CRANIUM 24, 2 - 2007

The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

Alexandra+A.E. der Geer George+A. Lyras and van

Samenvatting

in het deel het Griekse Vatera is een laat-pliocene (ongeveer twee miljoen jaar geleden) vindplaats zuidelijk van

eiland vertebraten is is De fauna Lesbos, waar een vastelandsfaunavan grote gevonden en systematisch opgegraven. die bestaat uit de typische Europese zoogdieren van tijd: paarden (Equus), giraffes (Mitilanotherium), gazelles (Gazella), antilopen(Gazellospira), runderen( Leptobos), wasbeerhonden(Nyctereutes ), dassen (Meles), sabeltandkatten

Mammuthus Bovendien (Homotherium), neushoorns(Stephanorhinus), mastodonten (Anancus) en mammoeten ( ). zijn

soort er ook resten van een zeldzame soort makaak(Paradolichopithecus) en van een reuzenschildpad (Cheirogaster)

ontdekt.

Summary

Pliocene the southern of Lesvos inwhich mainlandfaunaof Vaterais a Late (~2 Ma) locality in part Island, Greece, a large vertebrates has been found and systematically excavated. The fauna is composed of the typical European

of that time: horses (Equus), giraffes (Mitilanotherium), gazelles (Gazella), antelopes (Gazellospira), oxes

dirk-toothed cats Homotherium rhinoceroses (Leptobos), raccoon dogs ( Nyctereutes ), badgers (Meles), ( ), of of (Stephanorhinus), mastodonts ( Anancus) and mammoths ( Mammuthus). In addition, the remains a rare species also discovered. (Paradolichopithecus ) and a species of a giant tortoise ( Cheirogaster ) were

Introduction now of the of part of the collections Museum Paleontology and Geology of the University of Vatera is a holiday resort at the south part of Athens and are housed in the Vrissa Natural Lesvos Island (Greece), famous for its long History Collection on Lesvos Island. sandy beach. But the area is known for yet

antherreason: in the nearby sedimentary rocks

there are several sites with fossil vertebrates

et al. De Vos et The fossil sites and their (Dermitzakis 1991; al, 2002). age Although they all belong to the typical Late All the vertebrate fossils found in were expo- Pliocene fauna, which is common in many within the unit of the Vatera sures upper European localities, Vatera is a very special one, Formation. This unit is represented by fluvial as it is there, that the remains of two rare and deposits consisting of alternations of extraordinary creatures have been found. The breccia-conglomerates, sandy clays, sandy first is the primateParadolichopithecus, a large conglomerates and silt (Drinia et ah, 2002). In sized knownonly from Seneze macaque, total, 630 fossils havebeen recovered from 7 (France) andValea Grauneanu (Romania). sites (Fig. 1). All fossils were found within the Vatera is now the third locality in the world sandy clay layers, with the exception of the with this . The second is the giant proboscidean remains (sites H and U), which tortoise one of the largest tortoises Cheirogaster, This were found within the conglomerates. of the world and the last of its kind in Europe. seems to be related to the size of the

which The of this is to an illus- remains, large as they are, scope paper present proboscidean trated overview of the Vatera locality, which is require, just like pebbles, a faster stream flow to often missed within the details of the studies be transported.

published so far (Dermitzakis et al., 1991; F-Site Athanassiou, 2002; Eisenmann, 2002; De This is the richest site of the Formation. It has Lapparent de Broin, 2002; De Vos et al., 2002; been named after the of Drinia et al., 2002; Van der Geer & Sondaar, discovery Funny frag- of bones the initial of the 2002; Sondaar et al., 2006). The illustra- ments at stages research. The site excavated tions, used here, have been prepared by Alexis was systematically for 4 continues Vlachos under the guidance of the authors. The seasons (1997-2000), yielding 550 fossils to 15 different species of photographs of specimens have been shot by belonging mammals, one bird and one reptile (the Kyriakos Sykas. The fossils from Vatera are giant

11 The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

Fig 1 Location of the fossil sites and photos showing some sections of the VateraFormation

de de fossielen foto’s enkele secties de Vatera Formatie. Locatie van vindplaatsen van en met van

12 CRANIUM 24, 2 - 2007

discovered Till tortoise). Based on the fauna composition the which fossil plants were (Fig. 1). has been carried in site has been attributed to the MN 17 (about 2 now, no further study out these Ma) at the Late Pliocene (De Vos et al., 2002). two sites.

