GeolCarp_Vol70_No2_135

GEOLOGICA CARPATHICA

, APRIL 2019, 70, 2, 135–152

doi: 10.2478/geoca-2019-0008

www.geologicacarpathica.com

Sedimentological characteristics and paleoenvironmental

implication of Triassic vertebrate localities in Villány

(Villány Hills, Southern Hungary)

GÁBOR BOTFALVAI

1, 2,



, ORSOLYA GYŐRI

, EMÍLIA POZSGAI

, IZABELLA M. FARKAS

TAMÁS SÁGI

6, 7

, MÁRTON SZABÓ

1, 2

and ATTILA ŐSI

Department of Paleontology and Geology, Hungarian Natural History Museum, Baross Street 13, H-1088, Budapest, Hungary;



botfalvai.gabor@gmail.com

Department of Paleontology, Eötvös Loránd University, Pázmány Péter sétány 1/C, H-1117, Budapest, Hungary;

szabo.marton.pisces@gmail.com , hungaros@gmail.com

MTA–ELTE Geological, Geophysical and Space Science Research Group, Pázmány Péter sétány 1/c, H-1117 Budapest, Hungary;

gyori.orsi@gmail.com

Soós Ernő Water Technology Research and Development Center, University of Pannonia, Zrínyi Miklós Street 18, H-8800 Nagykanizsa,

Hungary; emily.pozsgai@gmail.com

Laboratories MOL, MOL Plc., Szent István Street 14, H-1039, Budapest, Hungary; izabella.farkas@gmail.com

Department of Petrology and Geochemistry, Eötvös Loránd University, Pázmány Péter sétány 1/C, H-1117, Budapest, Hungary;

sagi.tamas@ttk.elte.hu

MTA-ELTE Volcanology Research Group, Pázmány Péter sétány 1/C, H-1117, Budapest, Hungary

(Manuscript received September 26, 2018; accepted in revised form February 12, 2019)

Abstract: There are two Triassic vertebrate sites in Villány Hills (Southern Hungary), where productive and continuous

excavations have been carried out in the last six years resulting in a rich and diversified assemblage of shallow marine to

coastal animals. The studied formations belong to the Villány–Bihor Unit of the Tisza Megaunit, which was located at

the passive margin of the European Plate during the Triassic. The relatively diverse vertebrate assemblage was collected

from a Road-cut on Templom Hill and a newly discovered site at a construction zone located on the Somssich Hill. Four

main lithofacies were identified and interpreted in the newly discovered Construction vertebrate site consisting of

dolomite (deposited in a shallow, restricted lagoon environment), dolomarl (shallow marine sediments with enhanced

terrigenous input), reddish silty claystone (paleosol) and sandstone (terrigenous provenance) indicating that

the sediments of the Construction vertebrate site were formed in a subtidal to peritidal zone of the inner ramp

environment, where the main controlling factor of the alternating sedimentation was the climate change. However,

the recurring paleosol formation in the middle part of the section also indicates a rapid sea-level fall when the marine

sediments were repeatedly exposed to subaerial conditions. In the Road-cut site the siliciclastic sediments of

the Mészhegy Sandstone Formation are exposed, representing a nearshore, shallow marine environment characterized by

high siliciclastic input from the mainland.

Keywords: cyclic carbonate–siliciclastic deposits, dolomite, inner ramp, peritidal zone, vertebrates, Tisza Megaunit.

Introduction

The vertebrate remains from the Mesozoic of Hungary are

rela tively rare, and aside from a few isolated fossils only three

localities are known where productive and continuous excava-

tions have been conducted. Two of them provide vertebrate

fossils from Upper Cretaceous strata (Ajka and Csehbánya

Formations), deposited in freshwater and terrestrial environ-

ments. Fishes, amphibians, turtles, crocodiles and dinosaurs

were found in these sites of the Bakony Mountains (Ősi et al.

2012). The third vertebrate locality is situated in Villány,

Villány Hills (South Hungary) and includes two outcrops of

the Middle to Upper Triassic formations (Fig. 1). Field work in

these sites revealed rich and diverse assemblage of coastal to

shallow marine animals including scales and teeth of fishes,

cranial and postcranial elements of sauropterygians (notho-

saurs and placodonts), and vertebrae of Tanystropheus (Ősi et

al. 2013; Segesdi et al. 2017; Table 1). These Triassic fossils

and their embedding successions are of great importance,

since according to the relevant paleoreconstructions the Tisza

Megaunit, and within it the Villány area, was located at

the Northern Neotethys margin, on the shelf of the European

Plate southwards to the Bohemian Massif (Csontos & Vörös

2004; Haas & Péró 2004; Pozsgai et al. 2017).

Middle to Late Triassic marine to coastal vertebrate fossil

sites are well known from the Central European Basin and

Alpine successions representing the Western European realm

of the Tethys (e.g., Pinna 1990; Rieppel 2000; Schoch 2015;

Renesto & Dalla Vecchia 2018). However, much less is known

about the vertebrate faunal composition of the eastern regions

of the Northern Tethyan coast. With its abundant and relatively

diverse fauna (chondrichthyans, osteichthyans, nothosaurs,

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placodonts, archosauromorphs) is thus of great importance

since it extends our geographic and faunistic knowledge on

the shallow marine to seashore vertebrates of the Northern

Tethyan coast.

Besides the paleogeographic uniqueness, the vertebrate

fossils of the Villány sites help to gain a better understanding

of the Late Ladinian to Early Carnian evolutionary events of

sauropterygian and prolacertiform archosauromorphs, since

the record of these groups from this period of the Triassic is

much less known than from earlier periods. Thus, a detailed

evaluation of the sedimentological features and reconstruction

of the depositional environment of these recently discovered

bone-yielding beds are crucial in the paleoenvironmental

reconstruction, which is indispensable for the upcoming

paleontological research. Furthermore, besides the paleonto-

logical significances, the explored successions provide addi-

tional information about these sedimentological and

environmental processes. The bone-bearing successions of

the Villány Hills were deposited in an inner ramp environment

(Rálisch-Felgenhauer & Török 1993; Török 2000, and see

below) representing the transition zone between the upper

shoreface and fair-weather wave base (Burchette & Wright

1992). Mixed siliciclastic–carbonate deposits frequently

accommodate in such environments consisting of both

extrabasinal (e.g., terrigenous siliciclastic) and

intrabasinal (autochthonous carbonate) compo-

nents (Morsilli et al. 2012; Caracciolo et al. 2013;

Chiarella et al. 2017). The alternation of litho-

facies and/or the sediment mixing can be inter-

preted as the result of short-term sea-level or

short-term climate changes. Thus, the investiga-

tion of this type of successions is useful to under-

stand short-term sea-level and climatic changes

and processes (e.g., Brachert et al. 2003; Zeller et

al. 2015; Blanchard et al. 2016; Reis & Suss 2016).

