, 1986) and in the top of MN6, far above

al. The ual, though fluctuations in evolutionary rates are likely to Neudorf-Sandberg, by de Bruijn et (1992). sup- lockharti have occurred. Besides, the samples are not evenly posed presence of the suid Bunolistriodon (last

has spaced in time. In both cases a clear difference in evolu- occurrence in MN 5) at Neudorf-Sandberg played a

role in the estimation of the of the tive level can be seen between subsequent samples. Dou- age locality, though

these fossils have been have ble lines in a column indicate that for that particular line- subsequently supposed to

fossils with been reworked The of age, the localities above the line have yielded (Mein, 1986). presence Protrago-

evolved than those below. Gener- led Mein the in MN 8. a clearly more aspect cerus (1986) to place locality de al. the ally, but not invariably, this coincides with a change in Nevertheless, Bruijn et (1992) placed locality

in MN6, close to the with MN taxonomy. again very low boundary

Andrews et al. (1996, table 12.7) placed Engelswies 5. These localities have yielded mainly large mammals. and Pasalar in MN5-6 and Neudorf-Sandberg, Klein There are few recent revisions of the faunas from these

based Hadersdorf and Qandir in MN6. They assigned the localities, our current knowledge being mainly on

The detailed hominoids from Neudorf-Sandberg and Klein Hadersdorf studies published half a century ago. study to Griphopithecus darwini, those from Pasalar and of various lineages of large mammals would allow us to

these localities in framework. The Qandir to G. alpani and the one from Engelswies to place a stratigraphic

?i This classification is maintained in localities Pasalar and have been in Griphopithecus sp. Qandir usually placed MN5. subsequent papers (e.g. Köhler et al., 1999). Engelswies MN 6, though the former has also been assigned to

above yielded only a fragmentary tooth (Heizmann et al., Generally, (Jandir is placed Pasalar, though usu-

for this. The of 1996), which of course is a difficult basis for an assign- ally no arguments are provided position

Heizmann in MN 6 that it is much ment to species or genus. Although (1992) £andir high suggests younger placed Engelswies in MN5 and, though the current than Neudorf-Sandberg. opinion is that Pasalar is placed in MN6, Andrews et al. The best documented evidence for the relative strati-

(1996) assigned both localities to MN5-6, which gives graphic positions of Pasalar and Qandir comes from the the impression that in so doing they wished to minimise Bunolistriodon latidens lineage. The increase in meso-

of these suids indicates the gap between Engelswies and localities for which the distal diameter of the incisors

is older than der 1996a; presence of Griphopithecus has been accepted. The ma- that Pasalar Qandir (Van Made, terial from Klein Hadersdorf consists of two long bones Fortelius et al., 19%). and be with the material Similar trends in Listriodon incisors and cannot compared directly type splendens from Qandir, nor with that from Neudorf. Nevertheless, canines and increase in hypsodonty and size in the

Andrews et al. (1996) proposed to include Austria- Tethytragus langai lineage (Bovidae) suggest that

Val & copithecus weinfurteri, as defined on material from Klein Pasalar and £andir are older than Arroyo del IV

Hadersdorf, in G. darwini. This obviously is not based VI and Manchones I (Fig. 4; Van der Made, 1994,

Manchones is in the section of the on a comparative morphological study; the underlying 1996a). i type Arago-

in C2 & assumption seems to be that a geographical grouping nian zone (Daams Freudenthal, 1988). Changes corresponds to a taxonomical grouping. The possibility in the Taucanamo sansaniensis-inonuensis lineage (Pa- that one species is present in all four localities or the laeochoeridae, Suoidea) indicate that Pasalar and Inonii I possibility of different species with different ages was are younger than Sansan (Van der Made, 1993, 1997a). not considered. inonii I is older than Pasalar on the basis of two listrio- 31

4. Increase in in and = of the DT of the the Fig. length (DAP mm) hypsodonty (100 Hp/DT; Hp height entoconid, = width) M3 in

The localities in order from old Pasalar while in the Tethytragus langai lineage. are approximate to young: (studied IPS),

Manchones and Arroyo del Val (IVAU), Sari Çay (studied while in the IPS).

dont lineages. Sansan is placed low in MN 6 and in zone

F of the Aragonian. A short palaeomagnetic section in

Sansan be in various (Sen, 1997) may interpreted ways.

