GeolCarp_Vol67_No6_573

GEOLOGICA CARPATHICA

, DECEMBER 2016, 67, 6, 573 – 594

doi: 10.1515/geoca-2016-0036

www.geologicacarpathica.com

A new Middle Miocene selachian assemblage

(Chondrichthyes, Elasmobranchii) from the Central

Paratethys (Nyirád, Hungary): implications for

temporal turnover and biogeography

MÁRTON SZABÓ

1, 2

and LÁSZLÓ KOCSIS

MTA-ELTE Lendület Dinosaur Research Group, Pázmány Péter sétány 1/C, Budapest 1117, Hungary; szabo.marton.pisces@gmail.com

Department of Palaeontology and Geology, Hungarian Natural History Museum, Ludovika tér 2., Budapest 1083, Hungary

Geology Group, Faculty of Science, Universiti Brunei Darussalam (UBD), Brunei; laszlokocsis@hotmail.com, laszlo.kocsis@ubd.edu.bn

(Manuscript received February 23, 2016; accepted in revised form September 22, 2016)

Abstract: A new Middle Miocene (Langhian – early Serravallian) assemblage with shark and ray teeth from Nyirád

(Hungary, Transdanubia, Veszprém County) consists of nine families, with 15 different species. The assemblage shares

many common genera with other Middle Miocene assemblages in the Paratethys (Notorynchus, Carcharias, Otodus,

Cosmopolitodus, Hemipristis, Galeocerdo, Carcharhinus, and Aetobatus), and reflects a subtropical climate and a close

connection with the Mediterranean Sea. However, a detailed faunal compilation of Miocene selachians reveals that

several taxa that were still present in the Mediterranean or lived in the Paratethys during the Lower Miocene disappeared

or became very rare by the Middle Miocene in the Central Paratethys (e.g., Isistius, Centrophorus, Mitsukurina,

Carcharoides, Parotodus, Alopias). The taxa that went locally extinct in the Paratethys are mainly represented by

deep-water or pelagic forms. Their disappearance is most probably related to the gradual separation of the Paratethys

from the Mediterranean. The common presence of some large, rather pelagic sharks (e.g., Otodus, Cosmopolitodus) in the

Central Paratethys during the Middle Miocene is explained here by the widespread occurrence of their potential prey

represented by marine mammals (e.g., whales and dolphins).

Key words: Badenian, Central Paratethys, Hungary, Nyirád, Chondrichthyes, Carcharoides.

Introduction

The Central Paratethys was a part of a large epicontinental sea,

the Paratethys, which was isolated from the Tethys Ocean

during the late Eocene – early Oligocene (e.g., Báldi 1983,

Rögl & Steininger 1983; Rögl 1998). The separation was

driven by the Alpine orogeny, but global sea level changes

also played important roles in opening and closing seaways

towards the open ocean or between the different sub-basins.

Complete isolation and reopening of oceanic gateways

occurred repeatedly and led to development of a distinct

palaeobiological province (e.g., Báldi 1983; Rögl 1998).

Fossil remains of cartilaginous fishes (e.g., shark and ray

teeth) are often found in the Miocene sediments of the Para-

tethys and many field-reports and scientific papers have men-

tioned them since the 19

century (see references in Koch

1903 and Schultz 1971). Generally, in earlier times many fossil

species were described (e.g., Agassiz 1843; Probst 1878,

1979), however comparative studies with modern relatives has

allowed revising of several previously described fauna and the

number of species were reduced in the Paratethys as well

(Leriche 1927a,b; Vitális 1942; Schultz 1971; Holec et al.

1995; Kocsis 2007; Schultz et al. 2010).

Here a new Middle Miocene (Badenian) fauna is described

from Nyirád (Veszprém County, Hungary). This locality is

situated in the Transdanubian Range from where our know-

ledge on chondrichthyan fossils is very sporadic so far

(Fig. 1). On the other hand, from other sub-regions of the

Central Paratethys the marine Badenian beds often yielded

rich shark and ray faunas, among them the Vienna Basin (e.g.,

Schultz 1971, 2013; Holec 2001), Molasse Basin (Schultz

2003), Styrian Basin (Hiden 1995), the Carpathian Foredeep

(Radwański 1965; Schultz 1977; Brzobohatý & Schultz 1978;

Wysocka et al. 2012; Reinecke & Radwański 2015) and

recently Slovenia (Mikuž 2009; Mikuž & Šoster 2013; Mikuž

et al. 2015).

Regarding the Hungarian Badenian beds, so far no detailed

investigation was reported on the cartilaginous fish fauna.

From northwest Hungary the literature only provides fauna

lists (Ferenczi 1915; Noszky 1925; Kordos & Solt 1984),

sometimes with photo tables (Solt 1987). Regarding southern

Hungary there are some patchy Badenian shark and ray teeth

occurrences, but a very rich re-worked fauna is known from

the Pannonian (Late Miocene) freshwater deposits, still with-

out comprehensive taxonomical descriptions (Kazár et al.

2001; Sebe et al. 2015). The age of these selachian and ray

fossils is assumed to be Karpatian – Badenian (Kocsis 2002;

Sebe et al. 2015).

The lack of taxonomical and well-illustrated studies from

the Hungarian Badenian beds is evident. The aims of this

575

NEW MIDDLE MIOCENE SELACHIAN ASSEMBLAGE FROM THE CENTRAL PARATETHYS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

from boreholes and surface outcrops. It is made up of

the following sediments: abrasional gravel and con-

glomerate, shallow water – nearshore calcareous glauc-

onitic sand, sandstone and silt as well as clay marl and

marl consisting of coral linacean limestone lenses.

In some places tuff, tuffite or bentonite interbeddings

occur in the succession. Several transitions from the

looser, sandy-pebblic variation to the strongly

cemented, hard-faced, lime-bounded conglomerate

can be observed.

The sediments are grey, yellowish-grey, often with

reddish-brown tint derived from limonite (Selmeczi

1996, 2003; Selmeczi et al. 2002). The size of the peb-

bles ranges from 0.5 – 2 cm up to 10 – 15 cm; the quan-

tity of the latter is sub ordinated (in the wider area the

average size of the pebbles is 1– 2 cm). The pebbly

sediment of the Pusztamiske Formation is rich in

fossils (e.g., molluscs, mostly Pectinidae; Balanus

sp.-fragments; chondrichthyan and osteichthyan

remains, rarely sea mammals). There is a gradual tran-

sition from the Kolontár Member of the Pusztamiske

Formation into the Pécsszabolcs Member of the Leitha

Limestone Formation, which is very rich in fossil

remains. The Leitha Limestone is a common shallow

water facies in the Central Paratethys. It was named

after the Leitha Mountain in Austria (i.e., Leitha-kalk),

but in literature it is often referred to as coralli nacean

limestones, lithothamnium limestone or algal lime-

stone. In Hungary two members are distinguished: the

older Pécsszabolcs Member corresponding to late

Lower-Middle Badenian, while the Rákos Member

is Upper Badenian (Gyalog & Budai 2004). Chon-

drichthyan remains are often reported from these

limestones (see Table 2), but at Nyirád the majority of

the teeth come from the Pusztamiske Formation

underneath.

Regional outlook

The Middle Miocene Badenian stage of the Central

Paratethys generally corresponds to the Langhian and

early Serravallian age on the global chronostrati-

graphic chart (e.g., Kováč et al. 2007; Rögl et al.

2008). However, recently Hohenegger et al. (2014)

argued that the lower boundary of the Badenian

predates the Langhian/Burdigalian boundary by about

300 kys (Fig. 2). Nevertheless, the subdivision of the

Badenian is often confusing due to two- (lower and

upper) or threefold (lower–middle–upper or early–

middle–late) subdivisions, where also the boundaries

vary regarding different biostratigraphic approaches

and depending on the authors (e.g., Papp & Cicha

1968; Rögl 1998; Harzhauser et al. 2003; Piller et al.

2007). This often makes it difficult to correlate sites

from where chondrichthyan fossils are reported.

Table 1

: Compilation of Badenian fossil Chondrichthyes localities from the Central Paratethys. Regions (capital letters) with the most important localities (numbers) are plotted on the map in Fig. 1.

Region

Place

Selachian Lit

atur

Lithology and/

or Lithos

atigr

aph

e - Z

one

e (Ma)

Lit

atur

e on Str

atigr

aph

Vienna Basin (A)

Baden-Soos (s

otype loc

ality f

or the Badenian) (5)

Rögl e

t al. 2008; Schultz, 2013

Baden Fm. Cla

y - "Badener T

el"

Upper Lag

enidae Z

one - NN5

14.2

Rögl e

t al. 2008

Aus

tria

: e.

g. St. Mar

gar

ethen (2), Bruck a.d. Leitha (3), Bad V

öslau (4) ,

Hungar

y: F

ert

őr

ák

os (1)

Nos

zky 1925; Schultz 1971, 2013; Brz

oboha

tý & Schultz 1978; Schmid e

t al. 2001

Schultz, 2013

cor

allinacean limes

tones

a Leitha limes

tone

Upper Lag

enidae Z

one - NN5

Schmid e

t al. 2001,

Harzhauser e

t al. 2003

Slo

vakia:

vínsk

a K

yla (6), De

vinsk

a No

va V

es (7)

Nos

zky 1925; Schultz 1971, 2013; Brz

oboha

tý & Schultz 1978; Holec 2001

Studienk

a and Sandber

g F

orma

tions

Bolivina-Bulimina Z

one - NN6

13.58

žn

ý e

t al. 2012

ech R

epublic

: Kienber

g a

t Mik

ulo

v (8)

Schultz e

t al. 2010; Schultz 2013

Lanžhot F

orma

tion

Hrušky F

orma

tion

Upper Lag

enidae Z

one - NN5

Brz

oboha

tý e

t al. 2007

Molasse Basin (B)

Grund near Hollabrunn (9), Mühlbach am Manhartsber

g (10),

Schultz 2003, 2013; Da

xner

-Höck e

t al. 2004

Grund F

orma

tion

Gaindorf F

orma

tion

er Lag

enidae Z

one - NN5

15–15.1

Harzhauser e

t al. 2003,

xner

-Höck e

t al. 2004,

Ćorić e

t al. 2004

Styrian Basin (C)

g., St. Florian (1), T

obbiseg

g (2), W

eisseneg

g (3)

Brz

oboha

tý & Schultz 1978; Hiden 1995; Schultz 2013

eisseneg

g F

orma

tion

St. Florian Member

er Lag

enidae Z

one - NN5

Hiden 1995, Harzhauser e

t al. 2003

va Basin

Slo

venia

(D): Plesk

o quarr

y, near T

rbo

vlje (1);

eho

vic

a, Dolenjsk

a (2); Vir

št

anj (3)

Mik

už e

t al. 2013,

Mik

už 2009; Mik

už & Šos

ter 2013; Mik

už e

t al. 2015

marls

tones and lithothamnium limes

tones

marls

tone

upper part of NN5 ?Upper Badenian

Mik

už & Bart

ol 2011

Croatia

: Medv

ednic

Mtn.

apuk Mtn.