The sedimentological characteristics of one of

the vertebrate sites (Road-cut section) is well docu-

mented by several authors (Rálisch-Felgenhauer

1985; Vörös 2009, 2010), but the interpretation

of the depositional environment of this succes-

sion remained controversial. We performed here

new observations and paleontological data, which

may help to determine the depositional environ-

ment of this sediment accumulation. The other

bone-bearing excavation site (Construction site)

is less studied, because detailed paleontological

and sedimentological investigations of this suc-

cession has started only in 2012, when the section

was industrially excavated, and one of the authors

(E. Pozsgai) found a few bones and teeth in this

section and the area was recognized as a potential

Triassic vertebrate locality.

The aim of the present study is to give

an insight into the sedimentological history of

the bone-yielding successions. Based on the detai-

led description and interpretation of the identified

facies associations, the sedimentological and geological

significances of the shallow marine setting are discussed, and

the depositional paleoenvironments of the vertebrate sites are

identified.

Location and regional geology

The Villány Hills are situated in the southwestern part of

the Pannonian Basin in Hungary (Fig. 1A). The studied suc-

cessions are located 200–300 m northwest to the city of

Villány (Fig. 1B). The Villány Hills belong to the Villány–

Bihor Unit of the Tisza Megaunit, which formed a segment of

the passive Neotethys margin of the European Plate during

the Triassic (Csontos & Vörös 2004; Haas & Péró 2004; Feist-

Burkhardt et al. 2008; Fig. 2A).

The Lower Triassic sequence of the Villány–Bihor Unit is

predominantly characterized by clastic sediments (Bunt sand-

stein facies), which is overlain by Middle Triassic evaporitic

carbonate and shallow marine carbonate deposits (Röt and

Muschelkalk facies). These sequences show a close genetic

affinity with the Germanic-type Triassic sediments (Török

1997), while the Upper Triassic succession shows closer affi-

nities to the Carpathian Keuper facies (Bleahu et al. 1994).

Fig. 1. Map of the locality (A) and location of the two vertebrate sites in Villány (B).

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The Middle Triassic sequence was deposited on a homo-

clinal carbonate ramp of relatively uniform subsidence rate

(Török 1997, 2000). The geotectonic setting and the related

slow subsidence suggest that the sediment deposition may

have been primarily controlled by the eustatic sea-level

changes (Török 2000). In accordance with sea-level fluctua-

tions, three Middle Triassic deepening-upward and shallowing-

upward cycles were identified (Török 2000; Götz et al. 2003;

Götz & Török 2008).

The first cycle corresponds to the ramp initialisation and

the onset of carbonate sedimentation, coinciding with the Early

Anisian global sea-level rise (Török 2000; Götz & Török

2008). The beginning of the transgressive phase of that cycle

is represented by evaporitic sabkha sediments: dolomite, dolo-

marl and anhydrite (Hetvehely Anhydrite Formation; Fig. 2B).

The following regressive phase is characterized by thick-

bedded massive dolomite (Rókahegy Dolomite Formation).

The second depositional cycle took place during the Middle

to Late Anisian period. It shows striking similarities with

the first Muschelkalk cycle of the Central European Basin

(Török 2000; Götz & Török 2008). The dark grey, nodular,

fossiliferous clayey limestone (lower part of the Zuhánya

Limestone Formation; Fig. 2B) represents the transgressive

phase (Götz et al. 2003), whereas the dolomitic limestones and

the entirely dolomitized inner ramp carbonates (upper part of

the Zuhánya Formation) indicate the regressive phase of this

cycle (Haas 2001). The latest Anisian to Early Ladinian inter-

val is poorly documented in the Villány Hills. However, from

the early part of the Ladinian the general shallowing of

the basin can be assumed that is represented by yellowish grey

dolomite with dolomitic marl intercalations (Csukma Dolomite

Formation; Haas 2001).

The lower part of Csukma Dolomite Formation, known

from outcrops and boreholes in the middle part of the Villány

Hills, consists of grey, thick-bedded, locally laminated dolo-

mites with rare relicts of ooids, micro-tepee and fenestral

structures (Török 2000). Based on sedimentological characte-

ristics (e.g., presence of fenestral laminae, extensive dolomiti-

zation, tepees and exposure surfaces), the subtidal to peritidal

zone of the inner ramp was interpreted as the depositional

environment of the Csukma Dolomite (Rálisch-Felgenhauer

& Török 1993; Török 2000; Fig. 2B).

The uppermost part of the Csukma Dolomite, which is made

up by the alternation of yellowish grey dolomite and dolomarl

layers, was defined as the Templomhegy Dolomite Member of

this formation (Fig. 3).

Remains of a relatively diverse marine fish and reptile fauna

(Ősi et al. 2013) were recently encountered in this member

(Construction site; see below). Unfortunately, only a few

badly preserved casts of hard-shelled invertebrate fossils are

known from the dolomite and dolomarl beds of the Templom-

hegy Member, which cannot be used for more detailed paleo-

environmental reconstruction. However, a protected inner ramp

lagoon and connected tidal flat were interpreted as the deposi-

tional environment of the Templomhegy Member (Török 1998,

2000; Haas 2001; Bérczi-Makk et al. 2004; Ősi et al. 2013).

The age of the Templomhegy Dolomite Member of

the Csukma Dolomite Formation (including the bone-bearing

horizon of Construction site) is problematic since index fossils

of ammonites and conodonts are absent in this shallow marine

sequence. However, the Templomhegy Dolomite Member

probably belongs to the Ladinian stage based on its strati-

graphic position, because it is situated between the Late

Anisian Zuhánya Limestone Formation and the Carnian

Mészhegy Sandstone Formation (Rálisch-Felgenhauer &

Török 1993; Török 1998; Haas 2001; Fig. 2B). Furthermore,

the new vertebrate material (especially the remains of

Nothosaurus sp.), discovered from the Templomhegy

Dolomite Member, also suggests a late Middle Triassic

(Ladinian) age for this deposition (Ősi et al. 2013).

The relatively thick shallow marine Middle Triassic carbo-

nate succession is overlain by a thin formation that is made up

Vertebrate assemblage

Construction site

Road-cut section

Lifestyle

Taxon

Ladinian

(Templomhegy

Member)

Carnian

(Mészhegy

Sandstone

Formation

Ladinian

(Templomhegy

Member)

Carnian

(Mészhegy

Sandstone

Formation

Environment

References

Fishes

Hybodus sp.

marine, brackish, freshwater

Cuny 2012;

Klug et al. 2010

Palaeobates angustissimus

marine

Böttcher 2015;

Diedrich 2009

Polyacrodus sp.

marine

Diedrich 2009

Lissodus sp.

marine, freshwater

Cappetta 2012

Gyrolepis sp.

marine

Lakin et al. 2016;

Whiteside et al. 2016

Severnichthys acuminatus

marine

Actinopterygii indet

marine, brackish, freshwater

Nelson 2006

Reptiles

Nothosaurus sp. 1

marine

Rieppel 2000

Nothosaurus sp. 2

marine

Rieppel 2000

cf. Cyamodus sp.

marine

Rieppel 2000

Tanystropheus sp.

marine to coastal

Renesto 2005

?Archosauriformes indet.

coastal to terrestrial?