The date for the boundary of the G1/G2 zones

(Krijgsman et al., 1994, 1996) and the correlations by

Van der Made (1996a) suggest for Pasalar and Qandir

older and 12.5 ages slightly than close to Ma, respec- tively.

‘Gazella stehlini’, now referred to Tethytragus, is

present in Klein Hadersdorf, but is not typical of MN 8;

in Europe it is found in late MN 6 and in MN7. The re-

mains from that have rise Neudorf-Sandberg may given

to the citation of Protragoceros, are too poor for a defi-

nite assignment and the genus is likely to have appeared

prior to MN 8. The remains from Neudorf-Sandberg,

described as Bunolistriodon lockharti, belong to Listri-

odon splendens; there is no reason to assume that these

fossils are reworked (Van der Made, 1996a).

Trends in incisor width and canine size and morphol-

der 1996a; data from Mottl, ogy (Van Made, using 1957)

ofListriodon suggest that:

— Klein Hadersdorf is of about the same age as

and Arroyo del Val IV, i.e. close to 12.5 Ma;

— St Stephan is older than San Quirze and of the same

age as or older than La Grive oc (oc = old collections);

— Neudorf-Sandberg is about the same age as or

Grive younger than La oc (i.e. between 12.5 and 11 Ma,

but closer to the latter date. Fig. 5. Bivariate plot of length (DAP; in mm) vs width (DT; in Following Thenius (1952), Rabeder (1978) cited Cono- mm) of the P3 ofConohyus and Parachleua-stochoerus. hyus simorrensis steinheimensis from Klein Hadersdorf. Dot = Conohyus simorrensis from Klein Hadersdorf However, it has been shown that Para-chleuastochoerus (IPUW). Cross = Conohyus simorrensis from Pasalar steinheimensis is of (PDTFAU, MTA, PIMUZ). Oblique crosses = C. simor- Conohyusactually no subspecies

rensis from = simorrensis, but different in different Göriach (SLJG, NMW). Asterisks Cono- a species a genus hyus ebroensis from Fonte do Pinheiro (GML). Triangles (Chen Guanfang, 1984; Fortelius et al., 1996). Although

= Parachleuastochoerus steinheimensis from La Grive Conohyus reduced premolar size from C. simorrensis

(MGL, UCBL, NMB). Inverted triangles = P. stein- (crosses in Fig. 5) to C. ebroensis (asterisks), heimensis from Steinheim (SMNS, NMB). Parachleuastochoerus has still smaller premolars (trian- 32

The size of the from Klein Haders- of the gles). large premolar molars from Klein Hadersdorf(Fig. 6) suggest an

dorf (dot) indicates that these suid fossils represent evolutive level between Pasalar (early MN 6) and Le Within Conohyus. Conohyus, molar size increased form Fousseret (MN 7). Data on Listriodon and Conohyus

ebroensis C. simorrensis (MN 5-7) to C. (MN 8-9) (Van corroborate a position late in MN 6 for Klein Haders-

der Made, 1989, 1998; Fortelius et al., 1996). The size dorf.

6. Increase in in in the first and molars in the Fig. length (DAP; mm) second Conohyus simorrensis-ebroensis lineage. The localities are in

order from old Puente de Pasalar approximate to young: Göriach (SLJG, NMW, IGGML, NMB), Vallecas (MNCN), (PDTFAU,

MTA, PIMUZ), Klein Hadersdorf (IPUW), Le Fousseret (MNHN), El Buste (MPZ), Fonte do Pinheiro(GML).

