A fish fauna from the lowermost Bartonian of the Transylvanian Basin, Romania

Abstract

A fish fauna newly discovered in the middle Eocene marine sediments cropping out near the village of Luna de Sus, Romania, completes the fossil record of the Eastern European region. Teeth belonging to 15 species of Chondrichthyes and two species of Actinopterygii are herein recorded from the lowermost Bartonian deposits. These Paleogene fish document a marine tropical environment of medium deep waters in the northwestern area of the Transylvanian Basin. The vertical distributions of extant equivalent taxa allow a sea depth estimation of 100 to 200 m. The warm climate is documented by both the present faunal assemblage and previous palynological studies. It is important to note the presence of the scarcely known and poorly understood pycnodont species Phacodus punctatus and of the oldest representative of Labridae from this Carpathian area. The diversity of the fauna was found to be average compared to some areas from Western Europe or North Africa, but it falls within the regional diversity range of the Eastern European localities.

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Trif, Nicolae, Codrea, Vlad, and Arghiuș, Viorel. 2019. A fish fauna from the lowermost Bartonian of the Transylvanian Basin,
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A fish fauna from the lowermost Bartonian
of the Transylvanian Basin, Romania
Nicolae Trif, Vlad Codrea, and Viorel Arghiuș
ABSTRACT
A fish fauna newly discovered in the middle Eocene marine sediments cropping
out near the village of Luna de Sus, Romania, completes the fossil record of the East-
ern European region. Teeth belonging to 15 species of Chondrichthyes and two spe-
cies of Actinopterygii are herein recorded from the lowermost Bartonian deposits.
These Paleogene fish document a marine tropical environment of medium deep waters
in the northwestern area of the Transylvanian Basin. The vertical distributions of extant
equivalent taxa allow a sea depth estimation of 100 to 200 m. The warm climate is doc-
umented by both the present faunal assemblage and previous palynological studies. It
is important to note the presence of the scarcely known and poorly understood pycno-
dont species Phacodus punctatus and of the oldest representative of Labridae from
this Carpathian area. The diversity of the fauna was found to be average compared to
some areas from Western Europe or North Africa, but it falls within the regional diver-
sity range of the Eastern European localities.
Nicolae Trif. Department of Geology, Faculty of Biology-Geology, Babeş-Bolyai University, 1 Kogălniceanu
St., 400084, Cluj-Napoca, Romania and Brukenthal National Museum, Natural History Museum, Sibiu,
Romania, 1 Cetății St., Sibiu, 550160, Romania. nicolae.trif@gmail.com
Vlad Codrea. Department of Geology, Faculty of Biology-Geology, Babeş-Bolyai University, 1
Kogălniceanu St., 400084, Cluj-Napoca, Romania. vlad.codrea@ubbcluj.ro
Viorel Arghiuș. Environmental Sciences Department, Faculty of Environmental Sciences and Engineering,
Babeş-Bolyai University, 30 Fântânele St., 400294, Cluj-Napoca, Romania. arghius.viorel@ubbcluj.ro
Keywords: sharks; bony fish; Eocene; Căpuș Formation; Tethys Sea
Submission: 29 July 2018. Acceptance: 12 August. 2019.

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
2
INTRODUCTION
Eocene-age fish teeth in the Transylvanian
Basin were first described in the 1850s by Neuge-
boren (1850, 1851), who listed material belonging
to 65 species of sharks from the southern side of
this sedimentary basin. Almost half a century later,
Koch (1894) reported a few new occurrences of
Eocene fish teeth (again, mostly sharks) originating
from the northwestern side of the basin. It is likely
that in Koch's list, both middle and upper Eocene
strata were represented, but the actual stratigraphy
could not be confidently determined from this work.
This same author appended these data to a larger
subsequent list that includes the former Austro-
Hungarian Empire (Koch, 1900). Following these
researchers, only a handful of geologists focused
on this topic, surveying mostly the northwestern
area of the basin. The first one was Fuchs (1963,
1966), who described some fragmentary Myliobatis
teeth from the Priabonian of Cluj-Napoca surround-
ings. The diversity of the Eocene fish fauna only
started to be revealed a couple of decades later
(Șuraru et al., 1980; Șuraru and Șuraru, 1987), but
these studies focused exclusively on the same
geological age. The list of Priabonian species has
been supplemented again in the final decade of the
last century (Dica et al., 1996; Codrea et al., 1997).
The first definite mention of the middle Eocene fish
fauna was at the beginning of the twenty-first cen-
tury in an unpublished Ph.D. thesis (Dica, 2006).
From the type locality of the Căpuș Formation
(Căpușul Mic), Jaekelotodus sp., Striatolamia mac-
rota (Agassiz, 1843), Myliobatis sp., Aetobatus
irregularis (Agassiz, 1843) and Pycnodontidae
indet. have been illustrated and described.
Recent surveys of the middle Eocene depos-
its of the Luna de Sus locality revealed a rich
assemblage of fish teeth, a rostral and a dermal
spine, and a tail sting. Luna de Sus is situated on
the northwestern side of the Paleogene Transylva-
nian Basin, central Romania, at about 10 km west-
ward from the city of Cluj-Napoca (Figure 1). This
paper is the first to describe this locality for fossil
fishes in Romania.
GEOLOGICAL SETTING
According to the paleogeographic reconstruc-
tions of the Tethys Sea (Meulenkamp and Siss-
ingh, 2003), during the late Lutetian, the north-
western area of the recent Transylvanian Depres-
sion was part of a Paleogene sedimentary basin
covered by a shallow sea spreading over the
thrusting nappes of the Apuseni Mountains land-
mass. To the east, this sea was in connection with
the Eastern Carpathians Paleogene Flysh trough
(i.e., the outer “Moldavides”, sensu Săndulescu,
1984). The Căpuş Formation (Popescu, 1978) of
middle Eocene age is exposed in the northwestern
area of the Transylvanian Basin (Gilău sedimen-
tary area; Rusu, 1987).
The sedimentology illustrates a paleoenviron-
ment of an inner continental shelf of an open sea
FIGURE 1. Location of the Luna de Sus study site (after the Geological Map of Romania, 1:200.000 folio Cluj, simpli-
fied and modified).

PALAEO-ELECTRONICA.ORG
3
with a tidal regime (Rusu et al., 2004). The forma-
tion consists mainly of marls bearing several
important mollusk biohorizons with regional distri-
butions useful for correlating these deposits. Its
basal segment represents the level with Pycno-
donte brogniartii Bronn, 1831, while the top bears
the level with Nummulites perforatus Montfort,
1808 (Figure 2.1). The latest geological study
shows that the age of the formation is Lutetian-Bar-
tonian, but only its basal portion is Lutetian, with
the remaining portion belonging to the Bartonian
(Rusu et al., 2004).
The outcrop is situated on a very steep ravine,
a right tributary of the Feneș Creek, on the south-
ern side of the village of Luna de Sus. Its left bank
is covered by a landslide; therefore, the main inter-
est is restricted to only a few meters on the right
bank of the ravine. The nearest equivalent out-
crops situated at 650 m south-west, 400 m east
and 80 m north of the gully did not reveal any fish
fauna.
The Luna de Sus lithostratigraphic log is over
10 m thick, including: 7 m of marls bearing a basal
Pycnodonte brongniarti lumachelle; 0.1 m marl with
Sokolovia eszterhazyi Pávay, 1871, lumachelle;
0.17 m calcareous sandstone with some microcon-
glomerate elements. The last two layers are where
the fish remains were found. Above these layers
there is a very thin (0.07 m) layer of quartz micro-
conglomerate followed by 1 m of clayey shale and
2.4 m of marls bearing lumachelle of Nummulites
perforatus. The outcrop exposes rocks of a geolog-
ical age very close to the lowermost Bartonian, just
above the Lutetian/Bartonian boundary (Figure
2.2).
MATERIAL AND METHODS
The specimens were collected during 20 short
field excursions, from March 2009 to September
2017. The photographs were taken with a Nikon
D80 camera mounted on a Nikon SMZ 1000 binoc-
ular microscope and with a Nikon D700 camera
and a 105 mm Sigma lens. The described material
is stored at the Faculty of Environmental Science
and Engineering, Babeș-Bolyai University, Cluj
Napoca, Romania (abbreviated hereinafter, FES).
FIGURE 2. Lithostratigraphy of the studied outcrop. 1, Lithostratigraphic log of the Eocene deposits from the Gilău
sedimentary area (modified and simplified after Rusu et al., 2014). 2, Lithostratigraphic log at Luna de Sus.

