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GEOLOGICA CARPATHICA,  48, 5, BRATISLAVA,  OCTOBER 1997

341–343

UPPER JAW FRAGMENT OF THE ANANCUS ARVERNENSIS

(CROIZET & JOBERT 1828)

(MASTODONTIDAE, PROBOSCIDEA, MAMMALIA)

FROM THE VILLAFRANCHIAN OF HAJNÁČKA, SLOVAKIA

PETER LUPTÁK

Zoological Institute, Faculty of Sciences, Comenius University, Mlynská dolina, 842 15 Bratislava, Slovak Republic

(Manuscript received October 6, 1996; accepted in revised form March 18, 1997)

Abstract:  

Remains of a representative of the family Mastodontidae (Proboscidea, Mammalia) of the Villafranchian

fauna were found in Hajnáčka, southern part of Central Slovakia. The upper jaw fragment of an Anancus arvernensis
with preserved molars M

sin., M

2

 dex., M

3

 sin., M

3

 dex. is described in this paper. Anancus arvernensis is a relatively

common mastodontid in the Europaean Plio-Pleistocene deposits. Only isolated teeth and bones of mastodontids are
usually found on the territory of Slovakia and more complete fossil remains such as the presented specimen are very rare.

Key words: 

Villafranchian, Proboscidea, Mastodontidae, Anancus arvernensis.

Introduction

The deposits of the vertebrate locality Hajnáčka are of Euro-
paean importance. Since the year 1863, when was the locality
discovered, fossiliferous horizons have provided very rich
vertebrate fauna of the Villafranchian Age (Fejfar 1964). In
the history of the research many geologists and  paleontolo-
gists have been dealed with this locality (Kubinyi 1863; Sza-
bó 1865; Paul 1866; Krenner 1867; Schafarzik 1899; Kormos
1915 In: Fejfar et al. 1990) and from recent years Fejfar
(1964); Fejfar et al. (1985, 1990), etc.

The fossil remains of the locality originated from limnic

basin-fillings with pyroclastic intercalations (tuffs, tuffite,
lapilli, agglomerates) (Fejfar et al. 1990). The complete mam-
mal fauna consists of ca. 30 species. Tooth and bone frag-
ments of mastodonts and tapirs are most frequent. The com-
plete list of mammalian species is available in Fejfar (1964)
and Fejfar et al. (1990).

Systematic Paleontology

Class

Mammalia Linnaeus 1758

Order

Proboscidea Illiger 1811

Suborder

Mastodontoidea Osborn 1921

Family

Mastodontidae Girard 1852

Subfamily

Anancinae Hay 1922

Genus

Anancus 

 Aymard 1855

Anancus arvernensis

 (Croizet & Jobert 1828)

Synonymy: Mastodon arvernesis,

 Mastodon dissimilis, Di-

bunodon arvernensis
Type (lectotype): 

M

1

 sin., described and depicted by Croizet

& Jobert 1828, p. 139–140, Pl. 13, Fig. 1.
Paratypes: 

Dozens of isolated molars with fragments of up-

per and lower mandibles described and depicted by Croizet

& Jobert 1828, p. 133–138, Pl. 1, Fig. 1, 4, Pl. 2, Fig. 7 and
by Osborn 1936, p. 632–633, Fig. 596 (denoted as cotypes).
Type locality: 

Perrier, Auvergne, France

Type level: 

Lower Villafranchian, Upper Pliocene

Localities: 

Many sites in Europe and Asia

Material

A fragment of the upper jaw (ZIBA-2407/1986, Collec-

tions of Zoological Institute, Comenius University, Bratislava)
with the molars M

2

 dex., M

2

 sin., M

3

 dex., M

3

 sin. (see Fig. 2).

Only one molar (M

2

 dex.) is completely preserved, the other

teeth are incomplete. The fossil has a light-brown color with
features after burning in the lava (blackish inner part), while
the teeth are grey.

Locality

The locality of Hajnáčka is located 28 km east of the town

of Lučenec, in the Cerová vrchovina Upland (see Fig. 1).

Fig. 1. 

Geographical location of the Lower Villafranchian locality

Hajnáčka.