DS-Site

called after the and It was names (Daniela fauna of million The Vatera 2 years Sandra) of the ones who found the site. It has been excavated in the autumnof 2000. From ago

fossils The terrestrial this site sixty were collected, belonging macaque

bird and at least ten different mamma- to one kind of Two million years ago, a giant macaque lian species. Its is probably Late Pliocene age (Paradolichopithecus arvernensis). lived around (De Vos et al, 2002). Vatera Remains of this primate have been

found the and consist two E-site only in F-site, of

mandibles (one attributed to an adult female, This was the first site tobe discovered in the other to a juvenile male), some postcranials Vatera. It was named after a nearby building, and some isolated teeth (Fig. 2). The size and which was thought to belong to the Electric morphology of the material is similar to that Company. The only remains that have been from Seneze (holotype of the species) and Valea found are two bovid teeth, material of a large Graunceanuluiin Romania.

horse, a raccoon dog mandible and a femur

was as as the fragment of a large felid. Originally, this site Paradolichopithecus big largest baboons of considered to be of Late Pleistocene (Papio ursinus) today. However, was age what makes this is not its (Dermitzakis et al. 1991; and see below, under macaque so special

of due to the size, even if it is too for a but its the section horses). Later, presence large macaque, of life. Van der Geer & of the Late Pliocene was to raccoon dog, a age way According Sondaar Postcranial material from considered as more realistic Vos et al. 2002). (2002), (De r terrestrial Paradolichopithecu s suggests a highly of locomotion. has also been way It suggested U, H, T and V-site that this large monkey occupied a similar

H and had From each one of these sites (U for Unknown, ecological niche as , a

for T for and for similar of locomotion et House, Tortoise V Vatera), way (Sondaar al., 2006).

is known. In the tusk The authors demonstrated that only one species U-site, a same

fragment and two molars of a mammoth Australopithecus was not a perfect biped as

were found. The modern humans (.Homo but a (Mammuthus meridionalis) sapiens),, employed

a somewhat H-site was discovered while digging trench more all-round locomotion, compa- of house. that trench trained for the construction a In rable to a chimpanzee or macaque,

heavily fragmentated tusks and molars of a which can walk bipedally for quite a distance, found. On the mastodont (Anancus arvernesis) were but are also good at climbing trees. the 2 of the similarities in the From the T- and V-sites, only carapax of ground postcranial small small tortoises were excavated. The material, it has been suggested that the locomo-

tortoises from the T- and V-sites are of little tion of Paradolichopithecus was comparable to

stratigraphic value. The same is valid for the that of early hominids like Australopithecus. of Sondaar al. concluded that mammoth, because the stratigraphic range Finally, et (2006)

this mammoth species is from the Late Pliocene Paradolichopithecus lived at the forest edge till the early Middle Pleistocene. The mastodont bordering the savannah.

A. arvernensis lived from the Late Miocene till One of thetwo Paradolichopithecus mandibles the end of the Late Pliocene. found in Vatera belongs to a juvenile male (Van Other fossiliferous sites der Geer & Dermitzakis, unpublished data). It based was concluded that, on eruption pattern, The lower part of the Vatera Formation consists this animal died during his puberty. For male of lacustrine and brackish deposits (Drinia et al., baboon-like monkeys this is a very stressful Within of formation and 2002). this part the episode of their lives. During this life stage, close to there is of Vatera, a quarry (Fig. 1) leave their natal and become they group periph- claystone that contains plant and fish fossils. On eral. Social behaviour like grooming and a higher stratigraphic level, close to the village playing now decreases, while aggressive of there is second in Vrissa, a clay quarry, encounters and fighting with other males

13 The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

from The mandi- Fig 2 Two mandibles and a humerus from the baboon-like Paradolichopithecus arvernesis Vatera. male the ble on the left belongs to a juvenile and mandible on the right to an adult female.

arvernensis Twee onderkaken en een dijbeen van baviaan-achtige Paradolichopithecus van Vatera. De onderkaak

de rechts volwassen links is van een jong mannetje en onderkaak van een vrouwtje.