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
4
The single exception is the VT459 specimen
donated to the Babeş-Bolyai University Paleontol-
ogy-Stratigraphy Museum in Cluj Napoca (abbrevi-
ated hereinafter, BBUPSM). The taxonomic
identification of the described material included a
comparison with the high-resolution images of the
holotype and paratypes of Macrorhizodus nolfi
Zhelezko, 1999, from the Darwin State Museum,
Moscow (DSM) and of the holotype of Phacodus
punctatus Dixon, 1850, from the Natural History
Museum, London (NHM). The systematic paleon-
tology part follows Cappetta (2012) and Last et al.
(2016a). The use of the name Otodus Agassiz,
1843, follows Cappetta (2012), and it includes
Carcharocles as a subgenus. The terminology fol-
lows Deynat (1998), Cappetta (2012) and
Hovestadt and Hovestadt-Euler (2013).
During the field excursions about 175 kg of
sediment (sandstone) was collected. The high iron
content of the sandstone made it somewhat inert to
the acetic acid reaction that was used to dissolve
the rock. Therefore, the process of disaggregation
has been tedious and time consuming. A 0.5 mm
mesh sieve was used to sieve the resulting sedi-
ment. The disaggregation of the sandstone sam-
ples together with the direct field collecting yielded
ca. 75 fish teeth, rostral and dermal spines and a
tail sting.
SYSTEMATIC PALEONTOLOGY
Class CHONDRICHTHYES Huxley, 1880
Subclass ELASMOBRANCHII Bonaparte, 1838
Order HETERODONTIFORMES Berg, 1937
Family HETERODONTIDAE Gray, 1851
Genus HETERODONTUS Blainville, 1816
Heterodontus sp.
Figure 3.1-7
Material. Two teeth (FES 045 and 098).
Description. Both teeth are small. Only the speci-
men FES 045 is complete, with a length and width
of 8.1 by 3.2 mm. The teeth are labio-lingually
compressed and mesio-distally elongated. A retic-
ulated ornamentation is present on the labial side.
The reticulation becomes smaller and finer towards
the labial edge. The lingual face has an ornamen-
tation of weak parallel ridges perpendicular to the
mesial-distal direction. A longitudinal main ridge is
present on the same direction, interrupted by a
large central bump (in FES 098), or by a central
massive functional wear facet (in FES 045). In
occlusal view, the specimen FES 045 has a gen-
eral sigmoid shape, with a rounded end in the
mesial direction and an angled, pointed one dis-
tally. The crown overhangs the root in all directions.
The root is low and flat. We cannot observe any
marginal lingual or labial foramina because of
some strongly cemented sediment adherent to the
root.
Remarks. There are several Eocene species of
Heterodontus: H. pineti Case, 1981 (Priabonian,
Georgia, USA), H. vincenti Leriche, 1905 (Lutetian,
Belgium) and H. wardenesis Casier, 1966 (Ypre-
sian, England). Unfortunately, the preservation sta-
tus of the fossils at our disposal does not allow us
an assignation below the genus level.
Order CARCHARHINIFORMES Compagno, 1977
Family CARCHARHINIDAE Jordan and Evermann,
1896
Subfamily CARCHARININAE Jordan and
Evermann, 1896
Genus RHIZOPRIONODON Whitley, 1929
Rhizoprionodon ganntourensis Arambourg, 1952
Figure 3.8-11
Material. Two teeth (FES 076 and 082).
Description. The teeth are small, their height
around 2.5 mm and their width of almost 4 mm.
The cusp is strongly bent distally. No cusplets are
present, but the tooth exposes a distinct distal heel.
The mesial side is elongated and slightly convex.
The labial base of the enamel is quasi rectilinear,
while on the lingual side it is slightly arched. The
root foramen is small, oval and shifted distally.
Remarks. The teeth of this genus are character-
ized by a distinct gynadric heterodonty (Cappetta,
2012). In males, the central part of the cusp is
thicker, and the mesial cutting edge is less convex.
Our specimens are strongly compressed in the
labial-lingual direction, and the central part of the
cusp is strongly bent towards the rear. We interpret
our specimens as belonging to females. According
to Cappetta (1987), the teeth of this genus are very
similar to the ones found in the Scoliodon and Lox-
odon genera. No progress has been made so far in
the last quarter century to settle this issue, and the
data remain the same in the more recent works
(Cappetta, 2012).
Only a single Eocene species is known, Rhi-
zoprionodon ganntourensis. The species is also
present in the middle Eocene (Lutetian) of Morocco
(Arambourg, 1952), Uzbekistan (Case et al., 1996),
Togo (Cappetta and Traverse, 1988), the upper
Eocene of France (Cappetta and Nolf, 1981), as
well as the Eocene (Ciobanu, 2002) and the upper
Eocene (Priabonian) of Romania (Dica, 2006).
Subfamily GALEOCERDINAE Whitley, 1929
Genus GALEOCERDO Müller and Henle, 1838

PALAEO-ELECTRONICA.ORG
5
FIGURE 3. Fish fossils from Luna de Sus. 1-4, Heterodontus sp., (FES 045). 5-7, (FES 098); 8-9, Rhizoprionodon
ganntourensis, (FES 076). 10-11, (FES 082). 1, 5, occlusal views. 2, basal view. 3, 6, 8, 11, lingual views. 4, 7, 9, 10,
labial views. Scale bars equal 1-7 (5 mm), 8-11 (1 mm).

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
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FIGURE 4. Fish fossils from Luna de Sus. 1-2, Galeocerdo cf. eaglesomei, (FES 043). 3-4, (FES 044). 5-7, Physoga-
leus secundus, (FES 050). 8-11, (FES 053). 1, 3, 5, 8, lingual views. 2, 4, 6, 9, labial views. 7, 10, apical views. 11,
basal view. 1', detail of serration. Scale bars equal 5 mm.