Table 1: Synthetic faunal list of the Triassic marine vertebrate fauna from the Villány Hills (based on Ősi et al. 2013; Segesdi et al. 2017 and

Electronic supplement II).

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mostly of siliciclastic rocks. It was defined as the Mészhegy

Sandstone Formation and was assigned to the Upper Triassic

(Fig. 2B). The Mészhegy Sandstone Formation (Fig. 2B) is

composed of conglomerate, siltstone, sandstone, cellural dolo-

mitic limestone and marl (Rálisch-Felgenhauer & Török 1993;

Vörös 2009, 2010). The sedimentology of the Mészhegy

Formation exposed in the Road-cut section is well docu-

mented, however, the depositional environment of this succes-

sion has been interpreted in different ways. Some authors

argued that it is a shallow marine or littoral deposit (Rálisch-

Felgenhauer 1985; Török 1998), whereas

Vörös (2010) claimed fluvio-lacustrine

origin. The palynological investigation

indicates a Carnian age for the lower part

of the formation (Ősi et al. 2013 and see

below), but the age of the upper part is

still unknown. Vörös (2009) suggests that

this formation is composed of three sedi-

mentary parasequences, one definitely

Carnian, and two others, possibly Norian

and Rhaetian in age. However, this

hypothesis is not supported by paleon-

tological data due to the lack of age- con-

straining flora and fauna in the upper beds.

The discovered fish remains show a uni-

form taxonomical distribution through

the exposed section (see below), probably

indicating a shorter depositional time

for this formation (see below). The thin-

ness (up to 20 m) of the formation and

the seemingly continuous succession, as

well as the available paleontological data

rather suggest a Carnian age for the whole

formation (Ősi et al. 2013).

The Mészhegy Sandstone Formation of

the Road-cut site on Templom Hill is cove-

red by the Pliensbachian Somssichhegy

Limestone Formation. The lowermost

yello wish sandstone strata of the Pliens-

bachian Somssichhegy Limestone Forma-

tion unconformly overlies the Triassic

strata of the Mészhegy Formation (Vörös

1972, 2009, 2010, 2012).

Methods of investigations

The sedimentology of two Triassic

vertebrate sites has been investigated in

detail at Villány Hills (Fig 1B). Sedi-

mentary rocks were analysed in the field

and by hand specimens (1 kg from every

layers), collected from both sections.

The detailed field investigations included

the determination of grain size, colour,

bedding morphology along with record-

ing of paleontological data.

For microvertebrate faunal investiga-

tions, samples were taken from three pro-

ductive beds (L3–4–5) of the Mészhegy

Sandstone Formation at the Road-cut site,

Fig. 2. A — Paleogeographical map of the Tisza Megaunit in the Late Triassic (compiled by

Pozsgai et al. 2017). B — Triassic formations of the Tisza Megaunit (after Török 1998; Haas

2001).

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and four productive layers (14

, 18

, 20

and 22

layers) of

the Templomhegy Dolomite Member at the Construction site

(see below). The samples were screen-washed, by using tap

water and 5 % of acetic acid. The dried residue was sorted

under magnification, using a binocular microscope. The pic-

tures from the material were carried out with a Hitachi

S-2600N scanning electron microscope at the Department of

Botany, Hungarian Natural History Museum (NHMUS).

Detailed petrographic investigations were carried out on 22

thin sections, prepared at the Department of Physical and

Applied Geology, Eötvös Loránd University (ELU). A solu-

tion of alizarin red-S and potassium ferricyanide was used to

identify the carbonate phases in the samples (Dickson 1966).

The X-ray powder diffraction (XRD) measurements were

made using a Bruker D8 Advance powder diffractometer, with

parallel beam, 2θ – θ geometry equipped with LynxEye

detector. Before measurements the few grams, needed for

X-ray analysis were grounded as fine powder with a mean

diameter size between 1–5 µm using micronizing mill (Retsch

MM 400 type) for 3+2 minutes. The final grain size was

obtained using agate mortar and pestle. The measuring para-

meters were: step-scanning at 0.01 ° 2θ intervals, counting

time of 17.7 s (0.1). CuKα radiation at 40 kV and 40 mA was

used. The measurement range was 2–70 ° 2θ.

The identification of minerals was achieved using

the Diffrac EVA software by comparison of the X-ray diffrac-

tion pattern from the sample with the International Centre for

Diffraction Data PDF-2, release 2009 database. The (semi)-

quantitative data were obtained using TOPAS

software provi-

ding us a strandardless quantitative analysis (based on Rietveld

method).

Petrographic descriptions of the bed 10 were carried out

with a Nikon YS2-T polarizing microscope and an AMRAY

1830 I/T6 Scanning Electron Microscope (equipped with

a PU9800 EDX spectometer) at the Department of Petrology

and Geochemistry, ELU. Accelerating voltage of the SEM

instrument was 20 kV with and average beam current of 1 nA.

Back Scattered Electron (BSE) images were created from

characteristic areas of the thin sections, mineral phases were

identified by their EDS spectra.

Fig. 3. Schematic stratigraphic section of the Construction site (A). The lower part of the observed section is made up by dominantly micritic

dolomite beds with subordinate argillaceous deposits (B–C). There are main bone-bearing horizons of the Construction site (D). The bone-bearing

succession is covered by 3 meter thick massive dolomite succession with thin dolomarl intercalations (E). The scale bars represent 1 m.

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Sedimentological characteristics and facies

interpretation of vertebrate sites in Villány Hills

The two studied vertebrate sites in the eastern part of Villány

Hills expose the upper part of the Triassic succession.

• The Construction site is located on the Somssich Hill

(45°52’28.9” N, 18°26’46.2” E; Fig.1B), where productive

and continuous excavations have been carried out in the last

six years resulting in thousands of macro- and microverte-

brate remains of fishes and reptiles in several beds of

the Templomhegy Dolomite (Table 1), while the Mészhegy

Formation was proved to be unfossiliferous.

• The Road-cut section on the Templom Hill (45°52’31.6” N,

18°26’55.5” E) exposes the Mészhegy Sandstone where

a relatively diverse microvertebrate assemblage was disco-

vered (Fig. 1B and Table 1).

The aim of the following subchapters is to present a detailed

sedimentological description and interpretation of the Con-

struction site. In this article, after discussing the interpreted

depositional environment of this site, we provide an overview

of the previous sedimentological works carried out on the sili-

ciclastic succession of the Road-cut section and summarize

the most recent interpretation of its depositional environment

based on the newly discovered vertebrate microfossil

material.

Construction site on the Somssich Hill

The Construction site is located southwest of the Villány

railway station next to Arany János Street on the Somssich

Hill (Fig. 1B). The exposed succession is about 20 meter long

and 3–4 meter high, where around 17 meter thick succession

of the upper part of the Templomhegy Dolomite and a very

thin 3 meter thick sequence of the Mészhegy Sandstone crop

out. The dominant formation of this site, the Templomhegy

Member made up of southward-dipping (around 170/40 to

190/45) greyish dolomite and yellowish dolomarl beds with

dolomitic claystone interlayers (Fig. 3). Here only the bone-

bearing section of the Templomhegy Dolomite is discussed.