At Neudorf-Sandberg, there are two small suoids, and al., 1984). St Gaudens was placed in MN 8, when MN 7

and MN 8 still treated different units. the figures and descriptions by Thenius (1952) suggest were as Litho-

the is in the Ara- that these do not represent Taucanamo pygmaeum and stratigraphically, locality placed high

Taucanamo of the local and sansaniensis, as stated by that author, but gonian sequence Propotamochoerus

the palaeochoerid Taucanamo grandaeuvus and possi- palaeochoerus is cited from this locality (Ginsburg,

This bly the suid Albanohyus castellensis (see Van der Made, 1971). species appeared just prior to Hipparion

and indicates that the is 1996b, for systematics). If this observation is correct, locality younger than most of

this the MN 7+8 localities. However, we did succeed in would corroborate a late MN 7+8 age. A. pygmaeus not

(MN 7) evolved into A. castellensis (MN 8) and T. finding the P. palaeochoerus fossils on which the cita-

grandaevus replaced the T. sansaniensis lineage late in tion is based.

MN 6 or in MN 7. Agustf et al. (1996) correlated Rudabanya with Can

Neudorf-Sandberg is late in MN7+8, Qandir and Llobateres on the basis of a list of supposedly shared

Klein Hadersdorf are late in MN6 and Pasalar is earlier taxa. However, taxa such as Aceratherium incisivum

in MN 9- MN6. The difference in age between Qandir and (range 9-12), Hipparion primigenium (range MN

Dorcatherium Listriodon Neudorf-Sandberg might be over 1 Ma. Considering a 11), naui (MN6-11/12),

realistic rate of evolution, the possibility exists, that the splendens (MN 6-9, but not present at Rudabanya), Ko-

older localities have one species (G. weinfurteri) and rynochoerus palaeochoerus (MN 8-9/10), Myoglis

Neudorf-Sandberg another (G. darwini). This is of meini (MN5-10), Eomyops catalaunicus and Glirulus

but it is realistic than the aff. lissiensis course a speculation, more one are not particularly useful for such a pre-

by Andrews et al. (19%). cise correlation.

is in The of the of Cricetulodon subdivide MN Dryopithecus present St Stephan and in the old use entry to 9, collections from La Grive; these localities are as old as originally used by Agusti et al. (1984) and the evolution or older than Neudorf-Sandberg. The stratigraphic and of the suid Parachleuastochoerus huenermanni into P. geographic ranges of Dryopithecus and Gryphopithecus crusafonti during the later part of the Crictulodon zone overlapped. gives the following sequence: Hostalets, Can Ponsic I,

The lower levels of Hostalets (including Can Mata) Can Llobateres and La Tarumba (Fig. 7; Van der Made, are assumed to be younger than Sant Quirze (Agustf et 1990a). 33

Fig. 7. Decrease in length (DAP; in mm) of the M3 of the Parachleuastochoerushuenermanni-crusafonti lineage. The localities are in

order from old El Firal Can Can Llobateres and approximate to young: Wissberg (NMM), (MGB), Rudabánya (HGSB), Ponsic I,

La Tarumba (IPS).

The and Cricetulodon is its smaller relative presence of P. huenermanni species replaced by Microstonyx

(Agusti et al., 1996, fig. 5) at Rudabanya suggests that major. In Europe this took place in MN 10 (Van der

in this this locality is older than Can Llobateres 1 and close in Made & Moya Sola, 1989) and Turkey seems to

the et al., age to Can Ponsic (Fig. 7; Van der Made, 1998) and have been the case at same moment (Fortelius

than The transition The oldest is younger Hostalets. of P. huener- 1996). locality possibly

manni to P. crusafonti seems to be abrupt, no clearly Nikiti 1; this locality yielded M. major (Kostopoulos,

intermediate samples being known. The Vallesian is a 1994). Mein (1986), followed subsequently by Andrews

period of important changes and the MN 9-10 transition et al. (1996), placed Melchingen in MN 9 and Sal-

and the earlier of marked in MN Mein based his data part MN 10 are by important mendingen 11. (1986) on

faunal changes, probably caused by important changes the absence at Salmendingen of Anchitherium and the

in climate. This is called the 'mid-Vallesian crisis' and presence of the castorid Dipoides problematicus, which

one of the effects is an important decrease in suoid di- is common in the Turolian. Abusch-Siewert (1983) de-

versity (Van der Made, 1990a, b; Fortelius et al., 1994). scribed Anchitherium from Salmendingen and believed