PALAEO-ELECTRONICA.ORG
7
Galeocerdo cf. eaglesomei White, 1955
Figure 4.1-4
Material. Two incomplete teeth (FES 043 and
044).
Description. The teeth are broad but also high,
with a triangular crown that bears a central distinct
cusp inclined distally. FES 043 is 15 mm wide and
12 mm tall, while FES 044 is 15 mm wide and 10
mm tall. The mesial cutting edge is slightly convex
in one specimen and completely straight in the
other one, and it has numerous serrations that get
smaller towards the apex of the crown. The distal
cutting edge is almost straight, with serrations
increasing towards the middle. No nutritive groove
can be observed on the lingual surface of the root,
but this could be due to its poor state of preserva-
tion. Both mesial and distal cutting edges of the
main cusp are irregularly serrated. The serration of
the main cusp covers two thirds of its length. The
root is only partially preserved on both specimens.
Specimen FES 044 probably had a very lateral
position towards the commissure. It is interesting to
note the possible formation of a possible second-
ary serration in specimen FES 043 (Figure 4.1').
Remarks. At least three Eocene species of Galeo-
cerdo are known from the Eocene: G. latidens
Agassiz, 1843, G. aegyptiacus Stromer, 1905 and
G. eaglesomei White, 1955. Another possible spe-
cies, named Galeocerdo sp., from the Fayoum
Depression of Egypt, has also been described
(Case and Cappetta, 1990).
We found Galeocerdo eaglesomei to be the
species most similar to our material, with a compa-
rable serration that extends closer to the tip of the
cusp. The tall root is a morphological element also
found in our material. The size of the tooth and the
presence of the serration on the main cusp differ-
entiate our fossils from the species G. latidens.
Galeocerdo aegyptiacus from the Eocene of the
Fayoum Depression (Egypt) differs from our mate-
rial in being smaller in the basal-apical direction
and with a more convex mesial edge. Galeocerdo
sp. is similar to our specimens, but it has a smaller
serration on the lateral heels. This unnamed spe-
cies is considered to be distinct from G. latidens, G.
eaglesomei and G. aegyptiacus (Underwood et al.,
2011). Due to the fragmentary status of the fossils
we have at our disposal, we keep our assignment
to Galeocerdo cf. eaglesomei.
The species Galeocerdo eaglesomei can also
be encountered in the middle Eocene of Texas
(Westgate, 1989), Alabama (Maisch et al., 2014),
the upper Eocene of Mexico (Gonzales Barba,
2003), the Eocene of Nigeria (White, 1955) and
Egypt (Underwood et al., 2011).
Ciobanu (2002) documented Galeocerdo (as
G. latidens) from the Eocene of Turnu Roșu, Roma-
nia.
Incertae subfamiliae
Genus PHYSOGALEUS Cappetta, 1980
Physogaleus secundus Winkler, 1876
Figure 4.5-11
Material. Three lateral teeth (FES 050, 051, 052)
and one antero-lateral tooth (FES053).
Description. The lateral teeth measure only 6 to 8
mm mesio-distally and 4 to 5 mm in the apical-
basal direction. The main cusp is inclined distally
and has a triangular shape. The mesial side of
each tooth is long, almost straight, slightly convex
with two very irregular cusplets. The distal edge is
shorter and continues with two strong and triangu-
lar cusplets that are bent distally more than the
main cusp. The teeth are labio-lingually com-
pressed. On the lingual face the enameloid covers
about half of the root height and forms a straight
crown-root contact, while on the labial side the con-
tact has an arched shape. The lingual side of the
root has a deep central furrow while the labial side
has only some dispersed foramens.
The antero-lateral tooth measures 6.5 mm
mesio-distally and 5.5 mm in the apical-basal
direction. The main cusp is narrower than that of
the lateral teeth. The mesial edge is straight but
incomplete at its base. The distal edge is shorter
than the mesial one but the distal cusplets are
smaller and not as well outlined as they are in the
lateral teeth. The root has a much more pro-
nounced lingual protuberance than that of the lat-
eral teeth and the central furrow is very deep.
Remarks. Four Eocene species of Physogaleus
are known: P. te r ti us Winkler, 1874, P. latecuspida-
tus Müller, 1992, P. secundus Winkler, 1876 and P.
americanus Case, 1994. The species P. latecuspi-
datus is easily differentiated from P. secundus by a
wider main cusp, especially towards the tip of the
crown, hence its very illustrative name. Physoga-
leus americanus is a much smaller species with
teeth of around 3.0 mm and with a distinct cusp on
the mesial side. Physogaleus tertius lacks cusplets
in the lateral teeth; it only has a wavy lateral edge
that Case (1994 p. 121) considered an “unerupted
ridge”. We acknowledge another poorly known
species, P. rosehillensis Case and Borodin, 2000,
described from a single tooth. Taking into account
the limited description and the poor material, we
will not consider P. rosehillensis further, as it lacks
sufficient differentiating characters.

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
8
The morphology of our specimens is very sim-
ilar to one of the specimens figured by Cappetta
(2012, fig. 297), except for the labio-lingual com-
pression, which is more pronounced in our teeth.
We feel confident to assign our specimens to
Physogaleus secundus.
Physogaleus secundus is common in the
Eocene of Europe and Africa. The species is also
found in the lower-middle Eocene of Denmark
(Carlsen and Cuny, 2014), the Lutetian of Belgium
(Eckhaut and De Schutter, 2009), the Ypresian of
Morocco (Noubhani and Cappetta, 1997) and the
Eocene of Romania (Ciobanu, 2002; Dica, 2006).
Genus ABDOUNIA Cappetta, 1980
Abdounia sp.
Figure 5.12-13
Material. One anterior (FES 056) and one lateral
tooth (FES 057).
Description. The teeth measure only 7.4 mm
(FES 057) and 4.5 mm (FES 056) high in the api-
cal-basal direction. The main cusp is triangular with
a straight contact between the crown and the root
on the labial side. The lingual side is convex. The
lateral tooth has some very faint folds at the base
of the crown, on the labial side. The cusplets on
both teeth are triangular, divergent and not sepa-
rated by the crown on the labial side.
Remarks. Based on the described morphology we
assign our material to Abdounia sp. as it resembles
closely the description of the genus in Cappetta
(2012, p. 308). The scarce material and the imper-
fect preservation of the specimens prevent us from
assigning a species. The genus Abdounia is com-
mon in the Eocene of Africa, Europe and Americas
with at least seven valid species.
Order LAMNIFORMES Berg, 1958
Family LAMNIDAE Müller and Henle, 1838
Genus MACRORHIZODUS Glikman, 1964
Macrorhizodus praecursor (Leriche, 1905)
Figure 5.1-4
Material. Two teeth (FES no 041 and 042).
Description. The morphology of the specimens
indicates a lateral position. The specimen FES 041
is 49 mm high and 49 mm wide while specimen
FES 040 is 40 mm high and 41 mm wide. Both
teeth have a triangular crown that is inclined
slightly distally. The base of the crown is continued
by a heel on both sides. The heels are raised on
both sides of the tooth forming an almost straight
cusplet. The contact of the crown with the root fol-
lows a straight line. The root lobes are slightly
asymmetrical; the distal lobe has a sub-rectangular
outline, and the mesial lobe is pointed and a little
bit elongated. The specimen FES 041 has a visible
central foramen on the lingual face of the root.
Remarks. Two species and a subspecies of Mac-
rorhizodus are presumed to be present in the
Eocene: M. praecursor (Leriche, 1905), M. prae-
cursor americanus (Leriche, 1942) and M. nolfi
Zhelezko in Zhelezko and Kozlov, 1999. The differ-
ences are based on the outline of the crown, the
outline of the root lobes and the presence of so-
called heel bumps (or vestigial cusplets), respec-
tively.
Malyshkina and Ward (2016) consider that the
presence of high lateral cusplets (or heels) sepa-
rates the species Macrorhizodus nolfi from M.
praecursor, but they note that the characteristics
that separate the two species are unclear. Com-
pared with M. praecursor (illustrated by Cappetta,
2012, figure 207 A-P), M. nolfi (as figured in
Zhelezko and Kozlov, 1999, pl. 28, figs. 3a, 3b, 4a,
4b) has fully formed lateral cusplets.
Observations using high-resolution images of
these cusplets of the holotype (no. KP OF 15478/
41 DSM) and of the paratype (KP OF 15478/43
DSM) indicate that these cusplets are high and tri-
angular. We also observed that the cusplets are
positioned towards the margin in Macrorhizodus
praecursor and are closer to the main cusp in M.
nolfi. The root in M. nolfi is clearly higher than in
the specimen of M. praecursor figured by Leriche
(1906, pl. 16, figs. 12 and 12a), but similar to the
specimen of M. praecursor figured by Cappetta
(2012). Ignored for a long time, the existence of the
sub-species M. praecursor americanus has been
re-evaluated in recent decades and is mentioned
only rarely in the literature. However, we consider
the differences between M. praecursor and M.
praecursor americanus to be too subtle and rather
unclear. Cappetta (2012) regards M. praecursor
americanus as valid and specific to Priabonian. We
assign our specimens to Macrorhizodus praecur-
sor due to the low development of the lateral heels
and have decided not to assign the material to the
sub-species level.
Family MITSUKURINIDAE Jordan, 1898
Genus STRIATOLAMIA Glikman, 1964
Striatolamia macrota (Agassiz, 1843)
Figure 5.5-7’, 10-11
Material. Three anterior teeth (FES 068, 069, 070)
and five lateral teeth (FES 058, 059, 067, 091,
103).
Description. The anterior teeth measure up to 46
mm apico-basally and up to 21 mm mesio-distally.
The teeth are straight or trend slightly distally. The
labial face is strongly convex in the lower third of

PALAEO-ELECTRONICA.ORG
9
FIGURE 5. Fish fossils from Luna de Sus. 1-2, Macrorhizodus praecursor (FES 041). 3-4, (FES 042). 5-7', Striatola-
mia macrota (FES 068). 10-11, FES 067. 8-9, Otodus (Carcharocles) sp. (FES 071). 12-13, Abdounia sp. (FES 056).
14-16, Hypotodus verticalis (FES 061). 1, 3, 5, 9, 10, 12, 14 lingual views. 2, 4, 6, 8, 11, 13, 15, labial views. 6, 16, dis-
tal views. 7' detail of the cutting edge. Scale bars equal 5 mm (1-4, 12, 13) and 10 mm (5-11, 14-16).