Lithothamnium limes

tone

Sandy depoits

Lag

enidae Z

one - NN5

mos

t are re

work

ed --> ?Badenian

Ćorić e

t al. 2009

Pannonian Basin (Hung

Bak

y Moun

tains

Nyir

ád

this s

tudy

Pus

amisk

e Fm. &

Leitha Limes

tone Fm. - P

écss

zabolcs Member

Lag

enidae Z

one - NN5

Selmeczi 2003; K

csmár e

t al. 2015

alog & Buda 2004

Várpalot

a (Sz

abó-bán

ya)

not s

tudied y

et - pr

eliminar

y f

auna lis

t giv

en her

Pus

amisk

e Fm. &

Leitha Limes

tone Fm. - P

écss

zabolcs Member

Lag

enidae Z

one - NN5

tona e

t al, 2011; K

ók

, 2013;

(E) Mátra & Cserhát

Moun

tains

Má

zőllős (1); Sámsonház

a (2); Má

ebély (3);

?Má

ahas

znos;

Nos

zky 1925; Vit

ális 1915, 1942; Solt 1987, 1991

Leitha Limes

tone Fm. - P

écss

zabolcs Member

er Badenian ~Lag

enidae Z

one

Müller 1984, Moisse

tte e

t al. 2007,

alog & Buda 2004

Budapes

t and

its region

Zebeg

én

y (4), K

emence, T

ör

ökme

ző

Őr

s v

ér t

ér (5); Rák

os - Budapes

Solt & K

dos 1984

Leitha Limes

tone Fm. - P

écss

zabolcs Member

Leitha Limes

tone Fm. - Rák

os Member

er Badenian ~Lag

enidae Z

one

te Badenian ~ Bolivina-Bulimina Z

one

Müller 1984, Moisse

tte e

t al. 2007,

alog & Buda 2004

(F) Mecsek Moun

tain

ehely (1), F

asboda (2)

Sebe e

t al. 2015

Pécs

zabolcs Limes

tone Fm.

"Lo

er Leitha limes

tone"

Sebe e

t al. 2015

Danitz-pus

a (3)

Kaz

ár e

t al. 2001; K

ocsis 2002; Juhás

z 2006; Sebe e

t al. 2015

Pannonian sand with r

edeposit

ed fish r

emains

work

ed f

auna--> Badenian

Sebe e

t al. 2015

Hímesház

a (4)

Kocsis 2002

Pannonian sand with r

edeposit

ed fish r

emains

work

ed f

auna--> Badenian

Carpa

thian F

edeep

Poland

(G): K

ytnic

a (1)

Schultz 1977, 1979, 2013; R

eineck

e & Radw

ański 2015

Kor

ytnic

a Cla

er Lag

enidae Z

one

Rögl & Br

ands

tä

tter 1993

Pińcz

ów (2)

wlo

a 1960; Radw

anski 1965

"Lo

er Leitha limes

tone"

er Badenian ~ Lag

enidae Z

one

Ukraine

(H): e.

g. Khor

osno (1), Glebo

viti (2), Pidiarkiv

, Ber

ezhan

ysock

a e

t al. 2012

Myk

olaiv Sands - c

val with K

ytnic

a Cla

esence of

Orbulina suturalis

~ Lo

er Lag

enidae Z

one - NN5

Andr

a-Grig

ovich e

t al. 1997

ysock

a e

t al. 2002

ans

ylv

anian Basin

Gârbo

va de Sus (F

elsö-Orbó)

Nos

zky 1925

Sand & "Leitha limes

tone"

Gârbo

va de Sus Fm.

Upper Lag

enidae Z

one - NN5

Filipescu & Grî

bacea 1997

The c

apit

al le

ers and numbers are re

ferenc

es t

o Figure 1.

581

NEW MIDDLE MIOCENE SELACHIAN ASSEMBLAGE FROM THE CENTRAL PARATETHYS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

1843 Lamna (Odontaspis) acutissima sp. nov.; Agassiz: 294, pl. 37a:

33, 34.

1970 Odontaspis acutissima; Cappetta: pl. 1: 1-22. and pl. 2: 1-16.

1978 Odontaspis (Synodontaspis) acutissima acutissima (Ag.);

Brzobohatý and Schultz: 443, pl. 1: 12, 13.

1995Carcharias acutissima (Agassiz, 1844); Hiden: 57-58, pl. 1: 1, 2.

1995 Synodontaspis acutissima (Agassiz, 1844); Holec et al.: 40-41,

pl. 10: 3-5. and pl. 11: 1, 3.

2001 Carcharias acutissimus (Agassiz, 1844); Holec: 119-121, pl. 1:

6b, 7.

2007 Carcharias acutissima (Agassiz, 1843); Kocsis: 31-32, fig. 4.

6-11.

2013 Carcharias acutissimus (Agassiz, 1843); Schultz: 55-60, pl. 5:

7(a+b), 8(a+b)

2014 Carcharias acutissimus (Agassiz, 1843); Pollerspöck and

Beaury: 30-32, pl. 1: 7a, b

2015 Carcharias taurus Rafinesque, 1810; Reinecke and Radwański:

9, pl. 1:D-G

Collected tooth material: 11 teeth (Leitha Limestone Fm.:

MFGI V 2014.93.1-3., MFGI V 2014.96.1.; Pusztamiske Fm.:

MFGI V 2014.92.3.1-3., MFGI V 2014.91.1., MFGI V

2014.94.1., MFGI V 2014.95.1., MFGI V 2014.113.3.).

The species appeared in the Lutetian (Eocene), and it

became widespread in the Miocene epoch (Cappetta 2012).

The dentition is strongly heterodont, which is typical for the

genus (e.g., Taniuchi 1970

;

Purdy et al. 2001).

The crown is weakly convex on the labial, while, strongly

convex on the lingual face. The root is bifurcated with well

developed nutritive groove, and strong internal bulge. The

teeth have mostly one pair, but sometimes two pairs of

cusplets. The cutting edges are smooth both mesially and

distally and they run along the crown.

The anterior teeth (see Fig. 6E–H) are slender, thin, elon-

gated, the shape of the main crown is typically “S”-like in

lateral view. The cusplets of the anterior teeth are pointed,

thick at their bases and round in section (Cunningham 2000;

Antunes & Balbino 2003). The main cusp of the upper lateral

teeth (see Fig. 6 I, J) are straighter in lateral view, the cusplets

of these teeth are mostly flattened labiolingually. The crown is

thick at the base, and it slightly bends distally. On some lateral

teeth there are two pairs of cusplets (Antunes & Balbino

2003), although it is not typical.

The lingual side of the main crown is slightly folded verti-

cally, especially on the anterior teeth. This striation is not as

strong as in the family Mitsukurinidae (goblin sharks), more-

over the taxonomical relevance of this feature for the Carcha

rias teeth is questionable (Kocsis 2007). The striation is more

visible on the juvenile teeth, and it extends beyond the half of

the height of the main crown, while this striae pattern on the

teeth of adult animals is not very visible, and it becomes

weaker at about the middle height of the main crown (Cappetta

1970; Antunes & Balbino 2003).

Odontaspididae indet.

Collected tooth material: 302 teeth (Leitha Limestone Fm.:

MFGI V 2014.117.10.1-10., MFGI V 2014.101.4.1-4.;

Pusztamiske Fm.: MFGI V 2014.97.85.1-85., MFGI V

2014.100.39.1-39., MFGI V 2014.104.20.1-20., MFGI V

2014.105.2.1., MFGI V 2016.20.1.).

Tooth remains of Odontaspididae sharks (sand tiger sharks)

are typical chondrichthyan remains of the Miocene marine

sediments worldwide. The anterior teeth are easily distin-

guishable by their sigmoid profile. The lateral teeth are more

blade-like, they bend distally, but they are less curved in lateral

view than the anterior teeth.

At Nyirád the teeth of this family are the most common

(altogether 315 specimens), however, most of them can be

identified only as indeterminate Odontaspididae, because of

their missing root and cusplets.

Family: Lamnidae Müller and Henle, 1838

Genus: Carcharoides Ameghino, 1901

Carcharoides cf. catticus (Philippi, 1846)

Fig. 7 A– G

1846 Otodus catticus sp. nov.; Philippi: 24, pl. 2: 5-7.

1879 Otodus debilis sp. nov.; Probst: 155, pl.2: 78-81.

1970 Lamna cattica; Cappetta: pl. 4: 1-9.

1995 Carcharoides catticus (Philippi, 1846); Holec et al.: 42, pl. 12: 2.

2007 Carcharoides catticus (Philippi, 1851); Kocsis: 33, fig. 5. 1-3.

2011 Carcharoides catticus (von Philippi, 1846); Vialle et al.: 246,

fig. 2. 10.

Collected tooth material: 14 teeth (Pusztamiske Fm.: MFGI

V 2016.38.1., MFGI V 2016.39.1., MFGI V 2016.40.1., MFGI

V 2016.41.1.).

The Carcharoides remains from Nyirád are very fragmen-

tary. The material consists of isolated main crowns (Fig. 7A–C

and E–G) and other isolated, asymmetrical cusplets (typical

feature for lateral to distal teeth; see Fig. 7D), therefore we

refer to them only as Carcharoides cf. catticus. The pointed

anterior teeth are straight, relatively high and symmetrical,

while the laterals and the distals are distally bent, with asym-

metrical main crown and cusplets. The cutting edge is smooth,

a feature that distinguishes the species from Carcharoides

totuserratus (Ameghino, 1901). The main crown of the ante-

rior files is a little bit more concave on both faces, than those

of the lateral to distal files. The cusplets are relatively big,

pointed and roundish in cross section on anteriors, while trian-

gular and flattened on lateral-distal teeth. On lateral-distal

teeth the main cusp is pointed, flattened labiolingually, and

just weakly concave on both faces. Of these teeth the cutting

edges of the main cusp run down to the crown-root boundary,

where they connects to those of the cusplets.

The teeth of this extinct lamnoid shark are known from

several Tertiary sediments of Europe. They have been reported

from Hungary too, from the Eggenburgian of Ipolytarnóc

(Kocsis 2007), and from the Rupelian to early Chattian Kiscell

Clay Formation of the Buda Hills (Weiler 1933, 1938), which

is one of the oldest records of the species (see Reinecke et al.

2014). Regarding the Paratethyan shark faunas, this is the very

first Carcharoides discovery from the Badenian. The habitat

of this shark is not well known, but the sudden rarity of this

583

NEW MIDDLE MIOCENE SELACHIAN ASSEMBLAGE FROM THE CENTRAL PARATETHYS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

1843 Oxyrhina hastalis sp. nov.; Agassiz 1843: 277-278, pl. 34: 3,

6,13-18.