It is possible that these changes caused a short period of the locality to be MN 9 (last occurrence of that genus).

rapid evolution inParachleuastochoerus. An alternative However, the 'locality' yielded a possible Microstonyx

explanation could be a gap in the record, implying that major tooth (Fortelius et al., 1996), suggestive of the

there is time between Can Ponsic of MN 10-12 elements in the collection from more (last locality presence

with P. huenermanni) and Can Llobateres (first locality Salmendingen. There are several fissure fillings in the

with P. crusafonti) than we think. It is of interest that area and it is not clear where the old collections come

authors who recognise more than one species of from (Abusch-Siewert, 1983). It seems more prudent to

hominoid material Dryopithecus in the area assign fossils from Can Llo- assume that the comes from older

bateres and localities D. laietanus and the associations, i.e. from MN 9. younger to

material from Can Ponsic and older localities to D. This leaves the latest record of Dryopithecus in fontani or D. crusafonti (e.g. Begun, 1992a), while oth- Europe as early MN 10 (La Tarumba, Polinya II);

ers assign it also to D. laietanus (Ribot e/ a/., 1996). Dryopithecus went extinct during the mid-Vallesian

Middle Sinap has the large suid Hippopotamodon crisis and there is no proof that it extended into MN 11,

antiquus, like Eppelsheim (Fortelius et al., 1996). This as suggested by Mein (1986) and Andrews et al. (19%). 34

THE DISPERSAL OF DRYOPITHECINES INTO EUROPE The next event was but one million after the previous

cycle (15.5 Ma ago, lower boundary zone D, Set Illb,

MN 5). This short time interval makes it difficult to rec-

this ognise the possible dispersal events. During cycle or

In Aragonian times, Africa and Arabia were still con- the anterior one the first hominoid seems to have reached nected and the Balkan and Anatolia formed a subconti- There is Europe (the Engelswies hominoid). virtually no nent, that was not always connected to western and cen- record of large mammals of this age in Anatolia and tral & The Indian Sub- Europe (Rogl Steininger, 1983). southeast Europe. The locality of Ad Dabtiyah with continent in with but little was contact Asia, apparently leakyi (= Heliopithecus) should be placed faunal exchange occurred through the Himalayan area. In in Faunal Set Illb, on the basis of the presence of the suid the Middle East these land masses were separated by Bunolistriodon akatikubas, which is typical of this level shallow seas, that became land during periods of low and which is found also in Maboko (Africa; see Van der eustatic sea levels. At such moments faunal exchange Made, 1996a). The possibility of a relationship between occurred between the Indian Subconti- Africa-Arabia, the Engelswies hominid, Platodontopithecus, A. leakeyi nent, Anatolia-Balkanand Eurasia minus Balkan (Europe and Griphopithecus is difficult to assess at this moment, and Asia minus the Indian Five of such Subcontinent). but the possibility that the European and Chinese finds events are recognised in the Early and Middle Miocene represent a single dispersal event by Afropithecus into der Hominoid from Af- (Van Made, 1996a). dispersals Eurasia merits further investigation. rica to Europe are expected to have taken place in one or The about 14 Ma lower event at ago (TB2.5 cycle, two steps through the Turkish-Balkan subcontinent dur- and Set within boundary zone E IV, MN 5) again

periods oflow eustatic sea level. ing brought a wave of migrants to Europe that have their

et al. studied eustatic level fluctua- Haq (1987) sea origin in Africa (the pliopithecids Pliopithecus and Ple- tions, named and dated the cycles, each with a starting siopliopithecus), Anatolia (the rodent Cricetodon), the sea level low. Miller et al. (1996) slightly corrected the Indian Subcontinent (Conohyus) and from one or more ages and related eustatic sea level cycles to cyclical of these three Sanitherium The bovids areas ( ). Protrago-