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
10
the tooth, while the upper two thirds is flat. The lin-
gual side is strongly convex and has faint vertical
striations mostly in the lower half of the crown. The
cutting edges are sharp and continuous but do not
reach the base of the crown. One pair of very short
but sharp lateral cusplets is present in two of the
teeth. The cusplets do not reach the crown and are
more pronounced on the labial side. The root lobes
are long and pointed, with the exception of one
specimen, whose distal lobe is more flattened. The
lingual furrow is barely visible.
The lateral teeth have a main cusp with a dis-
tal inclination. The mesial edge is slightly convex in
the upper third and has a distal cutting edge that is
slightly concave in the lower third. The lingual face
of the crown is flat and smooth while the labial face
is slightly convex. The lingual face is ornamented
with very weak, parallel striations. These striations
extend only as far as the middle part of the crown
in the apical-basal direction. In some specimens
the striations are completely absent. The root lobes
are rounded with the mesial lobe being the longest.
The cusplets are well differentiated from the main
cusp and have a rounded-triangular shape.
Remarks. Two species of Striatolamia are known,
S. striata Winkler, 1876 and S. macrota Agassiz,
1843. S. striata is known only from the Paleocene
(Müller, 1992; Dutheil, 1992; Smith et al., 1999;
Moreau and Mathis, 2000), and it appears
restricted to this age (Cappetta, 2012). Striatolamia
macrota is very common in Ypresian, Lutetian
(Cappetta, 2012) and Bartonian (Zhelezko and
Kozlov, 1999). The characters we described and
the geological age clearly indicate that these teeth
are from Striatolamia macrota. The same features,
including the faint striations of the adult specimens,
have been reported by Zhelezko and Kozlov
(1999) and Cunningham (2000).
Family OTODONTIDAE Glickman, 1964
|Genus OTODUS Agassiz 1843 (sensu Cappetta,
2012)
Otodus (Carcharocles) sp.
Figure 5.8-9
Material. One fragmented tooth (FES 071).
Description. The root measures 51 mm wide and
both lobes are rounded. A single strong, wide and
unevenly serrated cusplet is present on either side
of the central cusp.
Remarks. The morphology agrees with that of Oto-
dus (Carcharocles) auriculatus, a large shark from
the middle Eocene, but the species could not be
positively determined due to the lack of a main
cusp.
Family ODONTASPIDIDAE Müller and Henle,
1839
Genus HYPOTODUS Jaekel, 1895
Hypotodus verticalis Agassiz, 1843
Figure 5.14-16
Material. One tooth (FES 061).
Description. The tooth has a triangular cusp with a
broad base and smooth labial and lingual sides.
The lingual side is convex, while the labial one is
flat and has a medial vertical ridge. In distal view
the crown is almost straight. Only one pair of cus-
plets is present on the sides of the crown from
which they are separated by a rounded notch. The
root lobes are moderately long and have pointed
ends. The distal lobe is shorter than the mesial
one. The lingual protuberance of the root is sharply
outlined and bears a central, pronounced furrow.
Remarks. Although somewhat similar in general
morphology, Hypotodus clearly differs from
Jaekelotodus by lacking the enameloid that covers
the upper part of the root lobes. Also, the labial
side of Hypotodus is convex to a certain extent,
while the labial side of Jaekelotodus is flat and has
a clear basal depression.
The validity of this genus has been widely dis-
puted. Over the years, authors such as Casier
(1946), Gurr (1962), Herman (1977) and Ward
(1988) misleadingly illustrated teeth belonging to
other taxa but which they attributed to Hypotodus.
Their opinions often changed the allocation of its
sole species H. verticalis, from Hypotodus to other
genera, hence invalidating the genus. Hypotodus
has since been re-considered as valid, as enough
characters have been found, and this made it pos-
sible to distinguish it from other genera (for further
discussion on this subject see Cappetta and Nolf,
2005, p. 244-246).
Genus JAEKELOTODUS Menner, 1928
Jaekelotodus robustus (Leriche, 1921)
Figure 6.1-13
Material. Three anterior teeth (FES 062, 063, 064)
and two lateral teeth (FES 060, 093).
Description. The teeth have a well-developed tri-
angular crown that is slightly inclined distally. Both
faces of the crown are smooth. The lingual face is
strongly convex while the labial one is flat. At the
base of the labial face is a central depression. On
this same side the enameloid descends from the
central depression and covers the upper portions
of the root lobes. On the lingual side, the contact
between the crown and the root is marked by a well
defined neck. The sharp cutting edge reaches the
base. A sharp cusplet occurs on either side of the

PALAEO-ELECTRONICA.ORG
11
FIGURE 6. Fish fossils from Luna de Sus. 1-3, Jaekelotodus robustus (FES 063). 4-8, (FES 065). 9-10, (FES 093). 11-
13, (FES 060). 1, 4, 9, 11, lingual views; 2, 5, 10, 12, labial views. 7, basal view. 8, apical view. 13, distal view. Scale
bars equal 10 mm (1-8) and 5 mm (9-13).