1879 Oxyrhina hastalis Ag.; Probst: 129-131, pl. 2: 1-6.

1965 Oxyrhina hastalis Agassiz, 1843; Radwański: 269-270, pl. 1: 3.

1970 Isurus hastalis; Cappetta: pl. 5: 1-13.

1978 Isurus hastalis hastalis (Ag.); Brzobohatý and Schultz: 443, pl.

2: 18, 19.

1995 Isurus hastalis (Agassiz, 1843); Holec et al.: 42-43, pl. 12: 4.

2007 Isurus hastalis (Agassiz, 1843); Kocsis: 34-35, fig. 5.7-8.

2010 Cosmopolitodus hastalis (Agassiz, 1843); Schultz et al.: pl. 1: 9-11.

2013 Cosmopolitodus hastalis (Agassiz, 1838); Mikuž et al.: 122-

125, pl. 1: 1, 2.

2013 Cosmopolitodus hastalis (Agassiz, 1843); Schultz: 43-47, pl. 4:

17, 21, 22(a+b)

2013 Cosmopolitodus hastalis (Agassiz, 1838); Šoster and Mikuž:

78, pl. 3: 19, 20.

Collected tooth material: 10 teeth (Leitha Limestone Fm.:

MFGI V 2014.11.5.1-2.; Pusztamiske Fm.: MFGI V

2014.114.5.1-5., MFGI V 2016.2.1.).

The genus is known from the Lower Miocene and wide-

spread till the late Pliocene (Cappetta 2012). The crown of the

upper teeth is strongly flattened labiolingually. The lower

anterior teeth are narrower, and more convex in their lingual

face, than the upper anteriors. The tooth crown is not consi-

derably thick on the upper teeth, but the lower dentition has

visibly thicker crowns. The thickness of the crown of all teeth

reaches its maximum near the root (Holec et al. 1995). Both

sides of the crown have smooth surface, the cutting edges run

from the apex to the crown-root boundary both on the mesial

and the distal side. The carinae are not serrated, cusplets can-

not be observed.

The crown of the anteriors is straight and is more elongated

apicobasally than that of the laterals (see Fig. 7H). The lateral

(see Fig. 7I) and anterolateral (see Fig. 7J) teeth have a typi-

cally triangle shaped crown. The distal teeth (see Fig. 7K) are

curved distally. The root is strongly bifurcated on the lower

teeth, but weakly bifurcated and mesiodistally widened on the

teeth of the upper dentition.

Family: Otodontidae Glikman, 1964

Genus: Otodus Agassiz, 1838

Subgenus: Otodus (Megaselachus) Glikman, 1964

Otodus (Megaselachus) megalodon (Agassiz, 1843)

Fig. 7 L – P

1843 Carcharodon megalodon sp. nov.; Agassiz: 247-249, pl. 29:

1-7.

1970 Procarcharodon megalodon; Cappetta: pl. 6: 2.

1978 Procarcharodon megalodon megalodon (Ag.); Brzobohatý and

Schultz: 443, pl. 3: 23.

1979 Procarcharodon megalodon (Agassiz, 1843); Schultz: 291, pl.

1: 1.

1995 Carcharocles megalodon (Agassiz, 1843); Hiden: 61-62, pl. 2,

pl. 3: 1, 2.

1999 Carcharocles megalodon (Agassiz, 1843); Mikuž: 144-146, pl.

1: 1a, b

2001 Carcharocles megalodon (Agassiz, 1843); Holec: 123,

pl. 3: 2.

2007 Carcharocles sp.; Kocsis: 34, fig. 5. 10.

2010 Megaselachus megalodon (Agassiz, 1835); Schultz et al.; pl. 1: 1, 2.

2012 Otodus (Megaselachus) megalodon (Agassiz, 1835); Cappetta:

224-227, text-fig. 210.

2013 Megaselachus megalodon (Agassiz, 1835); Schultz: 70-75, pl.

6: 1(a+b)-6(a+b)

2015 Megaselachus megalodon (Agassiz, 1835); Mikuž et al.: 79-83,

pl. 1-8 with all figures

Collected tooth material: 2 tooth fragments (MFGI V

2014.90.2.1-2.).

This species is the biggest, currently known macropredatory

shark that ever lived (Pimiento et al. 2010). Its massive tooth

remains are probably the most spectacular shark tooth fossils

of the Miocene sediments worldwide. Fossils of the species

were reported from the Middle Miocene to Pliocene, the

newest results show that this shark species went extinct around

2.6 million years ago, around the Pliocene-Pleistocene boun-

dary (Pimiento and Clements, 2014). The taxonomic assign-

ment of this shark species has been debated for decades, but

Cappetta (2012) classified it into the genus Otodus Agassiz,

1838, then he separated the genus into three subgenera: Otodus

(Otodus) Agassiz 1838, Otodus (Carcharocles) Jordan and

Hannibal 1923 and Otodus (Megaselachus) Glikman 1964

(Cappetta 2012).

The lingual side of the crown is strongly convex, while the

labial side is typically flat. The cutting edges are strongly serra-

ted in their full length both on the mesial and distal sides. The

root is bifurcated and usually symmetric. The crown of the ante-

riors is high, triangle shaped, wide with clearly visible crown-

root boundary. Going backwards distally the height of the teeth

reduces, and the crowns get more curved distally as well.

Most of the Otodus (Megaselachus) megalodon teeth fossils

from Nyirád were collected by private collectors.

Order: Carcharhiniformes Compagno, 1973

Family: Hemigaleidae Hasse, 1879

Genus: Hemipristis Agassiz, 1843

Hemipristis serra Agassiz, 1843

Fig. 8 A–E

1843 Hemipristis serra sp. nov.; Agassiz: 237, pl. 27: 18-30.

1970 Hemipristis serra; Cappetta: pl. 11: 1-18.

1878 Hemipristis klunzingeri sp. nov.; Probst: 146-149, pl. 1: 58-63.

1878 Hemipristis serra Ag.; Probst: 143-146, pl. 1: 49-57.

1978 Hemipristis serra Ag.; Brzobohatý and Schultz: 443, pl. 1: 6.

1995 Hemipristis serra Agassiz, 1843; Hiden: 64-65, pl. 4: 1-2.

1995 Hemipristis serra Agassiz, 1843; Holec et al.: 45-46, pl. 16: 1-4.

and pl. 17: 1-3.

2001 Hemipristis serra Agassiz, 1843; Holec: 127, 129, pl. 2: 10. and

pl. 3: 1.

2007 Hemipristis serra Agassiz, 1843; Kocsis: 36, fig. 6. 5-6.

2010 Hemipristis serra Agassiz, 1835; Schultz et al.; pl. 1: 3-5.

2013 Hemipristis serra Agassiz, 1835; Schultz: 90-93, pl. 7: 5(a+b),

8(a+b)-10(a+b)

Collected tooth material: 4 teeth (Pusztamiske Fm.: MFGI

V 2014.110.4.1-4.).

Snaggletooth sharks typically have dignathic heterodont

dentition. The upper teeth are labiolingually flattened, distally

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bent, triangle shaped with wide crown and visibly serrated cut-

ting edges, except the symphyseals. The distal cutting edge of

these teeth is concave, while the mesial is strongly convex.

These features make them similar to the teeth in the distal third

of the lower jaw. The lower and upper symphyseals, the lower

anteriors and the first few lateral teeth have an awl-like con-

tour in lingual view. The cutting edge is not serrated on these

lingually curved teeth, and it runs only to the apical half of the

tooth crown. Several cusplets can be also observed on these

teeth.

The tooth remains of Hemipristis serra are common and

abundant worldwide in Miocene marine sediments. The modern

species of the genus is the Hemipristis elongata (Klunzinger,

1871). Although the dentition of H. elongata and H. serra is

closely similar, the main difference between them are the more

numerous serrations on the upper laterals of the modern

species. Additionally, an increase in tooth size with age

was also reported for the genus from the Lee Creek Mine

(Chandler et al. 2006).

Family: Carcharhinidae Jordan and Evermann, 1896

Genus: Carcharhinus Blainville, 1816

Carcharhinus priscus (Agassiz, 1843)

Fig. 8 F–L

1843 Sphyrna prisca sp. nov.; Agassiz: 234-235, pl. 26a: 44, 47.

1970 Carcharhinus priscus; Cappetta: pl. 13:1-20 and pl. 14: 1-20.

1978 Carcharhinus priscus (Ag.); Brzobohatý and Schultz: 442, pl. 1: 9.

1995 Carcharhinus priscus (Agassiz, 1843); Hiden: 65-66, pl. 5: 2.

1995 Carcharhinus priscus (Agassiz, 1843); Holec et al.: 46, pl. 18:

1, 2.

1995 Carcharhinus similis (Probst, 1878); Holec et al.: 46-47, pl. 18: 3-4.

2001 Carcharhinus priscus (Agassiz, 1843); Holec: 123-125, pl. 2: 4, 6.

2001 Carcharhinus similis (Probst, 1878); Holec: 125, pl. 2: 5.

2007 Carcharhinus priscus (Agassiz, 1843); Kocsis: 36-38, fig.

6.7-12.

2010 Carcharhinus priscus (Agassiz, 1843); Schultz et al.; pl. 2: 11.

2013 Carcharhinus priscus (Agassiz, 1843); Schultz: 80-84, pl. 7:

6(a+b), 7(a+b)

2015 Carcharhinus priscus (Agassiz, 1843); Reinecke and Rad-

wański: 13, pl. 4: A-R

Collected tooth material: 28 teeth (Leitha Limestone Fm.:

MFGI V 2014.106.2.1-2.; Pusztamiske Fm.: MFGI V

2014.108.10.1-10., MFGI V 2016.22.1.).

The genus has a typical dignathic heterodonty in its denti-

tion. The lower dentition of the different species can be very

similar, and therefore the species are mainly best distinguished

by their upper dentition (see e.g., Bourdon 1997–2013,

Cappetta 2012). The lower teeth are simpler than the upper

ones, but their corresponding counterpart is similar in size.

The tooth crown of this genus is typically not higher than

the width of the tooth (anteriors are the highest), it is narrow

triangle shaped (sometimes weakly bent distally) on the ante-

rior teeth, while strongly bent distally on the laterals.

The cutting edges run downward, and continue on the lobes of

the root on both sides, while creating the enamel-shoulders.

The serration of the cutting edges is strongest on the

enamel-shoulders. The root is widened mesiodistally, usually

there is an axial groove on the middle of the lingual side.

The genus is known from the Lutetian (Cappetta 2012) and

become widespread and radiated during the Miocene and

Pliocene. The C. priscus was a very common form in the

Palaeo-Mediterranean Sea and the Paratethys. In the Lower

and Middle Miocene sometimes another species, C. similis is

mentioned (Probst 1878, Barthelt et al. 1991; Holec et al.