Sedimentological description and interpretation of

the Construction site

Composition of the Templomhegy Member exposed in

the Construction site is highly variable. It consists dominantly

of dolomite, quartz and clay minerals, and subordinate amount

of calcite (see Electronic supplement I). The grain size of

siliciclastic components ranges from clay to sand size. Four

lithofacies types were recognized, using a classification based

on colour, grain size, bedding, fossil content and sedimentary

structures (Figs. 3–9).

Dolomite lithofacies: This lithofacies is characterised by

white to light pink dolomite beds (Fig. 4A–C). Mineralogically,

this rock contains 80–96 % dolomite, while the clay minerals

and quartz content is consistently low (<10 %). Although

the carbonate rock is composed predominantly of dolomite,

the youngest, ~20 cm thick, layer of the succession (bed 29)

contains 20 % calcite (Fig. 5). The feldspar content is very low

throughout the investigated succession (<1 %). This litho-

facies is common in the lowermost and the uppermost part of

the site (Fig. 3), while the middle part of the section includes

thinner dolomite layers occurring between dolomarl beds

(Fig. 3D). The thickness of the beds decreases upwards in

the section (30–40 cm in the lower and 10–15 cm in the upper

part of the site).

The dolomite is usually aphanocrystalline and homoge-

neous in thin section (Fig. 6A), pointing to micritic precursor.

Some beds show mottled fabric, where the crystal size is

the same, but there are brownish patches, probably richer in

micrometer-sized solid inclusions (Fig. 6B). The shape of such

mottles is irregular. One sample is composed of aphanitic

“sphaerules” of 200 to 300 micrometers, surrounded by very

finely crystalline dolomite (Fig. 6C). The youngest dolomite

layer of the exposed section (bed 29) contains a fine fracture

system, along which the rock is calcitized/dedolomitized.

The calcite contains few micrometer-sized remnants of dolo-

mite and is intergrown with pyrite.

Vertebrate fossils are usually rare in this lithofacies, a few

dozen bones and teeth of sauropterygians (Ősi et al. 2013)

were discovered from it.

The dolomicrite fabric suggests a lime mud precursor sedi-

ment. Samples with moderate fabric preservation suggest

an originally ooidal carbonate sediment (Fig. 6C).

Therefore, this lithofacies can be interpreted as a carbonate

mud deposited in a low-energy, shallow, restricted lagoonal

environment (similar to Stockman et al. 1967; Brooks et al.

2003a, b; Blanchard et al. 2016), with episodic ooidal sedi-

ment transport. The probably reflux-related dolomitization of

these sediments may have taken place in near surface setting.

Dolomarl lithofacies: A dominant lithofacies of this verte-

brate site is yellowish to grey dolomarl with pale reddish

coloured mottles (Fig. 4A, B and D). The thickness of such

beds can vary from 10 cm to 50 cm. The clay content of

the dolomarl beds varies considerably (Fig. 7).

Mineralogically, this lithofacies consists dominantly of

dolomite (42–80 %), although the percentages of clay mine-

rals may reach 30–20 % (Fig. 7). The dolomarl beds of

the middle part of the section (beds 8–16) are characterized by

a higher quartz content (20–30 %), while this mineral is subor-

dinate (<5 %) in the other dolomarl horizons (see Electronic

supplement I). The feldspar content is predominantly low and

never exceeds 2 %. The more argillaceous dolomarl beds

(e.g., bed 14) show a complex system of red clay-filled cracks

(see below).

In thin section micrometer-sized dolomite crystals and clay

particles are visible. Slight changes in crystal size and mottled

fabric were commonly observed (Fig. 6D). There are brown

pressure solution seams.

The vertebrate fossils are more common in those dolomarl

beds which are characterized by lower carbonate and higher

siliciclastic content. Bed 14 is particularly important in terms

of chondrichtyan and osteichthyan fish remains; thousands of

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fish and sauropsid reptilian teeth and scales were collected

from here (see Electronic supplement II). Bed 14 shows

slightly different features than the other dolomarl layers at this

site, because its top is pedogenetically modified (Fig. 8D–F

and see below).

This lithofacies, comprising of mixed carbonate and fine

siliciclastic (clay and silt) components, indicates a periodi-

cally changing terrigenous input, which was probably con-

trolled by climatic and/or short-term sea-level changes (see

below). The dolomitization, the lack of grainstone textures

and the abundance of mud-dominated lithologies, as well as

the presence of pedogenically modified dolomarl beds (see

below) suggest that this facies was formed in a low energy,

restricted shallow marine environment. The pale reddish stain

in the argillaceous marl sediments probably indicates a more

intense oxidation due to the influence of meteoric water during

periodic subaerial exposure.

Reddish silty claystone: The reddish silty claystone with

pale yellow mottles represents the third lithofacies in

the obser ved section (Fig. 8). There are three claystone hori-

zons (top of bed 14, beds 21 and 27) in the section; their thick-

ness does not exceed 10 cm (Fig. 3A). These rocks are

characterized by a higher quartz (~20 %) and a lower dolomite

content (~20 %) than that of the dolomarl lithofacies (see

Electronic supplement I). Dispersed pale yellow mottles are

relatively common in this facies. Tiny cylindrical, poorly

preserved vertical root casts and weakly defined nodules can

also be detected (Fig. 8). Yellowish carbonate and reddish

brown clay rich patches show a mottled appearance in thin

section (Fig. 6E). Red colour of the clay rich part is due to

fine-crystalline hematite. Two slightly different types of this

lithofacies can be distinguished:

• Dark red, slightly calcareous, homogenous layer with tiny

irregular yellowish root traces. It usually forms a distinct

layer between two dolomarl beds (bed 21). No vertebrate

fossils has been found in it (Fig. 8A, B);

• Silty claystone of higher dolomite content, rich in carbonate

nodules. It is developing through a continuous transition

from the underlying dolomarl beds (top of bed 14).

Fragmented carbonate crusts showing poorly definable tepee

structure and laminated micritic horizons are present in

the uppermost part of this thin claystone layer (Fig. 8D–F).

The reddish colour and the vertical root casts indicate that

this claystone was better drained than the other argillaceous

carbonates in the section. Based on its mineral content (domi-

nated by non-carbonate minerals), macrofeatures (root moulds,

fine-grained layers with Liesegang-bands and mottles) and

colour, this horizon is interpreted as calcic paleosol (similar to

Klappa 1980a, b; Wright 1994; Kraus 1999). The red colour

implies the abundance of ferric oxides indicating oxidizing

conditions and well-drained environment during pedogenesis

(Wright 1994; Kraus 1999; Zand-Moghadam et al. 2014;

Fig. 4. The bone-bearing succession of the Construction site is made up by the alternation of yellowish grey dolomite and dolomarl layers

(A, B). Dolomite (C) and dolomarl beds (D) with bones at the Construction site.