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
12
crown and each cusplet is inclined towards the
main cusp. In the mesial or distal view it is
observed that the cusplets are not following the
same plan as the main cutting edge, but instead
are lingually inclined. In specimen FES 063 it is
observed the tendency towards the formation of a
secondary pair of cusplets partially separated from
the main pair. This tendancy is also observed for
the mesial cusplet of FES 093.
Remarks. At least two Eocene species of Jaekelo-
todus are known, J. trigonalis and J. robustus.
However, Cappetta (2012) lists also J. londonensis
Zhelezko, 1994 and an impressive number of sub-
species for J. trigonalis, all from Kazakhstan.
Teeth of Jaekelotodus are easily distinguished
from those of other genera of the same age by the
stout triangular crown with the central labial
depression and the medium sized, hook-shaped
cusplets. The teeth of J. robustus can be recog-
nized based on their single pair of lateral cusplets
that are shorter compared to J. trigonalis and J.
londonensis. Also, the root of J. robustus teeth has
a furrow that is more pronounced than that of J.
trigonalis (Cappetta and Nolf, 2005).
Superorder BATOMORPHII Cappetta, 1980
Order MYLIOBATIFORMES Compagno, 1973
Superfamily MILIOBATOIDEA Compagno, 1973
Family MYLIOBATIDAE Bonaparte, 1838
Subfamily MYLIOBATINAE Bonaparte, 1835
Genus MYLIOBATIS Cuvier, 1816
cf. Myliobatis sp.
Figure 7.1-3
Material. One almost complete dental plate (FES
074).
Description. The specimen measures 32.1 mm
labial-lingualy and has a maximum width of 29 mm.
The width-to-length ratio of the medial teeth is
5.3:1, while for the lateral ones it is 0.8:1 for the
inner row and 0.6:1 for the outer row. A slight cur-
vature is present in the labio-lingual direction. The
eight medial teeth are the widest teeth in the plate
and are slightly arched in occlusal view. All the
teeth are hexagonal. The medial teeth are framed
by two lateral roaws on each side. Longitudinal stri-
ations occur on the occlusal surface. The root is
partially filled with sediment but is no more than 2
mm high and of a polyaulacorhize type. The root
laminae are irregularly shaped, and the grooves
between them are wider than the laminae.
Remarks. The genus is described by Bigelow and
Schroeder (1948) and Cappetta (2012) as having a
dental plate consisting of seven rows of teeth, the
medial file being the widest, and each tooth being
hexagonal shaped.
The number of fossil Myliobatis species is
amazingly high. For instance, there are forty-five
species described for the Eocene (Hovestadt and
Hovestadt-Euler, 2013). However, Cappetta (1987)
listed only six valid species from the entire Paleo-
cene and later reduced this number to only five
(Cappetta, 2012). We note that an extensive com-
parative study by Hovestadt and Hovestadt-Euler
(2013) of the dental morphology of the extant rep-
resentatives of the Myliobatinae subfamily showed
that teeth morphological variation is very high even
within the same species. For example, several
species of Myliobatis show variation in the number
of rows and in the width/length ratios. Also there is
strong evidence that tooth morphology is cor-
related with dietary preference or prey availability
within a given species (Hovestadt and Hovestadt-
Euler, 2013, p. 22). As the specimen only partially
fulfills the genus diagnostic characters, we
assigned the specimen to cf. Myliobatis sp.
Family AETOBATIDAE Agassiz, 1958
Genus AETOBATUS Blainville, 1816
cf. Aetobatus sp.
Figure 7.4-7
Material. Two fragmentary upper teeth (FES 072,
073).
Description. The teeth are medium-sized (37 mm
width and 9 mm length for the figured specimen),
laterally arched and with a wavy outline, smooth
occlusal surface and roots divided longitudinally by
ridges and grooves, which continue from underside
towards backside.
Remarks. The antero-posterior arched, wavy or M-
shaped teeth from the Myliobatinae that have a
posterior displaced root are typically assigned to
Aetobatus. As with the genus Myliobatis, the lack
of data regarding the variability of tooth morphol-
ogy in extant genera and the poor knowledge of
genera such as Aetomylaeus led to erroneous
assignations.
Myliobatinae indet. 1
Figure 7.8-12
Material. One fragmented dental plate (VT459
BBUPSM).
Description. The fragment measures 28 mm
labial-lingually and has a maximum width of 46
mm. Only a single tooth is complete. The
width:length ratio of the complete medial teeth is
5:1. Only a small fragment of a lateral tooth is pres-
ent. The root is heavily worn and the laminae are
razed. Interestingly, the root contains deep grooves
between the teeth.

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13
FIGURE 7. Fish fossils from Luna de Sus. 1-3, cf. Myliobatis sp. (FES 074). 4-7, cf. Aetobatus sp. (FES 072). 9-10,
FES 093. 8-12, Myliobatinae indet. (1 VT 459). 13-15, Myliobatinae indet. 2 (FES 048). 16-18, Rajidae indet. (FES
046). 19-21, Batomorphii indet. (FES 086). 22-24, (FES 087). 6, 11, lingual views. 7, 10, labial views. 2, 5, 9, 13, 17,
21, 24, basal views. 1, 4, 8, 15, 16, 19, 22, occlusal views. 3, 12, 14, 18, 20, 23, lateral views. Scale bars equal 5 mm.

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
14
Remarks. Similar deep grooves are present on a
fragment assigned by Dica (2006) to Myliobatis sp.
1; this fragment was from the same formation, but
a different locality (Căpușul Mic).
Myliobatinae indet. 2
Figure 7.13-15
Material. One tail sting (FES 048).
Description. The specimen is a fragment of a tail
sting. The dorsal side is smooth. The ventral side
has a longitudinal ridge along its entire length. The
distal quarter of the sting exposes serrations on
both sides. The exact shape of these serrations
could not be determined due to the poor state of
preservation.
Remarks. Since the tail stings of rays lack diag-
nostic characteristics according to Hovestadt and
Hovestadt-Euler (2013), it could not be determined
below the subfamily level.
Superfamily DASYATOIDEA Whitley, 1940
Family DASYATIDAE Jordan, 1888
Genus DASYATIS Rafinesque, 1810
Dasyatis cf. jaekeli (Leriche, 1905)
Figure 8.1-4
Material. One tooth (FES 095).
Description. The tooth is 1.8 mm wide and 1.4
mm high. Both occlusal and labial surfaces are
covered by a network of alveoli and crests, while
the lingual face is smooth. The tooth is deeply worn
in the mesial direction of the occlusal face. The
general shape of the occlusal surface is hexago-
nal. On lateral profile, one can notice a depression
of the lingual side. The root is short and it has two
lobes with triangular ends. The central foramen is
barely visible between the root lobes.
Remarks. Tooth morphology of the genus is highly
variable, and it is complicated by the gynatric het-
erodonty, as breeding males have cuspidate teeth
and non-breeding males and all females have
rounded ones (Kajiura and Tricas, 1996; Cappetta,
2012). The above described characters likely indi-
cate a female tooth or a non-breeding male belong-
ing to Dasyatis. It differs slightly from D. jaekeli,
based on illustrations and descriptions from Ler-
iche, (1905), Case (1994) and Ciobanu (2002), as
it has a shallower lingual depression and the cen-
tral occlusal ridge is less obvious. For this reason
we treat the specimen here in an open nomencla-
ture.
Order RHINOPRISTIFORMES Naylor et al., 2012
Family RHINOBATIDAE Müller and Henle, 1838
Genus RHINOBATOS Linck, 1790
Rhinobatos cf. steurbauti Cappetta and Nolf, 1981
Figure 8.5-8
Material. Two teeth (FES 096, 097).
Description. The teeth are very small. The widest
crown measures 1.3 mm. The teeth have a general
globular shape with a very weakly marked trans-
versal ridge on the occlusal surface. A central,
elongated uvula is visible lingually. A pair of lateral
uvulae is present on the sides of the central uvula.
The lateral uvulae are shorter than the central one
and do not diverge. All uvulae have a rounded
basal end. The root is missing in both teeth.
Remarks. Two species of Rhinobatos are known
from the Eocene: R. bruxelliensis Jaekel, 1894 and
R. steurbauti Cappetta and Nolf, 1981. Rhinobatos
bruxelliensis has pointed and more divergent uvu-
lae while R. steurbauti has uvulae that are rounded
and less or non-divergent (Cappetta and Nolf,
1981, p. 96). Based on these differences, we
assign the specimens to R. cf. steurbauti. Rhinoba-
tos steurbauti is rarely reported but is present in
the middle Eocene of France (Cappetta and Nolf,
1981), England (Bone et al., 1991) and Uzbekistan
(Case et al., 1996).
Family PRISTIDAE Bonaparte, 1838
Genus PRISTIS Linck, 1790
Pristis sp.
Figure 8.9-13
Material. One incomplete rostral spine (FES 003).
Description. The rostral spine measures 48.0 mm
in apical-basal direction, 12.0 mm in anterior-pos-
terior direction and 4 to 6 mm dorsal-ventrally. The
spine is dorso-ventrally compressed with a
rounded anterior margin, and with a shallow longi-
tudinal groove along the posterior margin. Both
apical and basal ends of the spine are broken. The
apical surface has asymmetrical margins along the
longitudinal groove.
Remarks. Similar to the Myliobatidae, numerous of
fossil pristids have been described in the last few
centuries. Many of these species were described
from isolated rostral spines. Twenty-six species
belonging to four genera (Propristis Dames, 1883;
Mesopristis Farres, 2003; Pristis Linck, 1790; and
Anoxypristis White and Moy-Thomas, 1941) have
been described from the Eocene. Propristis has a
unique morphology, with rostral spines nearly as
long as high (Cappetta 2012, p. 396). The newest
genus, Mesopristis, was set in synonymy with
Anoxypristis by Cappetta (2012), but without expla-
nation. However, the rostral spines of Mesopristis
lack a posterior groove. Anoxypristis can be
excluded for the same reason. Pristis is the only
genus that has a posterior longitudinal groove.
Within the respective genus there are numerous
Eocene species but the species cannot be deter-

PALAEO-ELECTRONICA.ORG
15
FIGURE 8. Fish fossils from Luna de Sus. 1-4, Dasyatis cf. jaekeli (FES 095). 5-8, Rhinobatos cf. steurbauti (FES
096). 9-13, Pristis sp. (FES 003). 14, Labridae indet. (FES 057). 15-19, Phacodus cf. punctatus (FES 012-040). 1, 5,
15, labial views. 4, 16, lingual views; 8, 14, 18, occlusal view. 9, dorsal view. 10, ventral view. 11, posterior view. 12,
anterior view. 13, apical view. 2, 6, 7, 19, lateral views. 18', detail of the occlusal surface. Scale bars equal 1 mm (1-8),
5 mm (14-19) and 10 mm (9-13).