3 Hans Brinkerink full Vatera tortoise. The Fig standing next to a sized replica of the giant giant tortoise, named

Brinkerink and “Lolita” by France De Lapparent de Broin, was sculpted by Hans in Baarn (The Netherlands)

Lesvos Vrissa Collection in 1999. It is transported to for opening of the Natural History September of now stand- ing in the backyard of the building, which houses the collection.

Hans Brinkerink staande naast een levensgrote replica van de reuzenschildpad van Vatera. De reuzenschildpad,

genaamd ”Lolita” door France De Lapparent de Broin, was door Hans gemaakt in Baarn (Nederland) en toen naar

de de Vrissa Collectie in 1999. Lesvos getransporteerd voor opening van Natuurhistorische september Ze staat nu

in waarin de bevindt. de achtertuin van het gebouw zich collectie

14 CRANIUM 24, 2 - 2007

The material from Vaterahas been studied by De Lapparent de Broin (2002), who

determinated it as cf. Cheirogaster aff. schafferi.

The reason that she was not able to give a more firm determination (she preferred to use the

abbreviations cf. and aff.) is that she was

missing thebones on which the diagnosis of the and the based. The species genus is tortoises

can be distinguished by morphological features

of the plastron and anterior and posterior part of the the carapax. Since Vatera tortoise consists

only of a partial left hind limb, these features could not be used. Therefore, De Lapparent de

Broin (2002) based her determinationon simi-

larities on size and morphology of the Vatera

material with those of an almost complete skel-

eton of a Cheirogaster from Perpignan (Early Pliocene of France) and the limb elements of other giant tortoises from the Mediterranean.

On the ground of these comparisons De

Lapparent de Broin was able toestimate the size of the Vatera tortoise. According to her, the of could be maximal length its carapax 186 cm 4 Life of the Fig appearance spiral-horned antelope long, an estimation, which brings the Gazellospira, based on the fossils from Vatera, Sesklo from Cheirogaster Vatera as the largest indi- and of and Pyrgos (Museum Paleontology Geology vidual of this genus (Fig. 3). of the University of Athens, Greece). The animal’s

colorationand posture has been adapted from the living Tragelaphus of Africa.

Reconstructie de van spiraalhoornige antilope

de fossielen Gazellospira, gebaseerd op van Vatera, Sesklo en Pyrgos (Museum van Paleontologie en

Universiteit Geologie van de van Athene,

De vachtkleur het dier de Griekenland). van en

lichaamshouding zijn afgeleid van dat van de nu levende Tragelaphus uit Afrika.

increase. About half the male die at

this and also for this age, Paradolichopithecus

was true, with nearly half having died before the third molars erupted (Szalay & Delson 1979: 369)

The giant tortoise

Remains of a giant tortoise were found only in the F-site (De Vos et al, 2002). The material

consists of a femur, a tibia, carpal bones, phalanges and osteoderms, which probably all

belong to a partial left hind limb of a single

individual. Fig 5 Reconstruction of the small Gazella of Vatera,

based on the fossil material from the F-site and the In the past, the large terrestrial tortoises were living G. thomsonii of Africa. attributed to the Testudo and Geochelone. genera These genera are phylogeneticaly unrelated to Reconstructie van de kleine Gazella van Vatera, the forms fossiel large terrestrial and therefore, today gebaseerd op materiaal van de F-site en de nu these taxonomies considered levende G. thomsoniiuit Afrika. are inappropriate.

15 The Late Pliocene vertebrate faunaof Vatera (Lesvos Island, Greece)

The antilopes

the The true antilopes are among most common

fossils in Vatera. In total, five species of ante-

of lopes were recognized: one species Gazella and Gazellospira, three species of one, yet of small unidentified, species a very antelope, probably belonging to the neotragini (De Vos et

al., 2002).

The largest of the three antelopes is Gazellospira

is with characteristic (Fig. 4). It an antelope very morphology, which had a wide geographic

distribution (from Europe to China) during the Plio-Pleistocene (Duvernois & Guerin, 1989).