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Huggett et al. 2016). The laminated claystone horizons in

the uppermost part of bed 14 can be interpreted as dolocretes

(Fig. 8F) and may have been formed by microbial mediation

(similar to Wright 1994: fig.4). The presence of tepee structure

(Fig. 8E), dolocretes, as well as the vertical root moulds indi-

cate that these silty claystone sediments have been subaerially

exposed, during which they were modified by pedogenetic

processes. The peritidal environment is regularly exposed to

subaerial conditions during low sea-level stages resulting in

the development of thin paleosol horizons on top of the car-

bonate sediments (Ginsburg 1975; Wright 1994; Blanchard et

al. 2016; Huggett et al. 2016).

Sandy claystone: This is a single greenish, greyish, 80 to

100 cm thick, moderately to weakly calcareous sandy claystone

bed (bed 10) with subordinate clay intercalations (Figs. 3A

and 9). Quartz is the most abundant component (about 50 %),

while the clay mineral content shows an upward increasing

trend reaching about 50 % at the top of the layer. The rock is

characterized by a relatively high feldspar content (about

12 %), but this quantity suddenly decreases in the upper part

of the layer, where the clay content increases (Fig. 9B).

The microscopic texture is characterized by a very fine-

grained clay-rich matrix in which coarser grained, quartz-rich,

sometimes lenticular domains occur (Fig. 9C, D).

Based on the SEM analysis, the fine grained component

consists of illite (90–95 %), quartz (3–5 %), K-feldspar (<1 %),

calcite (1–2 %) and accessory minerals: glauconite, biotite,

magnetite, zircon (<1 % respectively). Quartz grains are

mostly 5–10 μm in size, biggest ones can reach 25–30 μm.

K-feldspar is usually much smaller, maximal grain size is

~15–20 μm. The laminae of the claystone is densely pene-

trated by slightly undulating cracks, which are more or less

parallel with the longer axes of the quartz rich lenses.

The transition towards the coarser grained domains and

towards the adjacent carbonate layers is continuous.

Based on the SEM and polarising microscopy analysis,

the fine-grained sandstone lenses consist mostly of subangular

to very angular, xenomorphic quartz (60–90 %) and K-feldspar

grains (5–10 %); kaolinite (5–10 %) and accessory minerals:

Ti-magnetite, muscovite, rutile, zircon, calcite and rock frag-

ments (<1 % respectively). The size distribution of the quartz

grains is bimodal with two peaks at ~50 and 200–250 μm.

Undulatory extinction of quartz is common. Amongst the quartz

crystals with normal extinction there are rounded grains —

similar to resorbed volcanic crystals. Feldspar grains have

an average size of 150–200 μm, and they are often strongly

fractured.

The rock forming minerals suggest a terrigenous prove-

nance. The significant amount of quartz crystals with undula-

tory extinction, the lack of plagioclase, amphibole or pyroxene

crystals and even altered volcanic glass fragments exclude

the pyroclastic origin. Although there are some minerals —

biotite, zircon, glauconite and presumably resorbed quartz

grains — which could be derived from resedimented pyroclas-

tics/volcanocalstics. The dominance of angular to subangular

grains and the significant amount of K-feldspar in the sand

fraction denote a short transportation path and a nearby source

area. The textural features suggest that the clay minerals are

probably allogenic (illite) in the matrix and most likely authi-

genic (kaolinite) in the lenticular sandstone domains.

Fig. 5. Mineralogical composition of dolomite beds of the Construction site, based on XRD analyses (see Electronic supplement I). The layer

numbers are shown in Fig. 3A.

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Sedimentological description of the Road-cut site

The Road-cut site („Siklóbevágás” in Hungarian) is nearly

30 meter-long and 3–5 meter-high, where three formations are

exposed (Fig. 10).

The uppermost beds of the Templomhegy Member is

exposed in the northern part of the Road-cut site. The dolomite

and yellowish dolomarl beds are similar to the uppermost

exposed dolomite beds of the Construction site (Pozsgai et al.

2017). This part of the sequence is dominated by whitish dolo-

mite and yellowish dolomarl beds, frequently intercalated by

very thin (few-mm-thick) reddish, yellowish or greenish

claystones.

Only one exception is a 15-cm-thick, variegated claystone

layer that is dominantly reddish and greenish, yellowish, pur-

plish mottled, and contains lenticular, indurated, calcareous

intercalations. This bed contains elongated carbonate con-

cretions and angular dolomite clasts as well. On the top of

the claystone bed, a friable carbonate crust occurs. This clayey

intercalation is probably identical to the reddish claystone

with calcareous crust in the Construction site (see Fig. 8).

The water-screened residue of Templomhegy Member at

the Road-cut site was not productive for fish fossils, but a few

poorly preserved sauropterygian bones were discovered from

these horizons (Ősi et al. 2013).

The siliciclastic content of the rocks increases upwards in

the section and the Templomhegy Member is overlain by

an almost 15-m-thick sequence of the Mészhegy Sandstone

Formation (Vörös 2009; Fig.10B), which

is made up

of cyclic

alternation of weakly cemented, greyish, yellowish, purplish

or greenish sandstone and siltstone layers, reddish, purplish or

variegated clay strata, and subordinately greyish dolomite and

yellowish dolomarl beds (Vörös 2010; Pozsgai et al. 2017).

Although several sharp surfaces dissect the succession (Vörös

2009), no unequivocal boundary can be recognised between

the Templomhegy Dolomite and the Mészhegy Sandstone in

this section. However, in agreement with Vörös (2010), we

also suggest to define the boundary between the two forma-

tions at the level where a continuous sandstone–siltstone–

claystone assemblage overlies the carbonate dominated strata

in the section.

The sediments of Mészhegy Sandstone Formation in

the Road- cut section can be divided into the following major

lithofacies (based on Rálisch-Felgenhauer 1985;

Vörös 2009,

2010; Pozsgai et al. 2017; Fig. 11

Conglomerate (Fig. 11A): Only one conglomerate bed

occurs in the section. It is 50-cm-thick and unconformly over-

lies a claystone horizon. It is a matrix supported (with a car-

bonatic, clayey, sandy matrix), polimict conglomerate includes

upward fining, rounded clasts (dolomite, dolomarl, limestone,

sandstone, claystone and polycrystalline quartz clasts).

Sandstone (Fig. 11B): These are fine to coarse grained,

(sublitharenite, subarcose) sandstones. In the lower part of

the section mainly greyish sandstones are present, containing

reworked rounded claystone (1–2 cm) and angular marl clasts

(1–10 cm) and smaller amount of quartz pebbles (<0.5 cm).

In the upper part of the section greenish, reddish, purplish,

greyish sandstones layers occur. Crossbedding is locally

visible.

Claystone, silty and sandy claystone (Fig. 11C, D): This is

the dominant lithofacies in the Road-cut section. The clay-

stone frequently include greyish, fine-grained, lenticular sand-

stone bodies. Greenish siltstone and variegated claystone beds

appear in the older part of the section whereas yellowish,

brownish claystone and claystone layers occur in the upper

part.