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
16
mined from the material at hand. Observation on
extant material belonging to Pristis, made by one of
us (NT), indicate a high degree of variability in ros-
tral spine length along with the width and depth of
the posterior groove on the same specimen. Con-
sidering the above-mentioned facts the specimen,
can be assigned with confidence to Pristis sp.
Order cf. RAJIFORMES Berg, 1940
Family, genus et species indet.
Figure 7.16-18
Material. Two dermal spine bases (FES 046, 088).
Description. These generally oval dermal spines
have a truncated cone shape in lateral view. A
shallow concavity and some very fine radial stria-
tions are present at the center of the dorsal side, as
are some obvious concentric growth lines. The
basal surface is slightly convex.
Remarks. The assignation of these thorn bases is
challenging, as both Rajiformes and Myliobati-
formes include numerous families where these der-
mal thorns are present (Deynat, 1998). Moreover,
the local taphonomy may have reduced through
abrasion some of the finer surface details. It is
debatable if these thorns should be allocated to
Rajiformes. Extant species show that thorn mor-
phology can be variable on the same individual
depending on the part of the body they occur
(Gravendeel et al., 2002). Nevertheless, we found
that several Rajidae genera have similarly shaped
dermal denticles. In addition, we note that very
similar fossil material from the upper Eocene of
Germany has also been assigned to Rajiformes,
based on comparison with extant material (Hillmer
and Mundlos, 1981).
Superorder BATOMORPHII Cappetta, 1980
Batomorphii indet.
Figure 7.19-24
Material. Two dermal spine bases (FES 086, 087).
Description. These two specimens differ greatly
from the above-described material. Specimen FES
086 has a pentagonal outline in dorsal view and a
truncated cone shape from a lateral view. A deep
concavity is present at the center of the dorsal
side. Striations and growth lines were absent,
unlike the cf. Rajiformes specimens. The base is
completely flat. Specimen FES 087 (Figure 7.22-
24) also differs from the cf. Rajiformes specimens
and specimen FES 086 (Figure 7.19-21) by having
a rounded outline and a torus-shaped lateral out-
line. The basal face is convex. This specimen has
a concavity in the central raised portion of the dor-
sal surface.
Remarks. Specimen FES 087 is reminiscent of a
pearl-shaped tubercle as defined by Deynat (1998,
p.158).
Order PYCNODONTIFORMES Berg, 1940
Family incertae sedis
Genus PHACODUS Dixon, 1850
Phacodus cf. punctatus Dixon, 1850
Figure 8.15-19
Material. Twenty-nine teeth (FES 012 - 040).
Description. The teeth are generally oval or round
and measure 4 to 11 mm along the long axis. The
oral surface is covered by fine pits (Figure 8.18')
that penetrate the enamel deeply. The basal side of
the crown has a deep oval-shaped depression. In
some specimens, the edges of the teeth show dif-
ferently-sized and geometrically-shaped protru-
sions (Figure 8.15-16, 8.19).
Remarks. The generally oval or round shape and
the presence of the fine pits that cover the entire
occlusal surface of the tooth are characteristic
traits of the monotypic Phacodus punctatus. The
presence of Phacodus in the Paleogene is
unusual. Until recently, Phacodus was known only
from the Cretaceous (Dixon, 1850; Santos and
Figueiredo, 1988; Hooks et al., 2013). In Romania,
a single occurrence of Phacodus after the K/T
boundary was known only from specimens found a
few years ago in the upper Eocene of Turnu Roşu
(Ciobanu and Trif, 2014). A Phacodus tooth has
also been found at Căpuşul Mic (Căpuş Fm.), but it
was referred only as Pycnodontidae indet. (Dica,
2006, plate 15, figure 2).
The species is based on a fragmentary asso-
ciated dentition (holotype NHM PV OR 25829),
representing possibly a splenial fragment, found in
the Cenomanian of the UK. For a long time the
species and the genus were rejected, the holotype
specimen being considered just a very worn and
fragmentary pycnodont (Woodward, 1888, 1895).
However, the genus and species were finally
accepted as valid by Woodward (1909), who had
previously rejected them. Since then the genus has
been studied only rarely. Arambourg (1952) found
a vomer and two fragmentary splenials in the
Maastrichtian of Morocco (Ouled Abdoun). Based
on this material, he erected a new variety, Phaco-
dus punctatus var. africanus, due to a slight differ-
ence in the outline of the central teeth and in the
smaller size of the lateral teeth. Santos and
Figuiredo (1988) described Phacodus sergipensis
almost three years later, from the Turonian of Bra-
zil. The difference between this new species and P.
punctatus, based mainly on two additional lateral
rows, was rejected by other authors (e.g., Hooks et

PALAEO-ELECTRONICA.ORG
17
al., 2013) who argued that the variation of the lat-
eral number of rows in pycnodonts had been noted
in other species too. While we have assigned our
Romanian teeth to Phacodus cf. punctatus, micro-
morphology and future research may lead us to a
new conclusion.
Order PERCIFORMES Bleeker, 1859
Suborder LABROIDEI Bleeker, 1859
Family LABRIDAE Cuvier, 1817
Labridae indet.
Figure 8.14
Material. One upper pharyngeal tooth plate (FES
057).
Description. A fragmented pharyngeal plate was
collected having numerous rounded teeth with
diameters of 0.25 to 2.5 mm. The teeth are longest
at the center of the plate and decrease towards the
edges. The central teeth have a rough, irregular
surface while the lateral ones have a completely
smooth exterior. The teeth are arranged in regular,
almost parallel rows. Together, they form a com-
pact grinding surface. Note the considerable thick-
ness of the plate in occlusal-basal direction, at 14.0
mm. The plate is triangular in transversal section,
including the grinding surface.
Remarks. Although some authors consider Labri-
dae tooth plates non-diagnostic (e.g., Long, 1992),
others, such as Dica (2002), consider the morphol-
ogy and the arrangement of the teeth on the pha-
ryngeal plates to be indicative characters on the
genus level. An example of this is the lower pha-
ryngeal plate of Lachnolaimus multidens from the
Priabonian of Romania (Dica, 2002, fig. 3). The fig-
ured specimen compares well with the previously
described lower pharyngeal plate of L. multidens
from the Miocene of the Vienna Basin (Münster,
1846, figs. 5a-c). However, we consider that the
assignment of other upper pharyngeal plates from
Transylvania to species level, as L. multidens
(Dica, 2002, pl. 1, fig. 4; Ciobanu, 2013, figs. 1-6) is
only arbitrary as the lower plate is missing.
The assignment of our specimen to Labridae
is however valid, based on similar fossil upper den-
titions (Münster, 1846; Wainwright, 1987).
The Labridae family is known from Eocene,
primarily from associated skeletons from Monte
Bolca (Bannikov and Carnevale, 2010), but iso-
lated pharyngeal triturating plates from Eocene
Labridae have also been reported (Bellwood et al.,
2019).
DISCUSSION
Taxonomy
The fish assemblage from Luna de Sus
includes at least 17 species from as many genera;
these fish belong to 12 families and seven orders.
Seven of these genera (Physogaleus, Abdounia,
Macrorhizodus, Striatolamia, Otodus, Jaekeloto-
dus, Hypotodus) are extinct as is the entire order of
Pycnodontiformes. One genus (Heterodontus) and
four species (Galeocerdo cf. eaglesomei, Jaekelo-
todus robustus, Hypotodus verticalis and Rhinoba-
tos cf. steurbauti) are new for Romania. This
locality represents the second Cenozoic occur-
rence of the pycnodontiform Phacodus cf. puncta-
tus in Romania and Europe. The first reported
occurrence was the one from the upper Eocene of
Turnu Roșu (Ciobanu and Trif, 2014). The Labri-
dae material reported here represents the oldest
record of this family in Romania.
Taphonomy
The relative abundance of teeth and dermal
spines at this level puts it in contrast with other lay-
ers from the same locality. Only a single other layer
contains fish teeth, namely the Nummulites perfor-
atus lumachelle, from where a single fish tooth of
Striatolamia macrota has been recovered.
The general state of preservation of the teeth
is good, as evidenced by their sharp cutting edges,
fine lateral cusplets and mostly intact roots. This
mode of preservation indicates that the remains
were transported only minimally before final burial;
therefore, we consider the specimens to be
autochthonous or parautochthonous. All the teeth
have a reddish-brown color specific to the fossils of
the iron oxide-rich Căpuș Formation (Stoicovici and
Mureșan, 1964a, 1964b).
Paleoecology
The environment of sharks, rays and bony
fishes from the locality of Luna de Sus can be
reconstructed with some degree of approximation,
considering the habitat preferences of extant rela-
tives. Heterodontus is found in a wide range of
depths, from intertidal to 275 m (Bass et al., 1975;
Compagno, 2002). Rhizoprionodon is found both
inshore and offshore, and at depths from less than
1 m to about 200 m, with some species reaching
500 m depth (Compagno, 1984). A transition
towards a shallower reef habitat has been encoun-
tered in some species (Sorenson et al., 2014).
Galeocerdo inhabits a wide range of depths and
habitats, from intertidal to over 200 m depth, but

