, APRIL 2012, 63, 2, 175—178 doi: 10.2478/v10096-012-0014-6
Incremental growth and mineralogy of Pannonian (Late
Miocene) sciaenid otoliths: paleoecological implications
, MIKLÓS KÁZMÉR
, MARIANN BOSNAKOFF
, TAMÁS VÁCZI
and LAJOS KATONA
Department of Palaeontology, Eötvös Loránd University, Pázmány Péter sétány 1/c, H-1117 Budapest, Hungary; firstname.lastname@example.org
Division of Climate and Environmental Physics, University of Bern, Sidlerstrasse 5, CH-3012 Bern, Switzerland; email@example.com
Institute for Geological and Geochemical Research, Hungarian Academy of Sciences, Budapest, Hungary
Hungarian Natural History Museum, Budapest, Hungary
Department of Mineralogy, Eötvös Loránd University, Budapest, Hungary
Natural History Museum of Bakony Mountains, Zirc, Hungary
(Manuscript received September 14, 2011; accepted in revised form January 4, 2012)
Abstract: Ontogenetic age and body dimensions were studied on three extremely well-preserved sciaenid fish otoliths
from sublittoral marls of Lake Pannon from Doba, Bakony Mts, Hungary. Macroscopic and microscopic observations
offered clear evidence for the preservation of the genuine structural characteristics, for instance the bipartite incremental
features. Ontogenetic ages were assigned for the three specimens as 16, 7 and 6 years by counting the annuli of the sagittae.
Analytical results prove that the original aragonitic mineralogy has been preserved making them, and probably other Late
Miocene teleost fossils, suitable for future microchemical analysis to reconstruct the past physicochemical environment.
Key words: Lake Pannon, paleoecology, littoral environment, sagitta, cathodoluminescence, Raman microspectroscopy,
Lake Pannon, a long-lived, giant lake, was a Late Miocene
successor of the disintegrated Paratethys Sea (Kázmér 1990).
Extensive freshwater discharge by the paleo-Danube and pa-
leo-Tisza rivers into the Pannonian basins (Kuhlemann et al.
2002) altered the composition of the water-body, reducing the
marine-derived fauna of the Late Sarmatian brackish-water
sea into a fully endemic, lacustrine fauna. While the mollusc
fauna is well-documented (e.g. Müller et al. 1999 and refer-
ences therein), the fish fauna is poorly known. Mostly oto-
conia (mineral material from the inner ear of any vertebrate;
see Carlström 1963) from sciaenids have been found relatively
frequently (e.g. Brzobohatý & Pana 1985; Bosnakoff 2008).
The endolymphatic organ of Sciaenidae, like in other te-
leost fishes, contains three pairs of small carbonate stones,
so-called otoliths. The largest of them are called sagittae, lo-
cated in the sacculus of the membranous labyrinth. Although
both incremental (e.g. Woydack & Morales-Nin 2001) and
microchemical information (e.g. Carpenter et al. 2003) of
fossil otoliths proved to provide unique information about
the coeval paleoenvironment, these parameters have never
been studied on the Pannonian otoliths. They are indepen-
dent proxies for the temperature and salinity of Lake Pannon.
Water temperature and salinity estimates show a wide range
of fluctuations during its history. A bottom-water tempera-
ture of ~15 °C and variable salinities (20—32 ‰) are estimat-
ed for the Early Sarmatian sea, while temperature and
salinity are estimated to have fluctuated from 15 °C to 21 °C
and from 15 ‰ to 43 ‰, respectively in the Late Sarmatian
sea (Tóth et al. 2010). These values are in range with esti-
mates from the composition of microfauna and molluscs. As
Lake Pannon hosted many endemic species (Müller et al.
1999; Geary et al. 2000) the isolation precludes direct esti-
mation of salinity from faunal composition or from oxygen
and carbon isotope studies of bivalve shells (Geary et al.
1989; Mátyás et al. 1996). The Late Sarmatian-Pannonian
samples have higher ratios of heavy isotopes (—2.11 ‰ for
O, and 1.27 ‰ for
C) than do the succeeding Pontian
Stage samples (—4.16 ‰
and —2.22 ‰, respectively). This
shift was interpreted as indicative of a basinwide drop in sa-
linity (Geary et al. 1989).