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342                                                                                                LUPTÁK

More detailed descriptions of the fossiliferous horizons, chro-
nostratigraphy and other geological aspects are available in
Fejfar (1964), Fejfar & Heinrich (1985) and Fejfar et al.
(1990). The described fragment of the upper jaw with pre-
served molars was discovered in the locality — Hajnáčka I
(area called Močiar (Swamp)), 600 m from the village
Hajnáčka southwest of the poplar alley. The fossil find origi-
nated from the fine sandy tuffites with abundant pyroclastic
agglomerates near Gortva stream gorge.

Description

The fragment of the upper jaw consists of four well pre-

served molars (M

2

 dex., M

2

 sin., M

3

 dex., M

3

 sin.) and the

particularly well preserved maxilla (see Fig. 2). Originally,
the jaw was desintegrated and composed of five major parts.
The size of the maxilla and teeth indicate a juvenile speci-
men of Anancus arvernensis. The number of the molar ridges
is lower than in the adult specimens. Total length of the frag-
ment is 400 mm and width is 320 mm. The palate bone tends
to narrow towards the front. The middle part of this bone has

a remarkable medial suture. Inner structure of the bone is
very cavernous and partly destroyed by burning.

M

dex. is the only molar entirely preserved. The trigon

and talon are same width. The root part is not observable. On
the chewing surface of the molar are 4 regular ridges and one
posterior semiridge. The first two ridges are particularly
abraded. The dentin is largely exposed only on the first ridge.
On the inner palatinal side the first and second ridge have
half  of the height on the external buccal side. The course of
the median sulcus can be traced on the second and third ridg-
es. On the first ridge it is impossible because of intense abra-
sion. The second ridge has one longitudinal groove. The
third and fourth ridges are typical with two longitudinal
grooves. Every halfridge of the third and fourth ridges has
two cusps. On the second ridge it is possible to see the con-
tinual fusion of both occlusial surfaces. The first ridge pre-
sents complete junction of the half-ridges into one surface.
The last fifth ridge consists of four major cusps and one
semicusp in the center of the ridge. The half-ridges of entol-
oph are shifted more distally than half-ridges of ectoloph. The
cement coat is preserved on the bottoms of the synclines. The
cingular thickening is preserved around the whole crown

Fig. 2.

 Upper jaw fragment of the Anancus arvernensis (Croizet & Jobert 1828) ZIBA-2407/1986 with preserved molars and part of max-

illa (photo P. Lupták).

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UPPER JAW FRAGMENT OF THE ANANCUS ARVERNENSIS                                                       343

base. Enamel on walls and crown base show tiny irregular
knolls, pits and grooves. The enamel has a rough pattern at
the end of the synclines. The thickness of the tooth enamel
was measured on the broken parts (M

2

 sin., M

3

 sin., M

2

 dex.)

and ranges from 5.2 to 6.1 mm. The adult specimens may
have 9 mm thickness (Fejfar 1964). Measurement results see
in the table (Table 1). The first ridge of the M

2

 sin. has broken

enamel on the mesial margin. The root is well preserved. The
length of the M

2

 sin. root (fourth ridge) is 89 mm.

M

dex. is nearly complete, only the fourth ridge has lost

half of the buccal cusp. The crown surface has a different
pattern to the third molars. Generally, the second molars are
much longer in relation to their  width, and in this juvenile
specimen without any abrasion. The structure of the bun-
odont ridges is typical with strongly developed half-ridges.
The half-ridges get an alternant position and the synclines
get closed. The fourth and fifth ridge of the M

3

 dex. are in

the bone, mainly the fifth. The crown base is elongated in
outline. Ectoloph and entoloph of the first ridge are com-
posed from several (4–5) secondary (adaxial) conelets. The
conelets (3) are also present in the second, third, fourth and
probably fifth ridge of the crown. Half of the fourth and the
whole fifth 

 

ridges are absent from M

sin.

Proboscidea has always presented special problems to pale-
ontologists. This difficulty is due to the unique manner in
which these teeth are worn during mastication, and the re-
sultant variation in morphology these molars display
throughout their height and length (Froehlich & Kalb 1995).
The alternation of half-ridges is explained by Lehman (1950)
by the tendency of molars to elongate their crown without en-
largement of jaws where the process took place. The speci-
men from Hajnáčka is placed in wear stage 1 (WS  I, after
Froehlich & Kalb terminology (1995)) when on the anterior
end of the second molars extending just to the first ridge.