The most typical features of this animal are the rather elongated metapodials and the robust and anti-corkscrew torsion of the horn cores.

There is only one species of Gazellospira in Europe (G. torticornis), however De Vos et al. (2002) attributed the Vatera specimens to G. cf. torticornis, due to their slightly smaller size.

The of three of has presence species gazelles been suggested by Athanassiou (in De Vos et ah,

2002 and Athanassiou, 2002). He attributed the

Vatera material to G. agaea (the largest), G. cf. Fig 6 Reconstruction of the giraffe Mitilanotherium

Volax bouvrainaeG.(the mediumaff.size) and based on the materialfrom Vatera and (Mu- of and of the borbonica (the smallest, Fig. 5). For the determi- seum Paleontology Geology University of nation of the two smaller gazelles, the abbrevia- Athens, Greece).

tions aff. and cf. were used, because their de Mitilanotherium Reconstructie van giraf gebaseerd identification is unsure: There are only a few op materiaal van Vatera en Volax (Museum van and badly preserved mediumsized horn cores Paleontologie en Geologie van de Universiteit van and the smaller horn cores are shorter that the Athene, Griekenland). minimum sizes for G. borbonica. This is not the

only problem concerning the gazelles from

Vatera. Van der Made (in De Vos et al., 2002) (F-site) belong to a giraffid (Fig. 6). The size and of this material is gave a different interpretation of the material. morphology postcranial very

He suggested that there are only two species of similar to material from otherPlio-Pleistocene

Gazella (G. aff. bouvrainae and G. cf. borbonica), sites, such as Valea Graunceanului (Romania),

that the differences is size are due to sexual Volax and Sesklo (Greece), which favors attri-

dimorphism and that the neotragine is in reality bution to Mitilanotherium inexpectatum (De Vos

a female individual of the small gazelle species. etMitilanotheriumal. (2002) attributed it to cf. Giraffid from has Contrary to only one species of Gazella Inexpectatum). material Volax there been namedMacedonitherium martinii (G. borbonica) in Western Europe, is a larger diversity of Pliocene gazelles in South (Sickenberg, 1967), and that from Kuruk Say Eastern Europe (Kostopoulos & Athanassiou (Tajikisthan) Sogdianotherium kuruksaense but metrical and 1997; Athanassiou, 2002). Therefore, the pres- (Sharapov, 1974), morpholog-

ence of three of Gazella in one is ical differences between these two and species locality genera

considered by us as the most probable scenario. Mitilanotherium are small, and they are to be

A more definite de considered most medium conclusion however, can synonyms; likely all drawn after of the sized Plio-Pleistocene only a more extensive study giraffids belong to one material. species only.

The giraffe, the deer and the ox Mitilanotherium inexpectatum is a medium sized giraffid, probably belonging to the subfamily Some postcranial remains, consisting of Palaeotraginae. The skull is elongated with a phalanges, carpals, an astragalus and an articu- flat cranial roof. The ossicones are long, and are lated distal humerus and proximal radius

16 CRANIUM 24, 2 - 2007

Fig 7 Reconstruction ofMetacervoceros rhenanus (the

small deerfrom Tegelen), based on a complete skel- 8 Life of based Fig appearance Leptobos, on a recon- eton from Seneze (Natural History Museumof structed partial skeleton from Valdarno(Museo di Basel, Switzerland). Storia Naturale, Firenze, Italy). The head is based on Adrie and Alfons the rhenanus klein a by Kennis; body is Reconstructie van Metacervoceros (het drawing adapted after the living Indianbison, Bos gaurus. Tegelse hert), gebaseerd op een compleet skelet van

Senèze (Natuurhistorisch Museumvan Bazel, Reconstructie van Leptobos, gebaseerd een Zwitserland). op gereconstrueerd gedeeltelijk skelet van Valdarno

(Natuurhistorisch Museum van Florence, Italië). De

above the inclined is situated exactly orbits, kop gebaseerd op een schildering door Adrie en

towards the front and bent towards the rear. Alfons Kennis; het lichaam is afgeleid van dat van de

nu levende Indiase bison, Bos The limb bones are elongated and moderately gaurus.

slender, with a size in between that of

Palaeotragus and Samotherium. In Europe

giraffids seem to occur only in Greece,

Romania, and Ukraine; they are missing in the

other European Plio-Pleistocene sites, although

refer cf. recently (Arribas et al, 2001) to a Mitilanotherium foundat the Spanish locality Fonelas (Late Pliocene).