in climate. of terrestrial mam- changes global Dispersals cerus and Gentrytragus entered Africa. Fort Ternan and mals that are allowed for eustatic sea level lows by Pasalar belong to this cycle, though they are closer to the should be simultaneous in the different continents. Dis- end of it. It is not clear whether Qandir still belongs to persal events on the continents correspond to dates of the this cycle or to the earlier part of the next. Not later than eustatic level lows et al. An sea as given by Haq (1987). in this cycle, Griphopithecus of African origin entered intercontinentalscheme of correlations was developed on Anatolia. It is possible that some species of African ori- the basis of suoid and bovid evolution der Made, (Van gin had lived already for a long time in Anatolia and

1992b, 1993, 1994, 1996a, 1997a, b, 1999a, and was b) southeast Europe. There is virtually no record of large used correlate in different continents to dispersal events mammals older than 14 Ma. Fossils from the Rotem ba- to each other and to the eustatic sea level cycles of Haq sin in Israel include a primate (Tchernov et al., 1987) et al. (1987). This scheme with its corresponding ages and be of this may age. was developed independently of the study of the palaeo- 12.5 Around Ma (TB2.6 cycle, lower boundary zone sections magnetic in the Aragonian area (Krijgsman type G and Set V, close to the end of MN 6). The primitive et al., 1994, 1996), but the results well. correspond very bovoid Hispanomeryx and the bovid Tethytragus with

For instance, new estimates of the of the MN 4/5 age previous records in Anatolia, Albanohyus, with a previ- boundary (then placed at the zone D/E boundary) by ous record in Fort Ternan, Africa, the bovid Turcoceros,

Krijgsman et al. (1994), based on a palaeomagnetic sec- of Asian or southeast European and Anatolian origin tion in the Aragonian area and the estimates based type (Van der Made, 1999a), the suid Parachleuastochoerus, on suoid correlations (Van der Made, 1992b) and the of Asian(?) origin, and the deer Euprox furcatus, of correlation ofmammal dispersals with Haq et al.'s cycles Asian origin entered Europe (Van der Made, 1993, 1994, der (Van Made, 1996a) are only some hundreds of thou- 1996a, b, 1997b, 1999a). Around this time, a number of sands of apart and both differ some 3 Ma with years pre- mammal lineages went extinct in western and central vious estimates. Europe; e.g. the suidHyotherium and the deer Dicroce- In this model of faunal dispersals (Van der Made, rus, and tragulid diversity in Europe was reduced from

1995, 1996a, 1997b, 1999a), the first major faunal ex- four to one species. Crocodiles disappeared from central change between Africa and Eurasia took place 21 Ma and there have been Europe seem to important changes

this does not seem to have involved primates. ago; in the avifauna (Van der Made, 1992a, 1993; Fortelius et

About 16.5 Ma of TB2.3 ago (beginning cycle, zone al., 1994, 1996) as well.

C, Set within MN of African Ilia, 4), Dionysopithecus Dryopithecus was present in Europe prior to 11 Ma origin entered the Indian subcontinent. Dionysopithecus (La Grive oc, St Gaudens, St Stephan) and may well and have entered China this Platodontopithecus may at have entered Europe as part of the faunal exchange moment, or the next & during cycle (Qiu Qiu, 1995). around 12.5 Ma. However, the Aragonian hominoid rec- 35

in The Valles-Penedes is the with increase of at the ord is very poor Europe. an in humidity in large parts Spain

area richest in hominoid localities in Europe, which pos- end of MN 7+8, as indicated by the distributionof Casto-

sibly is partially for ecological reasons, but for geologi- ridae (Daams & Freudenthal, 1988). If the Engelswies

fossil record in the this cal reasons it has a very poor range hominoid does not represent Griphopithecus, genus

the this in faunal MN5-7. Similarly, record for period is poor may have entered Europe as part of the exchange

several other areas in Europe that might have had fa- around 12.5 Ma. Like in the case of Dryopithecus, its

vourable environments for Dryopithecus (e.g. Styrian absence in western Europe may have been due to the lack

basin, southern Germany). Alternatively, Dryopithecus of record in areas with a favourable environment, though

may have arrived in Spain in the later part of MN 7+8 the ecological preferences of both hominoids are likely

this is the its arrival to coincide to different. (Fig. 9). If case, seems have been