1995, 2001), and its teeth are often confused with the

C. priscus (see the synonym list).

Genus: Galeocerdo Müller and Henle, 1837

Galeocerdo aduncus Agassiz, 1843

Fig. 8 M–R

1843 Galeocerdo aduncus sp. nov.; Agassiz: 231, pl. 26: 25-28.

1970 Galeocerdo aduncus; Cappetta: pl. 12:1-21.

1978 Galeocerdo aduncus Ag.; Brzobohaty and Schultz: 442-443, pl.

1: 10.

1995 Galeocerdo aduncus Agassiz, 1843; Hiden: 66-67, pl. 4: 4.

1995 Galeocerdo aduncus Agassiz, 1843; Holec et al.: 47-48, pl. 19:

1, 2, 4-6.

2001 Galeocerdo aduncus Agassiz, 1843; Holec: 125-127, pl. 2: 9.

2007 Galeocerdo aduncus Agassiz, 1843; Kocsis: 38, fig. 6.13-14.

2010 Galeocerdo aduncus (Agassiz, 1835); Schultz et al.: pl. 1: 6-8.

2013 Galeocerdo aduncus (Agassiz, 1835); Schultz: 84-87, pl. 7:

11(a+b), 12(a+b)

2015 Galeocerdo aduncus Agassiz, 1835; Reinecke and Radwański:

12-13, pl. 1: H-I

Collected tooth material: 9 teeth (Leitha Limestone Fm.:

MFGI V 2014.111.2.1-2.; Pusztamiske Fm.: MFGI V

2014.112.6.1-6., MFGI V 2014.105.2.2.).

The genus is known from the early Eocene. The species

G. aduncus appeared in the early Oligocene, and became

widespread in the Miocene Epoch (Cappetta 2012).

The dentition of the upper and the lower jaw is very similar

(Compagno 1984), but the teeth sitting in the same jaw can be

distinguished by their position (monognathic heterodonty).

The teeth reduce in height and increase in width distally from

the symphysis (Kocsis 2007). The mesial cutting edge is

convex and visibly serrated, while the distal cutting edge

wears a deep notch, which separates the main cusp from the

distal shoulder of the tooth. The tooth-shoulder wears a mas-

sive, distally reducing serration. The root lobes are slightly

curved lingually.

Genus: Negaprion Whitley, 1940

Negaprion sp.

Fig. 8 S–T

Collected tooth material: 1 tooth (Pusztamiske Fm.: MFGI

V. 2014.107.1.).

The genus is known from the Lower Miocene (Burgidalian)

and it exists today as well with two modern species: Negaprion

brevirostris (Poey, 1868) and Negaprion acutidens (Rüppell,

1837) (Compagno 1984; Cappetta 2012; Pimiento et al.

2013b).

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The teeth of the genus have unserrated, sometimes lingually

curved, pointed crowns perpendicular or nearly perpendicular

to the root. The labial side of the crown is flat, while the lin-

gual side is convex. The root runs mesiodistally, the axial

groove usually clearly visible with a central foramen.

Negaprion tooth remains are very similar to the lower teeth

of the Carcharhinus genus, but according to Pimiento et al.

(2013a,b), they are easy to distinguish from them by their

smooth, unserrated cutting edge, and their thicker root. How-

ever, many modern Carcharhinus species have lower teeth

with smooth cutting edge (Bourdon 1997-2013). Never theless,

based on the somewhat bulkier root, this tooth from Nyirád is

placed under this genus.

Carcharhinidae indet.

Collected tooth remains: 118 teeth (Leitha Limestone Fm.:

MFGI V 2014.99.1.; Pusztamiske Fm.: MFGI 2014.103.23.1-

23., MFGI V 2016.43.1.).

Requiem shark remains are abundant fossils in Miocene

marine sediments worldwide. The fragmentary tooth remains

of this family at Nyirád are small, none of them is bigger than

5 mm. On some teeth slight serration can be observed on the

cutting edges. The anterior crowns are straight, the laterals are

weakly curved, and all remains are pointed. The roots and

enamel-shoulders of all of these teeth are missing.

Family: Sphyrnidae Gill, 1872

Genus: Sphyrna Rafinesque, 1810

Sphyrna cf. zygaena (Linnaeus, 1758)

Fig. 8 U–V

2001 Sphyrna zygaena (Linnaeus, 1758); Holec: 127, pl. 2: 7.

2001 Sphyrna zygaena (Linnaeus, 1758); Purdy et al.: 158-160, text-

figs. 59-60

2007 Sphyrna cf. zygaena (Linnaeus, 1758); Kocsis: 38, fig.

6.18a-c

Collected tooth material: 1 tooth (Pusztamiske Fm.: MFGI

V 2016.42.1.).

Hammerhead sharks are known from the early Oligocene

(Cappetta 2012), and the taxon still exists with seven extant

species (Compagno 2005).

The teeth are flattened labiolingually, the crown is wide at

its base, and it slightly bends distally. The cutting edges are

smooth both on the mesial and distal sides. The mesial cutting

edge is slightly convex, while the distal one is straight, nearly

perpendicular to the root, and it creates a deep notch near to

the base of the crown. The distal cutting edge continues behind

this deep notch on a smooth, convex enamel-shoulder. This

feature can-not be observed on the Nyirád tooth as it is quite

fragmentary. Part of the root of MFGI V 2016.42.1. is also

missing, but it typically runs mesiodistally, and it bears

a clearly visible transversal groove.

The hammerhead shark tooth from Nyirád is similar to those

of Sphyrna zygaena, however, its poor preservation allows us

to describe specimen MFGI V 2016.42.1. only as Sphyrna cf.

zygaena.

Order: Myliobatiformes Compagno, 1973

Family: Myliobatidae Bonaparte, 1838

Genus: Aetobatus Blainville, 1816

Aetobatus arcuatus (Agassiz, 1843)

Fig. 9 A– C

1843 Aetobatis arcuatus sp. nov.; Agassiz; p. 327

1965 Aetobatis arcuatus Agassiz, 1843; Radwański: 272, pl. 1: 7.

1970 Aetobatis arcuatus; Cappetta: pl. 24: 6-9.

1995 Aetobatus arcuatus (Agassiz, 1843);Hiden: 73-74, pl. 7: 3.

1995 Aetobatis arcuatus L. Agassiz, 1843; Holec et al.: 48, pl. 20: 1.

2001 Aetobatus arcuatus (L. Agassiz, 1843); Holec: 129-130, pl. 3: 5.

2011 Aetobatus arcuatus (Agassiz, 1843); Vialle et al.: 253, fig. 4.11.

2013 Aetobatus arcuatus (Agassiz, 1843); Schultz: 106-109, pl. 11:

9-13

Collected tooth material: 4 teeth (Pusztamiske Fm.: MFGI

V 2014.89.2.1-2., MFGI V 2016.24.1., MFGI V 2016.25.1.).

This species is a relatively common eagle ray species in the

Miocene marine sediments of Europe (Radwański 1965), and

its tooth remains are easily distinguishable from those of other

eagle ray taxa. Most of the typical features of the lower teeth

can be well observed on all Nyirád specimens, however, they

are all fragmentary.

The lower teeth of A. arcuatus are curved distally, they

reach their maximal length at their mediolateral midline. The

occlusal surface is smooth and shiny. The crown is separated

from the root by a lingual bulge, which runs along the poste-

rior side of the plate. The lingually bent root is made up by

numerous laminae and grooves. The root reaches its maximal

height in the midline of the tooth.

Genus: Myliobatis Cuvier, 1816

?Myliobatis sp.

Fig. 9 D –K

Collected tooth material: 44teeth (Leitha Limestone Fm.:

MFGI V 2014.98.2.1-2.; Pusztamiske Fm.: MFGI V

2014.116.22.1-22., MFGI V 2016.26.1., MFGI V 2016.27.1.,

MFGI V 2016.28.1., MFGI V 2016.29.1., MFGI V 2016.30.1.).

This eagle ray genus is widely abundant in Miocene sedi-

ments. Numerous isolated, fragmentary ?Myliobatis teeth

have been collected at the Nyirád site. They are widened

medio laterally, and slightly curved distally. The occlusal

surface is shiny and smooth, the root lobes are high and medio-

laterally flattened. The lateral edges of the tooth contour are

angled in occlusal (and basal) view.

The ?Myliobatis sp. tooth remains are distinguished from

those of Rhinoptera here by their relatively lower crown, the

lingual extension of the root and the structure of the connec-

tions between the teeth. For the Rhinoptera genus the root not

or only slightly extended lingually, and the connection between

the tooth plates is more complex (so-called “tenon and

mortise” connections; see Bourdon 2002). Still, due to the

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SZABÓ and KOCSIS

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preservation of the teeth (e.g., broken and worn-off features),

it is not impossible that some of the teeth may come from the

Rhinoptera genus.

Family: Dasyatidae Jordan, 1888

Genus: Dasyatis Rafinesque, 1810

Dasyatis cf. probsti Cappetta, 1970

Fig. 9 L –N

1970 Dasyatis probsti sp. nov.; Cappetta: 91-92, pl. 21:15-23.

1977 Dasyatis aff. probsti Cappetta, 1970; Schultz: 202, pl. 1: 2-3.

1995 Dasyatis probsti Cappetta, 1970; Hiden: 70, pl. 6: 5. and text-

fig. 8C

2011 Dasyatis probsti Cappetta, 1970; Vialle et al: 252, figs. 4.5-6.

2014 Dasyatis probsti Cappetta, 1970; Pollerspöck and Beaury: 32,

pl. 2: 7 a, b

2015 Dasyatis probsti Cappetta, 1970; Reinecke and Radwański: 14,

pl. 6: A-D

Collected tooth material: 1 tooth (placed in private collec-

tion; stratigraphical origin is unknown).

The only known Dasyatis cf. probsti tooth from Nyirád is

well preserved. The tooth has a two lobed root, both lobes are

curved lingually, and have C-shaped contour in basal view.

The crown bears a well developed transversal crest, which

separates the crown into two visors: an anterior (labial) and

a posterior (lingual) one. This feature refers the tooth as

tooth of a female specimen. The lingual visor overhangs the

root-lobes. The labial visor and the transversal crest have

weakly reticulated ornamentation. The lingual margin has

two lateral facets and a medial facet in basal view (Fig. 9N),

a feature that is referred to this species by Hiden (1995, text-

fig. 8C).

Dasyatis rugosa (Probst, 1877)

Fig. 9 O –T

1877 Raja rugosa sp. nov.; Probst: 76, pl. 1: 5, 8, 9.

1970 Dasyatis rugosa; Cappetta: pl. 21: 1-14.

2011 Dasyatis rugosa (Probst, 1877); Vialle et al.: 253, fig. 4.7.

2014 Dasyatis rugosa (Probst, 1877); Pollerspöck and Beaury: 32,

pl. 2: 8.

2015 Dasyatis cf. rugosa (Probst, 1877); Reinecke: 20-22,

fig. 12.