Cellular marl (Fig. 11E, F): Yellowish marl beds, which are

characterized by polygonal cracks and cellular structure.

Its mineralogical composition is dominated by calcite (around

75 %); illite (10 %), kaolinite (10 %) and quartz (5 %) also

occur. Calcite veins (0.5–2 cm thick) are very common.

Several beds of the Mészhegy Sandstone Formation were

sampled for screen-washing, but only three brownish-greyish

sandstone beds (L3–L4–L5) yielded microvertebrate remains

(Fig. 10 and see Electronic supplement II). The most abundant

vertebrate material was recovered from the uppermost sand-

stone bed (L5 in Fig. 10B), which provided more than three

hundred teeth and scales from different marine fish and sau-

ropsid reptilian taxa (see Table 1). The Triassic sequence of

the Road-cut site on Templom Hill is covered by the Pliens-

bachian Somssichhegy Limestone Formation (Fig. 10).

Discussion

The succession of the Construction site is composed pre-

dominantly of carbonates (dolomite) with various clay con-

tent, i.e. it is made up of alternating dolomite and dolomarl

layers (Fig. 3). At the outcrop scale, we use the terms “dolo-

marl” and “dolomite” in a descriptive sense, the more massive

and solid, light-coloured beds with high dolomite content

Fig. 7. Ternary diagram of dolomarl and claystone beds of the con-

struction site based on the results of XRD analyses. The layer numbers

are shown in the Fig. 3A.

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production is slowed down or halted and the terrigenous influx

can increase, resulting in the predominance of siliciclast depo-

sition (Wilson 1967; Brachert et al. 2003; Carcel et al. 2010;

Caracciolo et al. 2013; Chiarella et al. 2017 and references

therein). The short-term climate changes play also an impor-

tant role in marine areas near the mainland (Coffey & Read

2007; Caracciolo et al. 2013; Zeller et al. 2015). The humidity

significantly influences the terrigenous sediment influx from

the land to the marine realm and thus the carbonates formed

during the dryer periods are frequently replaced by siliciclastic

sediments during humid conditions.

The observed succession was deposited near to the land in

an inner ramp lagoon and the related tidal flat environments

where the siliciclastic input and coeval carbonate production

were significantly controlled by different allocyclic (e.g.,

climatic or sea-level changes) factors (see also Brachert et al.

2003; Colombié et al. 2012; Chiarella et al. 2017). The studied

section of the Construction site was deposited between

the Late Anisian and Carnian based on its stratigraphic posi-

tion (Rálisch-Felgenhauer & Török 1993; Török 1998; Haas

2001) and vertebrate fossils (Ősi et al. 2013), which period

was frequently characterized by significant fluctuation in

the climatic conditions and rainfall intensity (e.g., Mutti &

Weissert 1995; Simms et al. 1995; Feist-Burkhardt et al. 2008;

Preto et al. 2010). We suggest that the interval of higher silici-

clast content observed in the middle part of the section (beds

10 to 22; Fig. 3), between two carbonate rich sequences,

reflects a temporary change in the prevailing climate (Fig. 12).

The lower and upper part of the studied section was formed

in the periods of more arid climate, when the siliciclastic

influx from the land was subordinate, while the middle part

of the section that is characterized by higher siliciclastic

content (including sandstone and claystone beds) was

deposited during a more humid phase of enhanced rainfall

intensity. Furthermore, the reddish silty claystone facies

(Fig. 8) situated in the middle part of the section, represents

recurring paleosol formation, and indicates that the marine

sediments were repeatedly exposed to subaerial conditions,

suggesting relative sea-level falls (Fig. 12). A global sea-

level fall was recognized around the Late Ladinian and

Early Carnian by Haq et al. (1988) and Ruffell (1991), which

period coincides with the assumed depositional age of the car-

bonate-dominated sedimentary sequence of the Construction

site and thus this eustatic change might be correlated with

Fig. 9. A — Sandy claystone bed of the construction site and its mineral composition (B) based on XRD analyses (see Electronic supplement I).

C — Claystone composed of illite, quartz, K-feldspar, calcite and accessory minerals: glauconite, biotite, magnetite, and zircon. Backscattered

electron image (bed 10). D — Fine-grained sandstone, consisting of subangular to angular, xenomorphic quartz and K-feldspar crystals;

kaolinite and accessory minerals: Ti-magnetite, muscovite, rutile, zircon, calcite and rock fragments. The transition towards the finer grained

domains is continuous. Backscattered electron image (bed 10).

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Fig. 3) is made up of dominantly aphanitic to very finely crys-

talline dolomite beds with subordinate argillaceous deposits.

This part of the section can be characterized by carbonate sedi-

mentation where the siliciclastic influx from the land was

subordinate. This part of the section was deposited in the inner

ramp zone was flooded due to sea-level rise. The top of this

interval corresponds to a relatively thick sandstone bed (bed 10;

see Figs. 3 and 9A), located between the massive dolomite and

the most marly interval. This sandstone with high quartz con-

tent indicates that the climate has become more humid and

significant siliciclastic influx arrived from the land (Fig. 9B).

Above this layer, more argillaceous sediment (dolomarl beds)

can be observed representing the middle part of the section

(beds 10 to 22; Fig. 3). This part was most likely deposited in

a period, when the fine terrigenous input significantly

increased most probably due to the enhanced humidity allo-

wing the accumulation of more argillaceous sediments.

The increase in clastic influx may have been also related to

a relative sea-level fall, because the presence of reddish silty

claystone bed in this part of the section shows evidence of

subaerial exposure. The pedogenesis took place under oxidi-

zing conditions, as indicated by the red colour of the claystone

bed (Construction site; top of beds 14 and 21). However,

neither karstic features nor downcutting by streams were

observed, indicating that the subaerial exposure was relatively

short (e.g. Wilson 1967). The middle part of the section is

cove red by beds of massive, homogenous very finely crystal-

line dolomite (beds 23–29;

Fig. 3) with thin dolomarl interca-

lations, where the vertebrate fossils are very rare. Dolomarl

beds in this section are characterized by a high dolomite con-

tent (about 85 %) indicating that the carbonate sedimentation

became dominant again while the amount of siliciclastic sedi-

ments decreased. The low siliciclastic content and the lack of

traces of pedogenesis in the youngest part of Templomhegy

Member exposed in Construction site suggest a more arid con-

dition and a sea-level rise during the accumulation.

The last dolomitic layer is unconformly covered by a thin-

ner siliciclastic sediment package which is composed of sand-

stone, siltstone and claystone beds. The vertebrate fossils are

completely absent from this sequence, but the petrographical

analyses indicates that this clastic succession is part of

the Carnian Mészhegy Sandstone Formation (Pozsgai et al.

2017). The sediments of the Mészhegy Sandstone Formation

can be better investigated at the Road-cut site where its

15-meter thick sequence is exposed (Rálisch-Felgenhauer

1985; Vörös 2009, 2010).