The upper jaw fragment with preserved four molars is the

next contribution to the Villafranchian fauna of Hajnáčka.
There are only two other localities in Slovakia where this spe-
cies is presented. From Lower Pleistocene of  southern Slova-
kia — Strekov (Holec 1985),  and few undiscribed findings
are known from nearly located place — Nová Vieska.

The fossil was discovered approximately thirty years after

the last intensive research on the locality. The complexity of
the jaw fragment and its state of preservation indicates the
possibility of important finds from this well known paleonto-
logical locality in future.

Acknowledgements: 

For helpful comments, literature I am

grateful to Dr. P. Holec from Department of Geology and Pale-
ontology (Comenius University), Prof. R.Musil (Masaryk Uni-
versity, Brno) and Dr. O. Fejfar (Charles University, Prague).

References

Fejfar O., 1964: The Lower Villafranchian Vertebrates from

Hajnáčka near Fi akovo in Southern Slovakia. Rozpr. Ústř.
Úst. Geol.,

  1–115.

Fejfar O. & Heinrich W.-D., 1985: Zur Bedeutung der Wirbeltier-

fundstätten von Ivanovce und Hajnáčka für die Säugetier-
paläontologie im Pliozän und frühen Pleistozän in Europa:
Kenntnisstand und Probleme. Věstn. Ústř. Úst. Geol.,  60, 4,
213–224.

Fejfar O., Heinrich W.-D. & Heintz E., 1990: Neues aus dem Vil-

lafranchium von Hajnáčka bei Filakovo (Slowakei, ČSSR).
Quart

ärpaläontologie (Berlin), 8, 47–70.

Froehlich D. J. & Kalb J., 1995: Internal reconstruction of ele-

phantid molars: Applications for functional anatomy and sys-
tematics. Paleobiology, 21, 3, 379–392.

Holec P., 1985: Finds of Mastodon (Proboscidea, Mammalia) Rel-

ics in Neogene and Quarternary Sediments of Slovakia
(ČSSR). Západ. Karpaty, Sér. Paleont., 10, 13–53.

Osborn H.F., 1936: Proboscidea. A Monograph of the Discovery,

Evolution, Migration and Extinction of the Mastodonts and El-
ephants of the World. Vol. I. Moeritherioidea, Deinotherioidea,
Mastodontoidea. Amer. Mus. Nat. Hist. New York, 1–802.

Kalb J. & Mebrate A., 1993: Fossil elephantoids from the homi-

nid-bearing Awash Group, Middle Awash Valley, Afar De-
pression, Ethiopia. Amer. Phil. Soc. Transactions, 83, 1–114.

Lehmann U., 1950: Über Mastodontenreste in der bayerischen

Staatsammlung in München. Palaeontographica 99, (Stut-
tgart),

 99,  A, 4–6, 121–228.

Table 1:

 Dental measurements of the molars M

2

 sin., M

2

 dex., M

3

sin., M

3

 dex. (Anancus arvernensis ZIBA-2407/1986, HEC —

height of ectoloph, HEN — height of entoloph, scale in mm).

RIDGE  No.

1

2

3

4

5

M2 sin. width

63.1

61.2

64.0

66.1

43.8

M2 sin. HEC

30.5

35.3

43.0

44.1

28.6

M2 sin. HEN

18.3

21.4

38.7

36.7

25.4

M2 dex. width

61.6

59.2

63.6

64.7

38.2

M2 dex. HEC

29.9

34.6

42.3

44.4

38.4

M2 dex. HEN

29.1

12.5

27.8

44.3

32.5

M3 sin. width

70.1

69.2

65.2

 -

 -

M3 sin. HEC

62.2

56.4

 -

 -

 -

M3 sin. HEN

47.6

49.0

40.4

38.6

 -

M3 dex. width

68.0

68.3

73.8

 -

 -

M3 dex. HEC

55.5

54.4

57.1

 -

 -

M3 dex. HEN

43.6

 -

61.8

 -

 -

Conclusions

Anancus arvernensis

 belongs to the progressive line of

the subfamily Anancinae. The progressivness is derived
from the presence of more than four ridges on the third mo-
lars. It is bunodont tetralophodont dibelodont brevirostrin-
ous mastodon form (Holec 1985). Pentalophodonty is also
presented in this group of proboscids — Anancus from the
Sagantole Formation, Ethiopia (Kalb & Mebrate 1993). The
process of the alternation of half-ridges, typical for the third
molars of this mastodontid, is also present on the crown sur-
face of the second molars. The description of molars of the