2015 Dasyatis rugosa (Probst, 1877); Reinecke and Radwański: 14,

pl. 7: D-F and pl.8: A-B

Collected tooth material: 9 teeth (Pusztamiske Fm.: MFGI

V 2016.34.1., MFGI V 2016.35.1., MFGI V 2016.36.1., MFGI

V 2016.37.1.).

This stingray species is abundant in the Miocene sediments

of Europe. A few female teeth are known from Nyirád, with

a relatively well preserved crown and missing root. Although

the root lobes are not preserved, the specimens have typically

robust lobes in basal view (see Hiden 1995, text-fig. 8B).

The labial visor of these teeth is ornamented with small crenu-

lations. The labial margin of the crown is smooth in basal view

(see Fig. 9Q, T and also Hiden 1995, text-fig. 8B).

Dasyatis sp.

Fig. 9 U–V

Collected tooth material: 14 teeth (Pusztamiske Fm.:

MFGI V 2016.31.1., MFGI V 2016.32.1., MFGI V 2016.33.1.).

These tooth remains clearly belong to the genus Dasyatis,

however, they are too fragmentary or worn to be identified on

the species level (they could belong to any Nyirád Dasyatis

form). Most of the remains have crown morphology typical

for female individuals. The transversal crest of one specimen

is elongated, and distally bent, which refers the tooth as a male

tooth (MFGI V 2016.31.1., see Fig. 9U–V).

Discussion

The shark and ray fauna of Nyirád includes 14 genera of

9 families, with 13 identified species: Notorynchus primigenius,

Araloselachus cuspidatus, Carcharias acutissima, Carcha

roides cf. catticus, Cosmopolitodus hastalis, Otodus (Mega

selachus) megalodon, Hemipristis serra, Carcharhinus

priscus, Galeocerdo aduncus, Negaprion sp., Sphyrna cf.

zygaena, Aetobatus arcuatus, ?Myliobatis sp., Dasyatis cf.

probsti and Dasyatis rugosa. The result indicates a moderately

diverse chondrichthyan fauna.

The dominant shark family of the ecosystem was the Odonta-

spididae (sand tiger sharks), represented by 315 teeth (due to

preservation bias 302 teeth could be identified only at family

level). In frequency the family Odontaspididae is followed by

the Carcharhinidae (requiem sharks) with 156 tooth remains.

The dominance of these families would indicate that the eco-

system was filled with smaller fishes, as avai lable potential

prey-animals. The remains of these possible prey-animals have

also been collected at the locality. The recovered ray taxa, like

the Aetobatus arcuatus, ?Myliobatis sp. and Dasyatis spp. can

be considered as part of the diet of the sharks. However, there

are abundant bony fish tooth remains at the locality as well,

such as Acanthurus sp., Dentex sp., ?Diplodus sp., Pagrus

cinctus, indeterminate Sparidae, indeterminate Tetraodontidae

and ?Trichiurus sp. teeth. Sea mammal remains have been col-

lected at the locality as well, such as an Odontoceti indet. tooth,

a vertebra fragment of a ?Sirenia, and several bone fragments

(other sea mammal fossils from the locality are also known in

private collections). However, there is still no direct evidence

for predational relations between any local shark and other ver-

tebrate taxon. Nevertheless, the top predator of this ecosystem

could have undeniably been the Otodus (Megaselachus)

megalodon, which is the biggest known macropredatory shark

of the Neogene.

Among the lifestyles of the species the nectonic (=freely

swimming) lifestyle and the tropical-subtropical distribution

dominates (Compagno 1984, 2005). The tropical-subtropical

shark fauna-elements, together with the other vertebrate and

invertebrate taxa found at the locality, represent a subtropical

climate with warm-temperate water and also indicate a con-

nection to Mediterranean marine realm. However, the rarity of

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, 2016, 67, 6, 573 – 594

the tropical genus Ginglymostoma is worth mentioning. This

taxon yielded only one tooth so far from the Central Paratethys

from this period (Hiden 1995).

The Nyirád fauna is very similar to the typical Middle

Miocene shark and ray assemblages of the Central Paratethys

(Fig. 1, Table 2) and most of the common genera are repre-

sented in our record (Notorynchus, Carcharias, Otodus, Cos

mopolitodus, Hemipristis, Galeocerdo, Carcharhinus,

Sphyrna). There are a few exceptions like Squalus or Squatina,

and a few rarely mentioned taxa (see Table 2 and e.g., Hiden

1995; Schultz 2003, 2013; Reinecke & Radwański 2015) that

have not been discovered at Nyirád. The reason for the lack of

these taxa could be (1) preservation bias for the micro- remains,

(2) the relatively low volume of sediment searched through

and/or (3) the palaeoenvironmental conditions were not sui-

table for these forms at Nyirád.

The compilation of the Badenian chondrichthyan remains

(Tables 1–2), however, also revealed that many groups dis-

appeared or became very rare in the Central Paratethys by

the middle Miocene, for example, Isistius, Centrophorus,

Mitsukurina, Carcharoides, Parotodus, Alopias. All these

genera were common in the Mediterranean during the Miocene

(Cappetta 1970; Vialle et al. 2011), and were also present in the

Paratethys during the Lower Miocene (Holec et al. 1995; Kocsis

2007; Pollerspöck & Beaury 2014). Moreover, most of them

still have modern representatives. The exceptions are the

Carcharoides that went extinct in the Langhian and the Parot

odus that died out in the Pliocene (Cappetta 2012).

It must be mentioned that Alopias is reported from a few

Karpatian and Badenian localities in Hungary (Kordos & Solt

1984), however, after re-examining these remains in the MFGI

collection these turned out to be Odontaspididae teeth. On the

other hand, very recently the presence of Alopias in the Badenian

was comfirmed by Reinecke and Radwański (2015) who

reported one tooth from this genus from Korytnica (Poland).

The disappearance or the rarity of these taxa in the Para-

tethys is quite intriguing. Most of these groups are deep-water

epi- to bathypelagic sharks (e.g., Isistius) or pelagic (e.g.,

Alopias) forms, and their vanishing most probably relates to

the gradual separation of the Paratethys from the Mediterra-

nean. Still, large predators and pelagic fishes like Otodus or

Cosmopolitodus were quite common in the Badenian. One

explanation of their presence may be linked to large prey ani-

mals namely to marine vertebrates like whales and dolphins.

Numerous fossil remains indicate the widespread present of

these animals in the Paratethys at this period (Kazár et al.

2001; Kazár & Venczel 2003; Vrsaljko et al. 2010; Banak et al.

2015). Moreover, some dolphins became endemic and also

lived in the Sarmatian period, when more or less brackish con-

ditions set in the Paratethys (Kazár et al. 2004).

Conclusion

The described shark and ray fauna from Nyirád includes 14

genera from 9 families, with 13 identified species. The fauna

is similar to other chondrichthyan assemblages reported from

the Badenian in the Central Paratethys. Comparison with

Mediterranean and older Paratethyan faunas revealed that

some deep-water and pelagic genera vanished or became very

rare by the Middle Miocene in the Central Paratethys, which is

best explained by the palaeogeographical evolution of the

region. Among these taxa, the Carcharoides is reported here

for the first time from Badenian beds of the Paratethys. Marine

mammals are proposed here as prey-animals for the still

common Badenian presence of other large sharks like Otodus

(Megaselachus) or Cosmopolitodus.

Aknowledgements: We especially thank Ildikó Selmeczi for

helping us to get to know and understand the geology of the

studied area. We thank László Makádi for his technical assis-

tance, and Emese Réka Bodor for critically reading the manu-

script and making useful suggestions for improve our work.

We are also very grateful to József Király, András Marton,

István Marton, Frank Puffer and József Szakonyi for being

cooperative and letting us study their private collections, and

taking photos of some of their fossil remains collected at the

locality. We thank Anna Rácz for her enthusiastic help, and for

finding the first local specimens of Carcharoides, Sphyrna and

Dasyatis. We would like to thank Péter Gulyás for his help

during the fieldwork. We are grateful for the help received

from Aleš Šoster regarding the Slovenian shark teeth locali-

ties, and Karmen Fio (University of Zagreb) in terms of the

Croatian Badenian faunas. The contributions from the Papuk

Geopark and Croatian Natural History Museum and Depart-

ment of Geology and Palaeontology are also much appre-

ciated. We also thank the Geological and Geophysical Institute

of Hungary (MFGI), the Hungarian Natural History Museum

(Budapest, Hungary), the Bakony Natural History Museum

(Zirc, Hungary) and personally Lajos Katona, and Árpád

David for widely helping our work via their collections or

information on the Hungarian Miocene fauna. L.K. received

support from his UBD-URC grant of UBD/PNC2/2/RG/1(325)

when this research was conducted. We are grateful to the

Readymix-Lesence Ltd. for their logistic help in the fieldwork.

References

Agassiz L. 1833–43: Recherches sur les Poissons Fossiles, Tome III

— Atlas. Neuchâtel 1–432, Table 1–83.

Ameghino F. 1901: L’âge des formations sédimentaires de Patagonie.

Anales de la Sociedad Científica Argentina 51, 20–39.

Andreyeva-Grigorovich A.S., Kulchytskiy Y.O., Gruzman A.D.,

Lozynyak P.Y., Petrashkevich M.I., Portnyagina L.O.,

Ivanina A.V., Smirnov S.E., Trofimovich N.A., Savitskaya N.A.

& Shvareva N.J. 1997: Regional stratigraphic scheme of

Neogene formations of the Central Paratethys in the Ukraine.

Geol. Carpath. 48, 123–136.

Antunes M.T. & Balbino A.C. 2003: Upper Miocene Lamniform

Selachians (Pisces) from the Alvalade Basin (Portugal). Ciências

de Terra (UNL), Lisboa 15, 141–154.

Antunes M.T., Balbino A.C. & Cappetta H. 1999: A new shark,

Galeorhinus gonclavesi nov. sp. (Triakidae, Carcharhiniformes)

591

NEW MIDDLE MIOCENE SELACHIAN ASSEMBLAGE FROM THE CENTRAL PARATETHYS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

from the latest Miocene of Portugal. Tertiary Research 19, 3–4,

101–106.

Ayres W.O. 1855: (Shark of a new generic type: Notorynchus macu

latus.) Proceedings of the California Academy of Sciences,

Series 1, 1, 72–73.

Báldi T. 1983: Hungarian Oligocene and Lower Miocene formations

[Magyarországi oligocén és alsómiocén formációk]. Akadémiai

Kiadó, Budapest, 1–293 (in Hungarian).

Banak A., Đuras M., Avanić R., Grizelj A. & Posilović H. 2015: Neo-

gene marine mammals from Vranić sand deposit. 5

Croatian

Geological Congress Program, Osijek, 24.

Barthelt D., Fejfar O., Pfeil H.F. & Unger E. 1991: Notizen zu einem

Profil der Selachier-Fundstelle Walbertsweiler im Bereich der

miozänen Oberen Meeresmolasse Süddeutschlands. Münchener

Geowissenschaftlicher Abhandlungen (A) 19, 195–208.