Fig. 8. Stratigraphy of European hominoid localities, the oldest Sivapithecus localities in the Indian Subcontinent(data from Kappelmann

et al., 1991), and selected African and Arabian localities (position according to Pickford, 1986; Van der Made, 1996a). 36

Si- The 12.5 Ma dispersal event is also noted in the Indian vapithecus entered the Indian Subcontinent (Kappelman

Subcontinent with the entry of Helicoportax and Gazella et al., 1991). The dispersals of Dryopithecus and Si-

der 12.5 Ma the hominoid have coeval. (Van Made, 1996a). ago, vapethecus seem to been

Fig. 9. Dispersal of Dryopithecinae into Europe; palaeogeographical reconstruction after Rögl & Steininger (1983) and Cahuzac et al.

There differences of in correlation of continental and marine which have their (1992). are opinion stratigraphy, very likely impact

the reconstruction of the H inclusive of Can K =Klein on palaeogeography. Localities: Ç = Çandir, = Hostalets, Mata, Hadersdorf,

L = La Nova P = SG Saint = San SS St Grive, NS = Neudorf-Sandberg(Dvinska Ves), Pasalar, = Gaudens, SQ Quirze, = Stephan.

The arrows indicate the dispersal during MN 6 and a possible dispersal during MN 7+8 (but see discussion in main text).

Agustf et al. (1996) thought that the common ancestor of have coincided largely with that of the hominoids. Many

Dryopithecus and Sivapethecus first dispersed outside species of Dryopithecus with overlapping temporal and

Africa 15.5 Ma ago. They refer to Steininger et al. geographic ranges have been recognised (e.g. four by

(1996), who did not discuss the first appearance of Andrews et al., 1996). Their morphology and size are

used demonstrate Dryopithecus in Europe, but who did use old dates for rather close and have not been to a the MN 5-6 boundary (16.5-15.2 Ma) and the date of separation in habitat and their geographic distribution

12.5 Ma for the MN6-7 As have and the continuous events not in favour of boundary. we seen, dispersal are there is no MN 6 Dryopithecus locality, so the entry must geographical separation. It is not clear how parallel have been later than 12.5 Ma. The first Sivapithecus is lineages should have evolved and maintained themselves, also of that age. Whether or not Dryopithecus and Si- if this is what occurred. vapithecus dispersed independently, the date of the dis- Ankarapithecus from Sinap (MN 9) was supposed to

related but is persals) is 12.5 Ma rather than 15.5 Ma. The 5 Ma pe- be to Sivapithecus (Andrews et al., 1996), riod of morphological stasis assumed by Agusti et al. now assumed not to be related to Dryopithecus and

(1996) for Dryopithecus is therefore much too long, Graecopithecus, but rather to Sivapithecus (Alpagut et

from the fact that there This is of southeast apart are nearly no early al., 1996). suggestive a European

Dryopithecus fossils to confirm this stasis, even for a and Anatolian branch that evolved from Dryopithecus much shorter period. when it passed through this area on its way to western

Subsequent dispersals of mammals in Europe show and central Europe, though alternatively, an Ankara- there moved inde- were few geographical or ecological barriers. In pithecus-Graecopithecus lineage may have

MN 8, Propotamochoerus palaeochoerus dispersed in pendently of Dryopithecus into this area.

mid-Vallesian crisis occurred either the transi- Europe, in MN 9 Hipparion and Hippopotamodon an- The at tiquus, still later in MN 9, hyraxes and murids and in MN tion from MN 9 to MN 10, early in MN 10 or in a short

10Microstonyx major and Schizochoerus. The distribu- period starting at the MN 9-10 boundary and lasting tion of the tapirs seems to have fluctuated and appears to some time into MN 10. The composition of the bovid 37

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