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
18
spends most of its time in less than 50 m of water
(Compagno, 1984; Vaudo et al., 2014). Aetobatus
is benthopelagic and typically occurs nearshore, in
1 to 60 m depth (Mundy, 2005; White and Last,
2016). Myliobatis is also benthopelagic and is con-
fined mostly to warm waters and shallow depths to
80 m (Love et al., 2005; Ebert and Stehmann,
2013). Dasyatis is demersal (to 40 m depth), with
most species preferring shallow water. Many
Dasyatis species are found inshore, in lagoons and
river-mouths (Struhsaker, 1969; Ebert and Steh-
mann, 2013). Rhinobatos lives in shallow inshore
estuarine to marine waters, to 100 m, but with
some species living as deep as 180 m (Ebert and
Stehmann, 2013). Extant Rhinobatidae live in
warm-temperate to tropical waters, but a few spe-
cies occur in deeper, cooler water offshore (Last
and Compagno, 1999). Representatives of Pristis
prefer shallow coastal waters and estuaries, typi-
cally in less than 10 m depth, but adults may be
found in the 100 m depth or more (Poulakis and
Seitz, 2004; Simpfendorfer, 2005; Ebert and Steh-
mann, 2013). Immature individuals are dependent
on shallow habitats, living especially around the
river-mouths (Simpfendorfer, 2005; Last et al.,
2016b). Some extant Pristis species have a prefer-
ence for freshwater habitats for nurseries (Peverell,
2005). The range for extant Pristidae is circumtrop-
ical (Nelson et al., 2016) and presumably the cli-
mate preference of the fossil Pristis was the same.
Labrids are generally shallow-water fishes, though
exceptionally, some species of Bodianus and
Decodon occur in deep water at 200 m or more
(Smith and Heemstra, 1986). Most Labridae live in
association with structure such as coral reefs or
rocky substrate in depths to 40 m (Allen et al.,
2003).
The presence of Phacodus cf. punctatus in
the Eocene of Luna is very unusual, as it is the only
pycnodont found in northwestern Transylvania. Lit-
tle is known about the ecological conditions of this
fish. Taking into consideration the associated fauna
from other Transylvanian occurrences (Ciobanu,
2002; Ciobanu and Trif, 2014), the warm, moder-
ately deep waters are also among its environmen-
tal preferences. The pycnodonts are very rare in
the northwestern portion of the Transylvanian
Basin as compared to their occurrence in Turnu
Roșu on the southern side of the basin. The pycno-
dont teeth are well represented in all three forma-
tions from Turnu Roșu (spanning the Ypresian-
Priabonian interval). The presence of a high num-
ber of teeth from this single species in Luna de Sus
indicates a well established population of pycno-
donts in an unspecified ecological niche that
allowed them to flourish among higher rank preda-
tors such as sharks.
The habitat preferences of modern relatives of
studied fishes (Figure 9) indicate a moderate water
depth of less than 200 m (more likely less than 100
m, as inferred by the preferred depth) in a warm
sea.
The knowledge of the middle Eocene fish
(documented by teeth) is still limited on the
regional level of Eastern Europe. We compared the
occurrences of taxa from Luna de Sus with those
from elsewhere in the region and with those from
other better-known localities. For this comparison
we only took into consideration the Chondrich-
thyes, given their numeric dominance at Luna de
Sus. We assessed the faunal diversity at this site
FIGURE 9. Depth range of extant equivalent taxa.

PALAEO-ELECTRONICA.ORG
19
with that of 13 sites from the European part of Rus-
sia, Hungary, Ukraine, Denmark, Germany, Bel-
gium, France, Egypt, and Morocco (Figure 10).
The data covers the early Lutetian through the late
Bartonian (Table 1).
We note a few genera that are probably more
adaptable and migratory and which are common
throughout the entire Tethys Sea and North-East
Atlantic coast, such as Striatolamia, Macrorhizodus
and the small Carcharhiniformes Abdounia and
Physogaleus (Dutheil, 1990; Udovichenko, 2006;
Eeckhaut and De Schutter, 2009; Udovichenko,
2009; Timircev and Popov, 2011; Diedrich, 2012).
The absence of Otodus from many sites of the
close-by region (Kocsis, 2002; Udovichenko, 2006;
Udovichenko, 2009; Malyshkina et al., 2013; Leder,
2013) is surprising, considering that this predator is
present in many localities of North-East Atlantic
coast and North Africa (Dutheil, 1990; Eeckhaut
and De Schutter, 2009; Adnet et al., 2010; Under-
wood et al., 2011; Diedrich, 2012; Carlsen and
Cuny, 2014). It is good to note that diversity on the
genus level in the North Sea Basin, Paris Basin
and in the south of the Tethys Sea is significantly
higher than in Eastern Europe. The presence of
Pristidae and small Carcharhinidae in southwest-
ern Morocco and in the Midawara Formation of
Egypt (Adnet et al., 2010), where tropical condi-
tions were well established, also indicates the exis-
tence of a warm habitat for Luna de Sus. Another
evidence to support this hypothesis is the lack of
deep, cold-water taxa such as Centrophorus,
Chlamydoselachus, Coupatezia, Echinorhinus and
Hexanchiformes that are present in other sites
(Dutheil, 1990; Eeckhaut and De Schutter, 2009;
Malyshkina et al., 2013; Carlsen and Cuny, 2014)
that indicates a shallow depth and warmer condi-
tions in Luna de Sus.
The climate reconstructions based on palynol-
ogy (Petrescu and Balintoni, 2003) confirms a
warm climate. This is supported further by recon-
structions using oxygen and carbon isotopes mea-
sured on Nummulites perforatus collected at the
same locality of Luna de Sus, on “Pavel Brook”.
For the interval involving the fish fauna, a mean
paleotemperature is calculated at 26 °C, while for
the levels immediately above this interval a slightly
FIGURE 10. Map of central and eastern Europe with similar localities.