Berg L.S. 1958: System der Rezenten und Fossilen Fischartigen und

Fische. Hochschulbücher für Biologie, Berlin, 1–310.

Blainville H.M.D. 1816: Prodrome d’une nouvelle distribution

systématique du règne animal. Bulletin des Sciences / par la

Société Philomatique de Paris 8, 105–124.

Bonaparte C.L. 1838: Selachorum tabula analytica. Nuovi Annali

della Science Naturali Bologna 1, 2, 195–214.

Bourdon J. 1997–2013: The Life and Times of Long Dead Sharks.

http://www.elasmo.com.

Bourdon J. 2002: Myliobatoid Teeth. In: The Life and Times of Long

Dead Sharks. http://www.elasmo.com

Brzobohatý R. & Schultz O. 1978: Die Fischfauna des Badenien.

In: Papp A., Cicha I., Seneš J. & Steininger F. (Eds.): M4,

Badenien (Moravien, Wielicien, Kosovien). Chronostratigraphie

und Neostratotypen, Miozän der Zentralen Paratethys 4, Verlag

der Slowakischen Akademie der Wissenschaften, Bratislava,

441–464. .

Brzobohatý R., Nolf D. & Kroupa O. 2007: Fish Otoliths from the

Middle Miocene of Kienberg at Mikulov, Czech Republic,

Vienna Basin: their paleoenvironmental and paleogeographic

significance. Bulletin de l’Institut royal des Sciences naturelles

de Belgique, Sci. Terre, Bruxelles 77, 167–196.

Cappetta H. 1970: Les Sélaciens du Miocéne de la région de Montpellier.

Palaeovertebrata, Mémorie extraordinaire, 1–139, 27 pl.

Cappetta H. 1987: Chondrichtyes II. (Mesozoic and Cenozoic

Elasmobranchii). In: Handbook of Paleoichthyology, Vol. 3B.

Gustav Fischer Verlag, Stuttgart, New York, 1–193.

Cappetta H. 2012: Handbook of Paleoichthyology, Vol. 3E: Chon-

drichthyes. Mesozoic and Cenozoic Elasmobranchii: Teeth.

erlag Dr. Friedrich Pfeil, 1–512.

Chandler R.E., Chriswell K.E. & Faulkner G.D. 2006: Quantifying

a Possible Miocene Phyletic Change in Hemipristis (Chondrich-

thyes) Teeth. Palaeontologia Electronica 9, 1–14.

Compagno L.J.V. 1973: Interrelationships of living elasmobranchs.

In: Greenwood P.H., Miles R.S. & Patterson C. (Eds.): Inter-

relationships of Fishes. Zoological Journal of the Linnean

Society 53, 1, 15–61.

Compagno L.J.V. 1984: FAO Species Catalogue, Volume 4: Sharks of

the World. United Nations Development Programme, Rome

1–655.

Compagno L.J.V., Dando M. & Fowler S. 2005: Sharks of the World,

Collins Field Guide. Harper Collins Publishers, London, 1–368.

Cope E.D. 1867: An addition to the vertebrate fauna of the Miocene

period, with a synopsis of the extinct Cetacea of the United

States. Proceedins of the Academy of Natural Sciences of Phila

delphia 19, 138–156.

Costa S.A.R.F., Richter M., Toledo P.M. & Moraes-Santos H.M.

2009: Shark teeth from Pirabas Formation (Lower Miocene),

northeastern Amazonia, Brazil. Boletim do Museu Paraense

Emílio Goeldi Ciȇncias Naturais, Belém 4, 3, 221–230.

Ćorić S., Harzhauser M., Hohenegger J., Mandic O., Pervesler P.,

Roetzel R., Rögl F., Scholger R., Spezzaferri S., Stingl K.,

Švábenická L., Zorn I. & Zuschin M. 2004: Stratigraphy and

correlation of the Grund Formation in the Molasse Basin,

northeastern Austria (Middle Miocene, Lower Badenian). Geol.

Carpath. 55, 207–215.

Ćorić S., Pavelić D., Rögl F., Mandic O., Vrabac S., Avanić R.,

Jerković L. & Vranjković A. 2009: Revised Middle Miocene

datum for initial marine flooding of North Croatian Basins (Pan-

nonian Basin System, Central Paratethys). Geologia Croatica

62, 1, 31–43.

Cunningham B.S. 2000: Tooth study of a recent sand tiger shark,

Carcharias taurus (Rafinesque, 1810). In: Bourdon J. (Ed.): The

life and times of long dead sharks. http://www.elasmo.com.

Cuvier G.L.C.F.D. 1816: Le Règne Animal distribute d’après son

organization pour server de base à l’histoire naturelle des

animaux et d’introduction à l’anatomie comparée. Les reptiles,

les poissons, les mollusques et les annelids. Deterville, Paris,

1–532.

Daimeries A. 1889: Notes ichthyologiques, IV. Annalesde la Société

Royale Malacologique de Belgique, Bulletin de Sciences 24, 5–10.

Daxner-Höck G., Göhlich U.B., Huttunen K., Kazár E., Nagel D.,

Rössner G.E., Schultz O., Miklas-Tempfer P.M. & Ziegler R.

2004: Marine and terrestrial vertebrates from the Middle

Miocene of Grund (Lower Austria). Geol. Carpath. 55, 2,

191–197.

De Buen F. 1926: Ichthyological catalogue of the Mediterranean of

Spain and Morocco, summary of the published fish of the coast

of the Mediterranean and the near region of the Atlantic Ocean

(’sea of Spain’). Resultados de las ampafias Realizadas por

Acuerdos Internacionales. Instituto Español de Oceanografia 2,

1–211 (in Spanish).

Ferenczi I. 1915: The Zalatna-Nagyalmás Tertiary Basin. Disserta-

tions. Földtani Közlöny, XLV. Kötet, 1915 January- February-

March, booklets 1–3 (in Hungarian).

Filipescu S. & Grîbacea R. 1997: Lower Badenian sea-level drop on

the western border of the Transylvanian Basin: Foraminiferal

paleobathymetry and stratigraphy. Geol. Carpath. 48, 5, 325–334.

Gill T. 1872: Arrangement of the families of fishes, or Classes Pisces,

Marsupiobranchii, and Leptocardii. Smithsonian Miscellaneous

Collections 11, 247, 1–49.

Glikman L.S. 1964: Sharks of the Paleogene and their stratigraphic

significance. Nakua Press, Moscow, 1–229.

Gradstein F.M., Ogg J.G., Schmitz M. & Ogg G. 2012: The Geological

Time Scale 2012, vols. 1 & 2. Elsevier Science Ltd., Oxford,

1–1176.

Gray J.E. 1851: List of the specimens of fish in the collection of the

British Museum. Part I. Chondropterygii. British Museum (Nat

ural History), London, 1–160.

Gunnerus J.E. 1765: Basking shark (Squalus maximus). Beskrvenen

ved J. E. Gunnerus, Det Trondhiemske Selskabs Skrifter 3,

33–49.

Gyalog L. & Császár G. 1982: Geological map of the Bakony Moun-

tains (without the Quaternary Formations). Published by the

Geological Institute of Hungary (Magyar Állami Földtani

Intézet), 1990 (in Hungarian).

Gyalog L.& Budai T. (Eds.) 2004: Proposal for new lithostratigraphic

units of Hungary. A Magyar Állami Földtani Intézet Évi Jelentése

2002-ről, 195–232 (in Hungarian).

Harzhauser M., Mandic O. & Zuschin M. 2003: Changes in Para-

tethyan marine molluscs at the Early/Middle Miocene transition:

diversity, palaeogeography and palaeoclimate. Acta Geol. Polon.

53, 4, 323–339.

Hasse C. 1879: Das natürliche System der Elasmobranchier auf Grun-

dlage des Baues und der Entwicklung ihrer Wirbelsäule. Eine

morphologische und paläontologische Studie. I. Allgemeiner

Theil, 1–76.

592

SZABÓ and KOCSIS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

Hiden H.R. 1995: Elasmobranchier (Pisces, Chondrichthyes) aus dem

Badenium (Mittleres Miozän) des Steirischen Beckens (Öster-

reich). Mitt. Abt. Geol. und Paläont. Landesmuseum Joanneum,

Graz, Heft 52/53, 41-109.

Holec P. 2001: Chondrichthyes and Osteichthyes (Vertebrata) from

Miocene of Vienna Basin near Bratislava (Slovakia). Mineralia

Slovaca 33, 111–134 (in Slovak with english abstract).

Holec P., Hornáček M. & Sýkora M. 1995: Lower Miocene Shark

(Chondrichthyes, Elasmobranchii) and Whale Faunas (Mamma-

lia, Cetacea) near Mučín, Southern Slovakia. Geologické práce

100, 37–52.

Hohenegger J., Ćorić S. & Wagreich M. 2014: Timing of the Middle

Miocene Badenian Stage of the Central Paratethys. Geol.

Carpath. 65, 1, 55–66.

Horváth F., Bada G., Windhoffer G., Csontos L., Dombrádi E.,

Dövényi P., Fodor L., Grenerczy Gy., Síkhegyi F., Szafián P.,

Székely B., Timár G., Tóth L. & Tóth T. 2006: Atlas of the present

geodynamics of the Pannonian-Basin: Euro-conform map-series

with explanatory notes. Magyar Geofizika 47, 4, 133–137.

Huxley T.H. 1880: On the application of the laws of evolution to the

arrangement of the Vertebrata, and more particularly of the

Mammalia. Proceedings of the Zoological Society 43, 649–662.

Hyžný M., Hudáčková N., Biskupič R., Rybár S., Fuksi T., Halásová

E., Zágoršek K., Jamrich M. & Ledvák P. 2012: Devínska

Kobyla — a window into the Middle Miocene shallow-water

marine environments of the Central Paratethys (Vienna Basin,

Slovakia). Acta Geologica Slovaca 4, 2, 95–111.

Joleaud L. 1912: Gélologie et paléontologie de la Plaine du Comtat et

de ses abords. Description des terrains néogènes. Montpellier:

impr. Montane, Sicardi et Valentin 2, 255–285.

Jonet S. 1966: Notes d’ichthyologie miocène. II. Les Carcharhinidae.

Boletim do Museu e Laboratorio Mineralógico e Geológico da

Faculdade de Ciȇncias 10, 2, 65–88.

Jonet S. 1968: Notes d’ichthyologie miocène portugaise. V. Quelques

Batoïdes. Revista da Faculdade de Ciȇncias da Universidade de

Lisboa 15, 2, 233–258.

Jordan D.S. 1888: A manual of the vertebrate animals of the northern

United States, including the district north and east of the Ozark

mountains, south of the Laurentian hills, north of the southern

boundary of Virginia, and east of the Missouri river, inclusive of

marine species. 5th edition. A.C. McClurg amd Company, Chi-

cago, i–iii. + 1–375.