Despite the several decades of research in the Road-cut sec-

tion, the age and the depositional environment was difficult to

define, because the mentioned authors did not find any fossils

which they could use for more detailed paleoenvironmental

reconstruction. The unfossiliferous nature of this formation

(except for some unidentified reptile bones) remained an

accepted viewpoint until the discovery of a relatively rich

vertebrate assemblage in 2012. The discovered fish remains

(e.g., Paleobates angustissimus, Gyrolepis sp.) indicates

a marine environment for at least the sampled beds, that are

situated in the lower (bed L3; Fig. 10B) and the upper part

(beds L4–L5; Fig. 10B) of the Road-cut section (Fig. 10 and

Electronic supplement II). The fish fauna of the Mészhegy

Sandstone Formation (dominated by Hybodus) is significantly

different from that of the Templomhegy Dolomite Member

(dominated by durophagous hybodontiforms, such as Palaeo

bates and Lissodus). Therefore the fish fossils of the Road-cut

site cannot be derived from the underlying marine deposits

(for detailed differences in the fauna compositions see

Electronic supplement II). Furthermore, besides collecting

vertebrate fossils, several samples were also taken for palyno-

logical investigations. One of these (from a sandy-claystone

bed from the lowermost part of the section; see Fig. 10B)

has provided a relatively diverse sporomoph assemblage

(Patinasporites densus, Infernopollenties sp., Aratrisporites

spp., Ovalipollis spp., and Triadispora spp.) indicating

a Carnian age for the lower part of the section (Ősi et al. 2013).

This also suggests that the sediments were deposited in

a nearshore environment characterized by a high input of land-

derived organic matter (Ősi et al. 2013). Nevertheless,

the col lected fossils are not sufficient to completely exclude

the possibility of fluviolacustrine sedimentation in the succes-

sion exposed in the Road-cut section (see Vörös 2009, 2010).

However, we suggest that at least four fossil-bearing sand-

stone beds, occuring in the lowermost and the uppermost part

of the section, were deposited in a nearshore (based on palyno-

logical data; Ősi et al. 2013), shallow marine environment

(based on marine fish assemblage) characterized by intense

terrigenous input.

Conclusions

1. Four main lithofacies were identified and interpreted in

the newly discovered Construction vertebrate site, which is

dominantly made up of alternating dolomite and dolomarl

layers. The four lithofacies units can be defined as follows:

• dolomite, deposited in a shallow, restricted lagoon

environment;

• dolomarl, as shallow marine sediment, where the enhan-

ced terrigenous input was the results of the more humid

climate;

• reddish silty claystone, formed as paleosol;

• sandstone indicating a terrigenous input.

2. The stratigraphical, sedimentological and paleontological

investigations revealed that the sediments of the Construction

vertebrate site were formed in the nearshore (subtidal to

peri tidal) zone of a ramp, where the alternating sedimenta-

tion was mostly controlled by climatic changes. However,

the recurring paleosol formation in the middle part of

the section also indicates episodic sea-level fall events,

when the marine sediments were repeatedly exposed to sub-

aerial conditions.

3. The Construction site can be divided into three main parts:

the lower part (from bed 1 to 9) can be characterized by

carbonate sedimentation where the siliciclastic influx from

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environment characterized by high siliciclastic input from

the mainland.

Acknowledgments: We thank Annette E. Götz, János Haas

and Attila Vörös for their useful comments and suggestions

that greatly improved our manuscript. The authors are emi-

nently thankful to Gyula Konrád, Krisztina Sebe, Tamás

Budai, István Dunkl, Emese Szőcs, Andrea Mindszenty,

Georgina Lukoczki, Orsolya Sztanó, György Czuppon and

Sándor Józsa for useful discussions and consultations.

The authors are grateful to Krisztina Buczkó and Kristóf Fehér

for their help in perfoming scanning electron microscopy

micrographs. We thank Réka Kalmár and János Magyar for

technical assistance. Our work was supported by the National

Research, Development and Innovation Office (NKFIH

K116665 and K124313), Hungarian Academy of Sciences

Lendület Program, Hungarian Natural History Museum,

Eötvös Loránd University, University of Pécs, University of

Göttingen, and the Danube–Dráva National Park.

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SEDIMENTOLOGY OF TRIASSIC VERTEBRATE LOCALITIES IN VILLÁNY HILLS, SOUTHERN HUNGARY

GEOLOGICA CARPATHICA

, 2019, 70, 2, 135–152

Electronic supplement

Supplement I:

Mineralogical composition based on X-Ray powder diffraction data of the Construction site

This file contains the detailed results of the X-ray powder diffraction analyses of the presented layers (beds). The results

shows the mineralogical compositional changes of the presented beds indicating changes in depositional environment.

Table S1: The bulk mineralogical composition of analysed beds showing the changes in depositional environment.

Samle ID

Bulk mineralogical composition [wt. %]

Silicates

Carbonates

Clay minerals

anatase

goethite

quartz plagioclase K-feldspar calcite Mg-calcite ankerite dolomite

10 Å phyllosilicate

(muscovite/illite/biotite?) kaolinite

swelling

clays

Layer 1

0.3

3.8

95.9

Layer 2

3.3

82.8

5.6

7.2

1.0

Layer 5

1.8

96.5

1.7

Layer 6

4.0

0.2

82.9

6.3

6.1

0.5

Layer 7

1.5

20.1

77.1

1.3

Layer 7

2.7

0.2

81.5

7.4

7.6

0.6

Layer 8

12.8

0.1

5.8

62.1

8.3

2.1

0.7

Layer 9

1.5

94.5

3.1

0.9

Layer 10_A

47.7

0.2

32.2

17.8

2.1

Layer 10_B

49.7

12.4

4.5

0.2

15.5

7.3

10.4

Layer 10_C

58.2

11.6

0.8

0.6

15.8

5.1

7.9

Layer 11

4.1

0.1

86.6

2.0

7.2

Layer 12

7.9

55.1

24.8

8.9

2.3

Layer 13

4.4

0.5

89.2

5.9

Layer 14

21.9

3.1

1.6

42.7

11.5

3.6

13.6

Layer 15_A

7.7

0.6

1.3

76.7

11.4

2.3

Layer 15_B

3.2

0.3

0.6

0.8

93.6

1.5

Layer 15_C

1.9

0.5

7.0

90.4

0.2

Layer 16_A

22.8

3.1

16.7

2.4

24.3

10.8

18.8

1.1

Layer 16_B

9.6

1.2

0.8

60.0

14.3

2.3

9.8

0.3

1.7

Layer 17_A

4.2

1.1

86.0

7.5

1.2

Layer 17_B

2.6

0.6

0.3

93.2

3.3

Layer 18

3.7

0.8

85.7

5.6

3.4

0.8

Layer 19

3.1

0.3

0.8

88.3

3.3

4.1

Layer 20

4.6

0.3

81.2

6.3

6.8

0.8

Layer 21

18.9

2.8

11.5

3.1

28.2

7.5

24.2

0.3

3.4

Layer 22

4.7

0.4

81.3

6.9

5.7

1.0

Layer 23

0.8

0.4

3.0

95.2

0.6

Layer 24

2.5

0.4

3.8

90.8

2.1

0.3

Layer 26_A

4.1

0.3

89.9

2.8

Layer 26_B

3.1

0.3

87.8

3.9

3.8

1.0

Layer 27

8.1

1.2

1.5

44.5

16.7

9.6

2.1

15.9

0.4

Layer 28

3.0

0.4

1.4

87.6

2.9

4.2

0.5

Layer 29

0.5

0.3

20.8

78.3

0.1

BOTFALVAI, GYŐRI , POZSGAI , FARKAS, SÁGI, SZABÓ and ŐSI

GEOLOGICA CARPATHICA

, 2019, 70, 2, 135–152

Supplement II:

Short summary of discovered fish remains from Triassic Villány vertebrate locality

This file contributes supplementary data to the fish faunas unearthed in the Triassic Villány vertebrate fossil sites. The data

include the summarized distribution, specimen numbers and habitat preferences of the Villány fish taxa, including Hybodus sp.,

Palaeobates angustissimus, Polyacrodus sp., Lissodus sp., Gyrolepis sp., Severnichthys acuminatus and Actinopterygii indet..

The data shows the quantitative dominance of Palaeobates angustissimus in the Templomhegy Dolomite Member at

the Construction site, and also that of Hybodus sp. in the Mészhegy Sandstone Formation at the Road-cut site. Three fish forms

(Palaeobates angustissimus, Gyrolepis sp. and Severnichthys acuminatus) are referable to marine habitats, while Hybodus sp.,

Polyacrodus sp., Lissodus sp. and the indeterminate actinopterygian remains are less informative in paleoenvironmental point

of view.

Table S2: Short summary of the discovered Villány fish remains.

Site

Formation

Age

Taxon

Description

Quantity

Paleoenvironment

References

Construction site

Templomhegy

Dolomite

Member

Ladinian

Palaeobates angustissimus

Crushing teeth; lentoid,

elongated or circular occlusal

view with reticulated

occlusal surface

1272

marine

Dalla Vecchia & Carnevale

2011; Diedrich 2003, 2009; Pla

et al. 2013; Schultze & Kriwet

1999

Lissodus sp.

Low-crowned crushing teeth

with sharp transversal crest

and no distinctive surface

sculpting

marine–brackish–freshwater Cappetta 2012

Gyrolepis sp.

Simple, pointed, conical

teeth and thick scales with

striated ganoine layer

180

marine

Allard et al. 2015; Cavicchini et

al. 2018; Diedrich 2003, 2009;

Landon et al. 2017; Nordén et

al. 2015; Whiteside et al. 2016

Severnichthys acuminatus

"Saurichthys"- and

"Birgeria"-type teeth; conical

teeth with fine, apicobasal

striae below and on the cap

marine

Allard et al. 2015; Cavicchini et

al. 2018; Korneisel et al. 2015;

Nordén et al. 2015; Mears et al.

2016; Whiteside et al. 2016

Actinopterygii indet.

Two different types of teeth

and badly preserved, simple

ganoid scales

300

marine–brackish–freshwater Nelson 2006

Mészhegy

Sandstone

Formation

Carnian

Yielded no fish fossils

Road-cut site

Templomhegy

Dolomite

Member

Ladinian

Yielded no fish fossils

Mészhegy

Sandstone

Formation

Carnian

Hybodus sp.

Isolated cusps with

apicobasal striation, circular

cross-sectrion, and smooth

cutting edges

194

marine–brackish–freshwater

Cuny 2012; Dica & Codrea

2006; Klug et al. 2010 and

references therein

Palaeobates angustissimus

Crushing teeth; lentoid,

elongated or circular occlusal

view with reticulated

occlusal surface

marine

Dalla Vecchia & Carnevale

2011; Diedrich 2003, 2009; Pla

et al. 2003; Schultze & Kriwet

1999

Polyacrodus sp.

Fragmentary teeth with large

central cusp, well-defined

lingual apron and occlusal

striation

marine–brackish–freshwater Böttcher 2015; Diedrich 2003,

2009; Hagdorn & Mutter 2011

Lissodus sp.

Low-crowned crushing teeth

with sharp transversal crest

and no distinctive surface

sculpting

marine–brackish–freshwater Cappetta 2012

Gyrolepis sp.

Simple, pointed, conical

teeth and thick scales with

striated ganoine layer

marine

Allard et al. 2015; Cavicchini et

al. 2018; Diedrich 2003, 2009;

Landon et al. 2017; Nordén et

al. 2015; Whiteside et al. 2016

Severnichthys acuminatus

"Saurichthys"- and

"Birgeria"-type teeth; conical

teeth with fine, apicobasal

striae below and on the cap

marine

Allard et al. 2015; Cavicchini et

al. 2018; Korneisel et al. 2015;

Nordén et al. 2015; Mears et al.

2016; Whiteside et al. 2016

Actinopterygii indet.

Two different types of teeth

and badly preserved, simple

ganoid scales

marine–brackish–freshwater Nelson 2006

iii

SEDIMENTOLOGY OF TRIASSIC VERTEBRATE LOCALITIES IN VILLÁNY HILLS, SOUTHERN HUNGARY

GEOLOGICA CARPATHICA

, 2019, 70, 2, 135–152

Table S3: Specimen numbers of taxa per sampled layers.

Site

Formation

Age

Taxon

Layer 22

Layer 20

Layer 18

Layer 14

∑

Construction site

Templomhegy Dolomite Member

Ladinian

Palaeobates angustissimus

1266

1272

Lissodus sp.

Gyrolepis sp.

173

180

Severnichthys acuminatus

Actinopterygii indet.

263

300

Site

Formation

Age

Taxon

∑

Road-cut site

Mészhegy Sandstone Formation

Carnian

Hybodus sp.

191

194

Palaeobates angustissimus

Polyacrodus sp.

Lissodus sp.

Gyrolepis sp.

Severnichthys acuminatus

Actinopterygii indet.

Fig. S1. Fish remains from the Villány vertebrate site. A — Hybodus sp. main cusp in lingual view. B — Palaeobates angustissimus lateral

tooth in occlusal view. C — Polyacrodus sp. fragmentary tooth in labial view. D — Lissodus sp. tooth in occlusal view. E — Gyrolepis sp. tooth

in profile view. F — Gyrolepis sp. scale in external view. G — Severnichthys acuminatus tooth in labial or lingual view. H — Actinopterygii

indet. tooth type A in labial or lingual view. I — Actinopterygii indet. tooth type B in occlusal view. J — Actinopterygii indet. scale in external

view. Scale bars: A, B, E, F, G: 1 mm; C, D, H, I, J: 500 μm.

BOTFALVAI, GYŐRI , POZSGAI , FARKAS, SÁGI, SZABÓ and ŐSI

GEOLOGICA CARPATHICA

, 2019, 70, 2, 135–152

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