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
20
TAB LE 1 . Orders and genera of Chondrichthyes documented in the Tethys Sea region. The numbers within brackets in
the first row refer to localities: (1) Luna de Sus, Romania, lowermost Bartonian (this article); (2) Gradizhsk, Ukraine,
Lutetian-Bartonian (Udovichenko, 2006); (3) Osinovo, Ukraine, lower Lutetian (Udovichenko, 2009); (4) Crimea, Rus-
sia, Gornyi Luch site, middle Eocene, most probably, Bartonian (Malyshkina et al., 2013); (5) Crimea, Russia, Bakh-
chisarai site, middle Eocene, most probably, Bartonian (Malyshkina et al., 2013; Leder, 2013); (6) Morozovsk, Russia,
middle Eocene (Timircev and Popov, 2011); (7) Gerecse, Hungary, middle Eocene (Kocsis, 2002); (8) Trelde Næs,
Denmark, upper Ypresian to middle Lutetian (Carlsen and Cuny, 2014); (9) Dalum and Osteroden, Germany (Diedrich,
2012); (10) Oosterzele, Lutetian, Belgium (Eeckhaut and De Schutter, 2009); (11) Paris Basin, the Lutetian-Bartonian
interval only (Dutheil, 1990); (12) Midawara Fm., Egypt, Lutetian (Underwood et al., 2011); (13) Dakhla, Morocco
(Adnet et al., 2010). (*) genera that are valid, but have an unclear/uncertain use in the Eocene; (**) Otodus is used
sensu Cappetta, (2012) and reunites Otodus and Carcharocles.
Orders Genera
(1) Luna ROMANIA
(2) Gradizhsk UKRAINE
(3) Osinovo UKRAINE
(4) Gornyi RUSSIA
(5) Bakhchisarai RUSSIA
(6) Morozovsk RUSSIA
(7) Gerecse HUNGARY
(8) Trelde Næs DANEMARCK
(9) Dalum GERMANY
(10) Oosterzele BELGIUM
(11) Paris Basin FRANCE
(12) Midawara EGYPT
(13) Dakhla MOROCCO
HETERODONTIFORMES Heterodontus ––– – ––
ORECTOLOBIFORMES Chiloscyllium ––
Eostegostoma – –
Hemiscyllium –
Pararhincodon –
Nebrius – ––––
Palaeorhincodon ––
Protoginglymostoma –
CARCHARHINIFORMES Abdounia ––– –– ––––
Carcharinus ––
Crassescyliorhinus –
Foumtizia – –
Galeocerdo – ––– –––––
Galeorhinus – –– –
Hemipristis ––
Iago – ––
Leptocharias –
Megascyliorhinus –
Misrichthys –
Pachygaleus ––
Paragaleus –
Physogaleus –––––– – ––––
Premontreia –
Rhizoprionodon ––– –––
Scyliorhinus –– –
Sphyrna –
Tri aki s – –
Triakidae nov. genus –

PALAEO-ELECTRONICA.ORG
21
LAMNIFORMES Alopias – –– – ––
Anomotodon –––
Carcharias –––– –
Carcharodon * –
Cretalamna – –
Brachycarcharias – –– –
Hypotodus ––– –––
Isurolamna –––– ––––
Isurus * – –
Jaekelotodus ––– – ––––
Macrorhizodus –––– – ––– ––
Mitsukurina –
Otodus sensu
Cappetta, 2012
– – – ––––––
Odontaspis –– – ––––
Palaeohypotodus –
Parotodus –
Sylvestrilamia ––
Striatolamia –––––––––––
Turania –
Usakias –––
Woellsteinia –
Xiphodolamia ––– –
HEXANCHIFORMES Chlamydoselachus –
Heptranchias –
Hexanchus – – –
Notorhynchus – – –
Weltonia –
SQUATINIFORMES Squatina –– – – –
SQUALIFORMES Centrophorus –
Echinorhinus – –
Isistius – – – –
Squalus –
MYLIOBATIFORMES Aetobatus ––– – ––––
Archaeomanta –– –
Aturobatis –
Burnhamia –– – –– –
Coupatezia – – ––
Orders Genera
(1) Luna ROMANIA
(2) Gradizhsk UKRAINE
(3) Osinovo UKRAINE
(4) Gornyi RUSSIA
(5) Bakhchisarai RUSSIA
(6) Morozovsk RUSSIA
(7) Gerecse HUNGARY
(8) Trelde Næs DANEMARCK
(9) Dalum GERMANY
(10) Oosterzele BELGIUM
(11) Paris Basin FRANCE
(12) Midawara EGYPT
(13) Dakhla MOROCCO
TABLE 1 (continued).

TRIF, CODREA, & ARGHIUȘ: BARTONIAN FISH FROM TRANSYLVANIA
22
cooler tendency was recorded, calculated at 23-24
°C (Bartholdy et al., 2000).
CONCLUSIONS
The richness of the middle Eocene fish fauna
of the Luna de Sus in the northwestern portion of
the Transylvanian Basin is significant, with at least
17 species belonging to as many (17) genera. The
vast majority of the teeth found here belongs to
Chondrichthyes and apparently originate from
large-to-medium-sized fish, but this could be a con-
sequence of the sampling bias. As revealed by the
teeth taphonomy, the layer of sediments the teeth
were recovered from most likely accumulated
under low energy depositional conditions, and we
therefore consider these fossils to be either
autochthonous or parautochthonous. The faunal
Dasyatis –––––––
Garabatis –
Gymnura –––
Heterotorpedo –
Himantura * ––
Hypolophodon –
Jacquhermania –
Leidybatis –––
Lophobatis –
Mobula –
Myliobatis ––– – ––––
Neotrygon –
Pseudoaetobatus –
Rhinoptera –
Ouledia ––
Taeniura –
RHINOPRISTIFORMES Anoxypristis –––
Platyrhina –
Platyrhinoidis –
Pristis – ––––
Propristis ––
Rhinobatos – –– –– –
Rhynchobatus ––––
PRISTIOPHORIFORMES Pristiophorus –
TORPEDINIFORMES Narcine ––
SELACHII INC. SEDIS Odontorhytis ––
Total number of genera 15 18 17 7 9 15 7 32 17 41 33 36 38
Genera in common with Luna 10 13 4 4 8 6 8 6 14 12 10 11
Orders Genera
(1) Luna ROMANIA
(2) Gradizhsk UKRAINE
(3) Osinovo UKRAINE
(4) Gornyi RUSSIA
(5) Bakhchisarai RUSSIA
(6) Morozovsk RUSSIA
(7) Gerecse HUNGARY
(8) Trelde Næs DANEMARCK
(9) Dalum GERMANY
(10) Oosterzele BELGIUM
(11) Paris Basin FRANCE
(12) Midawara EGYPT
(13) Dakhla MOROCCO
TABLE 1 (continued).

PALAEO-ELECTRONICA.ORG
23
analysis of the northeast Atlantic coast and of the
Tethys Sea reveals in part a common elasmo-
branch fauna with a variable number of common
genera. These common genera are mostly lamni-
forms (Hypotodus, Jaekelotodus, Macrorhizodus,
Otodus, Striatolamia) that were opportunistic pred-
ators of more open water or at least with broader
habitat preferences.
Four species and one genus are reported
here for the first time from the Eocene deposits of
Romania. Aside from them, one of the most import-
ant occurrences in these deposits is Phacodus cf.
punctatus. This second report in the northwestern
Transylvanian Basin confirms its survival of the K/
Pg boundary.
ACKNOWLEDGMENTS
We thank both anonymous reviewers for their
critical reading of the manuscript and for their valu-
able suggestions. The authors are very grateful to
J. Seitz for not only reviewing the English language
but also for the suggestions and references he
sent. The authors also would like to thank A. Fuciu
for the help in translation of the early version of the
manuscript. The manuscript greatly benefited from
the discussions with P. de Schutter and T. Malysh-
kina.
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