Jordan D.S. & Evermann B.W. 1896: The fishes of North and Middle

America, a descriptive catalogue of the species of fish-like

vertebrates found in the waters of North America, north of the

isthmus of Panama. Part. I. Bulletin of the United States Naitonal

Museum 47, I–LX + 1–1240.

Jordan D.S. & Hannibal H. 1923: Fossil sharks and rays of the Pacific

slope of North America. Bulletin of the Southern California

Academy of Sciences 22, 22–63.

Juhász T.J. 2006: Observations on the chondrichthyan remains of the

Danitz-puszta sand pit. Folia Historico Naturalia Musei

Matraensis 30, 9–24 (in Hungarian).

Katona L.T., Kókay J. & Berta T. 2011: Badenian mollusc fauna from

Várpalota (Faller street). Földtani Közlöny 141, 1, 3–22 (In

Hungarian).

Kazár E. & Venczel M. 2003: Kentriodontid remains (Cetacea: Odon-

toceti) from the middle Miocene of Bihor County, Romania.

Folia naturae Bihariae 30, 39–66.

Kazár E., Kordos L. & Szónoky M. 2001: The Danitz-puszta sandpit.

Pannon sand with reworked vertebrate remains. Hungarian Geo

logical Society, Palaeontology and Stratigraphy Section: 4th

Hungarian Palaeontological Convention, Pécsvárad, Abstracts

and Fieldguide, 42–43 (In Hungarian).

Kazár E., Vremír M. & Codrea V. 2004: Dolphin remains (Cetacea:

Odontoceti) from the Middle Miocene of Cluj-Napoca, Roma-

nia. Acta Paleontologica Romaniae 4, 179–189.

Kercsmár Z., Budai T., Csillag G., Selmeczi I. & Sztanó O. 2015:

Surface geology of Hungary. Explanatory notes to the Geologi-

cal map of Hungary (1:500.000). Geological and Geophysical

Institute of Hungary, Budapest, 1–66.

Klunzinger C. B. 1871: Synopsis der Fische des Rothen Meeres.

II. Theil. Verhandlungen der Königlischen ZoologischenBota

nischen Gesellschaft in Wien 21, 441–688.

Koch A. 1903: Tarnócz, in Nógrád county, as new and rich locality for

fossil shark teeth. Földtani Közlöny 33, 22–44 (in Hungarian).

Kocsis L. 2002: Miocene Chondrichthyes remains from the area of

the Mecsek Mountains. Hungarian Geological Society, Palaeon

tology and Stratigraphy Section: 5th Hungarian Palaeonto

logical Convention, Pásztó, Abstracts and Fieldguide, 20–21

(in Hungarian).

Kocsis L. 2007: Central Paratethyan shark fauna (Ipolytarnóc,

Hungary).

Geol. Carpath. 58, 1, 27–40.

Kókay J. 2013: Study of the Middle Miocene (Badenian and Sarma-

tian) formations in the Várpalota Neogene Basin. Földtani

Közlöny 143, 2,145–156.

Kordos L.& Solt P. 1984: Sketch of the marine vertebrate fauna levels

of the Miocene of Hungary. A Magyar Állami Földtani Intézet

Évi Jelentése 1982-ről, Budapest, 347–351 (in Hungarian).

Kováč M., Andreyeva-Grigorovich A., Bajraktarević Z., Brzobohatý R.,

Filipescu S., Fodor L., Harzhauser M., Oszczypko N., Pavelic D.,

Rögl F., Saftić B., Sliva L. & Studencka B. 2007: Badenian

evolution of the Central Parathethys sea: paleogeography,

climate and eustatic sea level changes. Geol. Carpath. 58,

579–606.

Lawley R. 1876: New study on the fossil fishes and other vertebrates

of the Tuscan mountains. Tipografia dell Arte della Stampa,

Firenze, 1–122, pl. 1–5 (in Italian).

Linnaeus C. 1758: Systema Naturae per regna tria naturae, regnum

animale, secundum classes, ordines, genera, species, cum

characteribus differentiis synonymis, locis. Ed. X. 1. Stockholm

(L. Salvius) 1–824.

Leriche M. 1927a: Les Poissons de la Molasse Suisse I. Mémoires de

la Société Paléontologique Suisse 46, 1–56.

Leriche M. 1927b: Les Poissons de la Molasse Suisse II. Mémoires de

la Société Paléontologique Suisse 47, 57–120.

Mikuž V. 1999: The great-teeth shark Carcharocles megalodon (Agassiz)

also from Middle Miocene–Badenian beds above Trbovlje, Slo-

venia [Velikozobi morski pes Carcharocles megalodon (Agas-

siz) tudi v srednjemiocenskih-badenijskih plasteh nad

Trbovljami]. Geologija 42, 141–150 (in Slovenian).

Mikuž V. 2009 : Mackerel shark found also in the Miocene beds in

Dolenjska (Slovenia) [Morski volk najden tudi v miocenskih

plasteh na Dolenjskem]. Folia Biologica et Geologica 50, 2,

91–97 (in Slovenian).

Mikuž V. & Bartol M. 2011: Prva najdba sipine kosti (Sepiidae)

v miocenskih skladih Slovenije. (The first find of cuttlefish shell

(Sepiidae) in Miocene beds of Slovenia.) Folia Biologica et

Geologica 52, 3, 5–22 (in Slovenian).

Mikuž V. & Šoster A. 2013: A Mackerel Shark (Megaselachus mega

lodon) find in Orehovica, Dolenjska, Slovenia. Folia Biologica

et Geologica 54, 1, 109–119.

Mikuž V., Šoster A. & Ulaga S. 2013:Miocene fish teeth from the Plesko

Quarry, Slovenia. Folia Biologica et Geologica 54, 1, 121–133.

Mikuž V., Šoster A., Stare F. & Sukič Prekmurski M. 2015: Megalo-

don teeth from Miocene marlstone at Virštanj, Slovenia [Megal-

odonovi zobje iz miocenskih laporovcev Virštanja]. Folia

Biologica et Geologica 56, 2, 77–107.

Moissette P., Dulai A., Escarguel G., Kázmér M., Müller P. & Saint

Martin J.-P. 2007: Mosaic of environments recorded by bryo-

zoan faunas from the Middle Miocene of Hungary. Palaeogeogr.

Palaeoclimatol. Palaeoecol. 252, 530–556.

593

NEW MIDDLE MIOCENE SELACHIAN ASSEMBLAGE FROM THE CENTRAL PARATETHYS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

Müller J. & Henle F.G.J. 1837: Gattungen der Haifische und Rochen

nach einer von ihm mit Hrn. Henle unternommenen gemein-

schaftlichen Arbeit über die Naturgeschichte der Knorpelfische.

Berichte der Königlichen Preussischen Akademie der Wissen

schaften zu Berlin 111–118.

Müller J. & Henle F.G.J. 1838: On the generic characters of cartilagi-

nous fishes, with descriptions of new genera. Magazine of natu

ral history and journal of zoology, botany, mineralogy, geology

and meteorology (n.s.) 2, 33–37, 88–91.

Müller J. & Henle F.G.J. 1841: Systematische Beschreibung der

Plagiostomen. Veit, Berlin, 1–200.

Müller P. 1984: Decapod Crustacea of the Badenian. Geol. Hung.

Series Palaeontologica 42, 1–317.

Münster G.G. 1846: Ueber die in der Tertiär-Formation des Wiener

Beckens vorkommenden Fisch-Ueberreste, mit Beschreibung

einiger neuen merkwürdigen Arten. Beiträge zur Petrefacten

kunde 7, 1–31.

Noszky J. 1925: Additional informations for the Leitha Limestones of

Hungary. XXII. Annales Musei Nationalis Hungarici 1925 (in

Hungarian).

Papp A. & Cicha I. 1968: Definition der Zeiteinheit M — Badenien.

In: Papp A., Cicha I., Seneš J. & Steininger F. (Eds.): M4 —

Badenien (Moravien, Wielicien, Kosovien). Chronostratigraphie

und Neostratotypen, Miozän der Zentralen Paratethys 6. VEDA,

Bratislava, 47–48.

Pawłowska K. 1960: Fish remains from the Miocene limestone of

Pińczów. Acta Palaeontol. Polon. 5, 4, 421–432 (in Polish).

Péron F. 1807: Voyage de Découvertes aux Terres Australes, exécuté

par ordre de sa majesté l’Empereur et Roi, sur les Corvettes la

Géographe, la Naturaliste et la Goulette la Casuarina, pendant

les années 1800, 1801, 1803 et 1804. Voyage de Découvertes aux

Terres Australes, Paris 1, 1–496.

Piller W., Harzhauser M. & Mandic O. 2007: Miocene Central Para-

tethys stratigraphy — current status and future directions. Stra

tigraphy 4, 151–168.

Philippi R. A. 1846: Tornatella abbreviata, Otodus mitis, Otodus cat

ticus, und Myliobatis testae. Palaeontographica 1, 23–25.

Pimiento C. & Clements C.F. 2014: When Did Carcharocles mega

lodon Become Extinct? A New Analysis of the Fossil Record.

PLoS ONE 9, 10, e111086.

Pimiento C., Ehret D.J., MacFadden B.J. & Hubbell G. 2010: Ancient

Nursery Aera for the Extinct Giant Shark Megalodon from the

Miocene of Panama. PLoS ONE 5, 5, e10552.

Pimiento C., Gonzalez-Barba G., Ehret D.J., Hendy A.J.W., Mac-

Fadden B.J. & Jaramillo C. 2013a: Sharks and rays (Chondrich-

thyes, Elasmobranchii) from the Late Miocene Gatun Formation

of Panama. J. Paleontology 87, 5, 755–774.

Pimiento C., Gonzalez-Barba G., Hendy A.J.W., Jaramillo C., Mac-

Fadden B.J., Montes C., Suarez S.C. & Shippritt M. 2013b:

Early Miocene chondrichthyans from the Culebra Formation,

Panama: A window into marine vertebrate faunas before closure

the Central American Seaway. J. South Amer. Earth Sci. 42,

159–170.

Poey F. 1868: Synopsis of the fishes of Cuba. Catalogue of the fishes

of the Island of Cuba. Repertorio FisicoNatural de la Isla de

Cuba 2, 279–484 (in Spanish).

Pollerspöck J. & Beaury B. 2014: Eine Elasmobranchierfauna (Elas-

mobranchii, Neoselachii) aus der Oberen Meeresmolasse

(Ottnangium, Unteres Miozän) des Heigelsberger Grabens bei

Teisendorf, Oberbayern. Zitteliana A 54, 23–37.

Probst J. 1877: Beiträge zur Kenntniss der fossilen Fische aus der

Molasse von Baltringen. II: Batoidei A. Günther. Jahreshefte des

Vereins für vaterländische Naturkunde in Württemberg 33,

69–103.

Probst J. 1878: Beiträge zur Kenntniss der fossilen Fische aus der

Molasse von Baltringen. Hayfische. Jahreshefte des Vereins für

vaterländische Naturkunde in Württemberg 34, 113–154.