A fragmentary male skull with shed antlers and

its maxillaries, a partial skull, a left maxillary

and some postcranial elements (F-site) are

ascribed to the Dama- like deer (Fig. 7) Metacervoceros cf. rhenanus (Dama cf. rhenana in

De Vos et al., 2002). The size of the maxillaries and the metapodials is similar to that of the material from Tegelen (type locality of the

species). Dama-like deer occur in Europe from

the Late Pliocene until and show a today, 9 Life of the rhino Fig appearance Stephanorhinus metrical gradual and transition Mauricio morphological etruscus. Redrawn from a painting by from the earlier the later forms. Cladistic to Antón (Agusti & Antón, 1997). analysis indicates that they actually all belong

to the der Made Reconstructie van de neushoorn Stephanorhinus genus Dama (Van 1999), in

van een door contrast to the study of Azzaroli (1953;1992), in etruscus. Nagetekend schildering Mauricio Antón (Agusti & Antón, 1997). which a separate genusPseudodama is proposed

17 The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

for the early forms. However, at present, the The rhinoceros

generic nameMetacervoceros is in use for the The rhino remains (a vertebra and some limb earlier Dama-like forms. bones) have been found in the DS- site. The of Remains yet another small deer were found material is attributed by De Vos et al. (2002) to

to cf. Croizetoceros(F-site). They may belong Stephanorhinus etruscus (Fig. 9). The Etruscan but this rhinoceros ramosus, is by no means certain at (Stephanorhinus etruscus) was an

present. C. ramosus has more or less the same inhabitant of the forests. This rhinoceros was

size as M. rhenanus (Valli 2004: C. philisi), but small and had relatively long limbs. It bore two

has a more complicated antler. It is tempting to horns: one larger horn on its nose and one much smaller horn assume that this deer preferred the plains (Valli on its forehead.

2004), contrary to M. rhenanus, which is a forest The horses deer.

There of are two kinds horses in Vatera. A large distal A part of a metacarpal from the F-site and robust one (Dermitzakis et al. 1991; very r might be attributed to the larger bovid Leptobos Eisenmann, 2002) and smaller and more slender If then sp (Fig. 8). so, it is slightly larger than one (Eisenmann, 2002). the Leptobos metacarpals from St. Vallier.

Another is similar. The slender horse (Fig. 10) is represented metacarpal (DS-site) very by some teeth and several remains Other elements are a distal radius and a postcranial (Fig. limb bones phalanx (DS-site). Due to these uncertainties, 11). Its are very long and slender and those of De Vos et al. (2002) attributed the material to cf. morphologically comparable to

stenonis Vos et E. stenonis is Leptobos sp. Equus (De al, 2002). the typical horse of the Plio-Pleistocene of

Eurasia. Therefore, its morphology is very well known. However, because of its large temporal and geographical occurrence it developed a Fig 10Life reconstructionof the small horse (Equus

cf. stenonis) from Vatera.

Reconstructie het van kleine paard (Equus cf.

stenonis Vatera. ) van

18 CRANIUM 24, 2 - 2007

Fig 11 A front limb of a horse (Equus cf. stenonis),

which was found in anatomical position in the

F-site. Scale bar is 5 cm.

cf. dat Voorbeen van een paard (Equus stenonis), was gevonden in de F-site inanatomische positie.