Probst J. 1879: Beiträge zur Kenntniss der fossilen Fische aus der

Molasse von Baltringen. Hayfische. Jahreshefte des Vereins für

vaterländische Naturkunde in Württemberg 35, 127–191.

Purdy W.R., Schneider P.V., Applegate P.S., Mclellan H.J., Meyer

L.R. & Slaughter H.B. 2001: The Neogene sharks, rays, and

bony fishes from Lee Creek Mine, Aurora, North Carolina.

In: Clayton E. Ray & David J. Bohaska (Eds.): Geology and

Paleontology of the Lee Creek Mine, North Carolina, III. Smith

sonian Contributions to Paleobiology 90, 71–202.

Radwański A. 1965: A contribution to the knowledge of Miocene

Elasmobranchii from Pińczow (Poland). Acta Palaeontol. Polon.

10, 2, 267–276.

Rafinesque C.S. 1810: Caratteri di alcuni nuovi generi e nuove specie

di animali e pinate della Sicilia, con varie osservazioni sopera

i medisimi lère partie. (Part 1 involves fishes, i–iv + 3–69

[70 blank], Part 2 with slightly different title, ia–iva + 71–105

[106 blank])

Reinecke T. 2015: Batoids (Rajiformes, Torpediniformes, Myliobati-

formes) from the Sülstorf Beds (Chattian, Late Oligocene) of

Mecklenburg, northeastern Germany: a revision and description

of three new species. Palaeovertebrata 39, 1–32.

Reinecke T. & Radwański A. 2015: Fossil sharks and batoids from the

Korytnica-clays, early Badenian (Langhian, Middle Miocene),

Fore-Carpathian basin, central Poland — a revision and updated

record. Palaeontos 28, 1–32.

Reinecke T., Stapf H. & Raisch M. 2001: Die Selachier und Chimären

des Unteren Meeressandes und Schleichsandes im Mainzer

Becken (Rupelium, Unteres Oligozän). Palaeontos 1, 1–73.

Reinecke T., Balsberger M., Beaury B. & Pollerspöck J. 2014: The

elasmobranch fauna of the Thalberg Beds, early Egerian (Chat-

tian, Oligocene), in the Subalpine Molasse Basin near Siegsdorf,

Bavaria, Germany. Palaeontos 26, 3–129.

Rögl F. 1998: Palaeogeographic considerations for the Mediterranean

and Paratethys seaways (Oligocene to Miocene). Annalen des

Naturhistorischen Museums in Wien 99 (A), 279–310.

Rögl F. & Steininger F.F. 1983: Vom Zerfall der Tethys zu Mediterran

und Paratethys. Die neogene Paläogeographie und Palinspastik

des zirkum-mediterranen Raumes. Annalen des Natur

historischen Museums in Wien 84 (A), 135–163.

Rögl F. & Brandstätter F. 1993: The foraminifera genus Amphistegina

in the Korytnica Clays (Holy Cross Mts, Central Poland) and its

significance in the Miocene of the Paratethys. Acta Geol. Polon.

43, 1/2, 121–146.

Rögl F., Ćorić S., Harzhauser M., Jimenez-Moreno G., Kroh A.,

Schultz O., Wessely G. & Zorn I. 2008: The Middle Miocene

Badenian stratotype at Baden-Sooss (Lower Austria). Geol.

Carpath. 59, 5, 367–374.

Rüppell W.P.E.S.E. 1837: Fische des rothen Meeres. Frankfurtam

Main. 53–80, Pls. 15–21.

Schmid H.-P., Harzhauser M. & Kroh A. 2001: Hypoxic Events on

a Middle Miocene Carbonate Platform of the Central Paratethys

(Austria, Badenian, 14 Ma). 1–50, 8 pls., 8 figs.

Schultz O. 1971: Die Selachier-Fauna (Pisces, Elasmobranchii) des

Wiener Beckens und seiner Randgebiete im Badenien (Miozän).

Annalen des Naturhistorischen Museums in Wien 75, 311–341.

Schultz O. 1977: Elasmobranch and teleost fish remains from the

Korytnica Clays (Middle Miocene; Holy Cross Mountains,

Poland). Acta Geol. Polon. 27, 2, 201–209.

Schultz O. 1979: Supplementary notes on elasmobranch and teleost

fish remains from the Korytnica Clays (Middle Miocene; Holy

Cross Mountains, Central Poland). Acta Geol. Polon. 29, 3,

287–293.

Schultz O. 2003: The Middle Miocene Fish Fauna (excl. otolithes)

from Mühlbach am Manhartsberg and Grund near Hollabrunn,

Lower Austria. Annalen des Naturhistorischen Museums in

594

SZABÓ and KOCSIS

GEOLOGICA CARPATHICA

, 2016, 67, 6, 573 – 594

Wien 104 (A), 185–193.

Schultz O. 2013: Pisces. In: Piller W. (Hg.) Catalogus Fossilium Aus-

triae, Bd. 3; 576 pp. Verlag der Österreichischen Akademie der

Wissenschaften, Wien, (ISBN 978-3-7001-7238-3).

Schultz O., Brzobohatý R. & Kroupa O. 2010: Fish teeth from the

Middle Miocene of Kienberg at Mikulov, Czech Republic,

Vienna Basin. Annalen des Naturhistorischen Museums in Wien

112 (A), 489–506.

Sebe K., Csillag G., Dulai A., Gasparik M., Magyar I., Selmeczi I.,

Márton Sz., Sztanó O. & Szuromi-Korecz A. 2015: Neogene

stratigraphy in the Mecsek Region. 6

Workshop on the Neogene

of Central and South-Eastern Europe. Hungarian Geological

Society. In: An RCMNS Interim Colloquium. Programme,

Abstracts, Field Trip Guidebook. 31 May–3 June 2015, Orfű,

Hungary, 102–124.

Selmeczi I. 1996: The Pusztamiske Formation. In: Gyalog L. (Ed.):

Explanatory notes of geological maps and short descriptions for

the stratigraphical units. A Magyar Állami Földtani Intézet

Alkalmi Kiadványa 187, Budapest, 81 (in Hungarian).

Selmeczi I. 2003: Prepannonian Miocene formations in the

south-western area of the Transdanubian Mountains (Devecs-

er-Nyirád Basin, Tapolca Basin, northern part of the Keszthely

Mountains). PhD thesis, Pécsi Tudományegyetem Természet

tudományi Kar, Földrajzi Intézet, Földtudományok Doktori

Iskola. Kézirat, 1–130 (in Hungarian).

Selmeczi I., Bohnné Havas M., Szegő É. & Lelkes Gy. 2002: The

Lower Badenian of the Devecser-Nyirád Basin. Investigations

on Macro-, Microfauna and Microfacies. 5. Őslénytani Vándorg-

yűlés. Programme, Abstracts, Field Trip Guidebook. 3 May–4

May 2002, Pásztó, Hungary, 28–29 (in Hungarian).

Solt P. 1987: Following Ferenc Légányi in Mátraszőlős, a locality for

Procarcharodon. Folia HistoricoNaturalia Musei Matraensis

12, 15–18 (in Hungarian).

Solt P. 1991: Marine fish remains from the Upper Miocene of

Hasznos. A Magyar Állami Földtani Intézet Évi Jelentése

1989-ről, Budapest, 473–478 (in Hungarian).

Šoster A. & Mikuž V. 2013: Fish remains from Miocene beds of

Višnja vas near Vojnik, Slovenia [Ostanki rib iz miocenskih

plasti Višnje vasi blizu Vojnika]. Geologija 56, 1, 73–86 (in

Slovenian).

Taniuchi T. 1970: Variation in the teeth of sand shark, Odontaspis

taurus (Rafinesque) taken from the East China Sea. Japan. J.

Ichthyology 17, 1, 34–44.

Vialle N., Adnet S. & Cappetta H. 2011: A new shark and ray fauna

from the Middle Miocene of Mazan, Vaucluse (southern France)

and its importance in interpreting the paleoenvironment of

marine deposits in the southern Rhodanian Basin. Swiss J.

Palaeontology 130, 241–258.

Vitális I. 1942: Dentition of extant Notidanus and fossil Notidanus

primigenius Ag., in regard to the Miocene Notidanus teeth from

Mátraszöllős (Hungary). Geol. Hungarica, Series Palaeontolog

ica 18, 1–38 (in Hungarian).

Vrsaljko D., Japundžić S., Kovačić M., Grganić-Vrdoljak Z. & Pleše

P. 2010: Vranić: The most important finding place of fossil

whales in Northern Croatia. 4

Croatian geological congress,

Zagreb, 118.

Weiler W. 1933: Zwei oligozäne Fischfaunen aus dem Königreich

Ungarn. Geol. Hungarica. Series Palaeontologica 11, 1–54.

Weiler W. 1938: Neue Untersuchungen an mitteloligozänen Fischen

Ungarns. Geol. Hungarica. Series Palaeontologica 15, 1–31.

Whitley G.P. 1940: The fishes of Australia. Part 1. The sharks, rays,

devil fishes and other primitive fishes of Australia and New Zea-

land. Royal Zoological Society of New South Wales, Sydney,

1–230.

Wysocka A., Radwañski A. & Górka M. 2012: Mykolaiv Sands in

Opole Minor and beyond: sedimentary features and biotic con-

tent of Middle Miocene (Badenian) sand shoals of Western

Ukraine. Geol. Quarterly 56, 3, 475–492.

Appendix

Placoid scales from the Pusztamiske Formation: MFGI V 2016.10.1.
Selachimorpha indet. teeth from the Pusztamiske Formation: MFGI V 2016.1.1., MFGI V 2016.15.1., MFGI V 2016.21.1.
Acanthurus sp. teeth from the Pusztamiske Formation: MFGI V 2016.11.1.
Dentex sp. teeth from the Pusztamiske Formation: MFGI V 2016.16.1.
?Diplodus sp. teeth from the Pusztamiske Formation: MFGI V 2016.4.1.
Pagrus cinctus teeth from the Pusztamiske Fm.: MFGI V 2014.88.25.1-25., MFGI V 2016.13.1., MFGI V 2016.17.1., MFGI V 2016.18.1.,

MFGI V 2016.19.1.
Indeterminate fish vertebra from the Leitha Limestone Fm.: MFGI V 2014.87.1.
Indeterminate Sparidae teeth from the Pusztamiske Formation: MFGI V 2016.7.1.
Indeterminate Tetraodontidae teeth from the Pusztamiske Formation: MFGI V 2016.3.1.
?Trichiurus sp. teeth from the Pusztamiske Formation: MFGI V 2014.102.1., MFGI V 2016.9.1.
Odontoceti indet. tooth from the Leitha Limestone Formation: MFGI V 2014.109.1.
?Sirenia vertebra from the Leitha Limestone Formation: MFGI V 2016.45.1.

Indeterminate bone fragments from the Pusztamiske Formation: MFGI V 2016.44.1.