Maatstreep is 5 cm.

she concluded that it animal to very large morphological variation, compli- was an adapted this the about its conditions and cating in way, views dry open landscapes.

taxonomy and exact phylogenetic position, The second horse is a more puzzling one. Its which still remain unclear. remains have been found only in the E-site

Eisenmann (2002) estimated that the size of the (Fig.12). There are only three fossils known, an

small Vatera horse is comparable with a astragal, a metatarsus and a tibia. The astragal

Grevy's zebra (more the 400 Kg) and that its was the first fossil to be found in Vatera, about withers should be height 157 cm. Because its twenty years ago. Although its proportions are

metapodials and phalanges are very slender, comparable to E. stenonis (see Dermitzakis et al.,

1991), it is far more robust thanthat of the F-site

(Eisenmann, 2002). The body size of this horse

has been estimated to 605 up Kg (Eisenmann, 2002).

of two different kinds of horses The presence in

Vatera can not be easily explained, as there are localities theworld with only a few Pliocene in

coexistence of two types of horses (Eisenmann,

2002). One possibility is that the horse from the

of later thanthat of the E-site is a age F-site (originally, Dermitzakis et ah, 1991, thought that There the E-site was Late Pleistocene). are many

faults cutting the Vatera Formation and, at least,

one unconformity is present within the forma-

tion. Therefore, it is possible that there is a

considerable time difference betweenthe two

sites. Such a possibility was not supported by

De Vos et al. because of the of (2002), presence

raccoon dog remains in both sites (Fig. 13). 12 metatarsals from the robust Fig Astragals and Raccoon dogs went extinct from Europe at the horse of the E-site and and the slender horse of (A B) end of the Pliocene and that was sufficient the F-site (C and D). evidence for them to suggest that the E- and of the should be F-sites are same age. It noted Enkelbeenderenen middenvoetsbeenderenvan het

here that the raccoon from the E-site is robuste de E-site het dog paard van (A en B) en van much smaller from the one from the F-site (see slankere paard van de F-site (C en D). below, under the section of carnivora), so

19 The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

Fig 13 Mostof the biostratigraphic data come from the F-site. The site was systematically excavated for 4 seasons

fossils. The mandibleof by a Greek-Dutch team. A. John De Vos supervising the cataloging and packing of the B.

the female C. the of fossils around the site Paradolichopithecus, as it was found in situ. Although presence was known actual location site discovered after Paul Sondaar the in for a long time, the of the was only surveyed area

1997. D. Jan Van der Made taking a photo of the site.

De komen de F-site. De site vier meeste biostratigrafische gegevens van was systematisch opgegravenvoor door Grieks-Nederlands de Vos houdt toezicht het seizoenen een team. A. John op catalogiseren en inpakken van

de het in situ C. Hoewel de fossielen. B. De onderkaak van Paradolichopithecus vrouwtje, gevonden. aanwezigheid

van fossielen rond de site al langer bekend was, werd de juiste locatie van de site pas ontdekt toen Paul Sondaar

der Made het gebied in 1997afzocht. D. Jan van neemt een foto van de site.

20 CRANIUM 24, 2 - 2007

et al. another left femur, (Dermitzakis , 1991),

missing the distal condyles (F-site), a damaged

ulna (DS-site; olecranon and distal end

missing), a distal tibia (DS-site), and a complete tibia (DS-site) (De Vos et al, 2002).

The femur proximal fragment might belong to a dirk-toothed cat (cf. Homotherium latidens; Fig.

16); the same is true for the ulna and the distal tibia. The complete tibia is somewhat smaller and could belong to the dirk-toothed cat (cf. Meganterion cultridens). The almost complete left

femur is again smaller; its morphology is

similar to that of Lynx issiodorensis, but is some- Fig 14 Life reconstructionof the raccoon-dog

based cranial and what smaller. Nyctereutes megamastoides on

postcranial remains from Sesklo (Museum of Paleon-

tology and Geology of the University of Athens, Greece).

Reconstructie de wasbeerhond van Nyctereutes

megamastoides gebaseerd op schedel- en skeletresten

Sesklo van (Museum van Paleontologie en Geologie

van de Universiteit van Athene, Griekenland.

indeed there small differ- maybe is some time

between the sites alternative ence two or, as an

hypothesis, the E-site fauna represents a migra-

tion from a different ecological niche or land-

scape. The of Meles Fig 15 badger Vatera, thorali, based on

the of the The carnivores appearance living European badger.

The of canid mustelid remains a a De das van Meles het (Nyctereutes ), Vatera, thorali, gebaseerd op

(Meles) and felids have been found in the sites uiterlijk van de nu levendeEuropese das.

E, F and DS of Vatera.

The small tortoises and birds The raccoon dog ( Nyctereutes ) has been found in the E- andF-sites Their remains (Fig. 14). are In Vatera the remains of small tortoises and

limited to a hemi-mandiblewith preserved birds have been found as well. At the begin- c-m3 and toothrow (F-site) a right c-m2 (E-site). remains ning, only the of one individual, Their morphology is characteristic for of and consisting a partial carapax a plastron Nyctereutes megamastoides, a canid which went of Testudo ibera have been iden- (T-site) a graeca extinct the end of the Pliocene from at Europe tified (De Lapparent de Broin, 2002). Later, in (Tedford & Qiu, 1991). The Nyctereutes from the the second of year 2003, a partial carapax from E-site is much smaller that the one the in Testudo graeca was found a site very close to

F-site and one of the smallest fossil actually Vatera beach (V-site). Morphologically they do

ever found. Nyctereutes not differ from the living tortoise of the region.

The Meles thorali; of Vatera is badger ( Fig. 15) The bird remains have not been determined on

represented in the F-site by a front part of a and genus species level (Aves gen. et sp. indet.,

skull and a mandible Vos et al., The (De 2002). in De Vos et al., 2002). The material is restricted and lower dentition are both complete. upper to a radius fragment and a humerus (F-site) and Measurements of the lower carnassial show that two associated thoracic vertebrae (DS-site). The the the that from Vallier species is same as St. size of the humerus approaches that of an eagle (France). (Aquila).

The only fossils of larger representatives of the cat family (Felidae; Machairodontinae) from

Vatera are a left femur fragment from the E-site

21 The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

16 Life of the dirk-toothed Homotherium. Based Mauricio Antón & Fig appearance cat upona painting by (Turner Antón, 1997).

de sabeltandkatHomotherium. Gebaseerd door Antón & Antón, Reconstructie van op schildering Mauricio (Turner 1997).

The proboscideans fragmentary metapodial (F-site). The dental elements belong without doubt to the In Vatera two kinds of proboscideans were gomphothere Anancus arvernensis (Fig. 17); the found: a mastodontand a mammoth. size of the metapodial is suggestive for the The of mastodonts of remains consist a same species. complete, very worn left molar, a fragment of a right molar, two tusks (all from H-site) and a

17 Anancus skeleton from Fig The mastodont arvernensis, based on a complete Valdarno, Italy (Museo di Storia Naturale, Firenze, Italy).

De mastodont Anancus skelet Italië arvernensis, gebaseerd op een compleet van Valdarno, (Natuurhistorisch Mu- seum van Florence, Italië).

22 CRANIUM 24, 2 - 2007

The southern mammoth (Mammuthus have coexisted in Europe for some time during

meridionalis; Fig. 18) is represented only by a the Late Pliocene, although their fossils have so

tusk and two molar fragments (U-site) and far notbeen found at exactly the same site.

some molar fragments (DS-site). The attribution Anancus became extinct in the Early Pleistocene,

to M. meridionalis is likely, but not decisive at whereas the southern mammoth Mammuthus

et al., 2002 determinate as cf. meridionalis and rise to the present (De Vos it persisted, gave Mammuthus meridionalis). steppe mammoth (M. trogontherii, Middle Pleis-

tocene) and eventually to the woolly mammoth The gomphothere mastodontAnancus and the (M. primigenius, latest Pleistocene) in Eurasia. elephantine Mammuthus meridionalis seem to

The Fig 18 southernmammoth Mammuthus meridionalis, based on a complete skeleton from Valdarno, Italy (Museo di Storia Naturale, Firenze, Italy). The southern mammoth is the ancestor of the woolly mammothand is the largest proboscidean of all times.

De Mammuthus Italië zuidelijke mammoet meridionalis, gebaseerd op een compleet skelet van Valdarno, de (Natuurhistorisch Museum van Florence, Italië). De zuidelijke mammoetis de voorouder van wolharige mammoet en is de grootste slurfdrager die ooit geleefd heeft.

23 The Late Pliocene vertebrate fauna of Vatera (Lesvos Island, Greece)

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