In addition, the isotope data clearly indicate that Lake
Pannon was not a huge, closed freshwater lake. Assuming a
riverine outlet somewhere along the southern margin, salini-
ty would have been lost and full freshwater conditions ap-
pear in a couple of tens of thousands of years. In contrast, a
closed basin with no outflow needs evaporation of all in-
flowing water and rainfall – impossible under present-day
climatic conditions (Leeder 2007).
Three otolith samples have been selected for the purpose
of this pilot study. The scope was to investigate the preserva-
tion of original incremental and mineralogical structure and
decide if the chemical signature remained unaffected by
post-depositional processes. The preservation of the genuine
microchemistry of the fossil otoliths is a substantial pre-
requisite for any future paleoenvironmental or paleoeco-
KERN, KÁZMÉR, BOSNAKOFF, VÁCZI, BAJNÓCZI and KATONA
GEOLOGICA CARPATHICA, 2012, 63, 2, 175—178
Materials and methods
Fossil otolith samples
Sagittal otoliths were collected from the upper 0.9 m of a
2.6 m deep trench profile dug in a Late Miocene sequence at
Doba village, Bakony Hills, Hungary (Fig. 1). The age of the
fossiliferous littoral marl layer of the Lymnocardium ponti-
cum Zone is estimated as 8.8—9.2 Ma (Magyar et al. 2007).
Twenty-nine otoliths were found after sieving (mesh size:
500 micrometers) of ca. 300 kg bulk material. Most of the
samples (28) were determined as Umbrina subcirrhosa
Schubert, 1902 based on the morphological characteristics.
For further details about the material, the reader is referred to
Bosnakoff & Katona (in print).
Three samples have been selected from the collection to
represent large (UC1) medium (UC2) and small (UC3) size
categories (Fig. 2). Basic morphological parameters have
been measured to the nearest 0.1 mm on each sagittae by a
Preparation for structural analysis
Samples were embedded in epoxy resin and transverse
sections had been prepared using an IsoMet
and a Buehler MiniMet grinding-polishing machine. Cut fac-
es were ground using the standard sequence of water-based
diamond suspensions (9, 6, 3, 1 µm; METADI
) and alumi-
na (0.05 µm; Gamma Micropolish
II) was used for final
polishing. Samples were placed in a distilled water bath and
agitated by an ultrasonic cleaning device for 15 min. Sec-
tioning trials were made both perpendicular to (UC1) and
parallel with (UC2, UC3) the sagittal plane to see any poten-
tial directional difference of structure or mineralogy.
Cathodoluminescence (CL) is an electron-excited phe-
nomenon. Upon bombardment by a high-energy electron
beam originating from a cathode, certain materials emit pho-
tons in the visible range of the electromagnetic spectrum.
Excitation is generated at luminescence centres (activators)
in the material, which are crystal lattice defects and chemical
impurities. It is generally accepted that Mn
is the most im-
portant activator of CL in carbonate minerals (Marshall
1988; Machel 2000). CL microscopy is a common technique
in sedimentary petrology to study the mineralogy, texture
and diagenetic history of carbonate rocks (Machel 2000).
Biogenic carbonates (Barbin 2000) and fish otoliths (Halden
et al. 2004) are also routinely analysed.
Cathodoluminescence examination was performed using a
Reliotron cold-cathode equipment mounted on a Nikon E600
microscope. The equipment was operated at 8 to 10 kV accel-
erating voltage and 0.5 to 1.0 mA current, and a defocused
electron beam was used. Photos were taken using a Nikon
Coolpix 4500 digital camera using automatic exposure.
Raman spectroscopic analysis was done using a HORIBA
JobinYvon LabRAM HR dispersive, edge-filter based confo-
cal Raman spectrometer (focal length: 800 mm) equipped
with an Olympus BXFM microscope. Spectra were taken us-
ing the 785 nm emission of a diode laser, a 100 (N.A. 0.9)
objective, a grating with 600 grooves/mm and a pinhole of
100 µm, which acted also as the entrance slit to the spec-
trometer. Net counting times were between 30 and 60 s. Ra-
man spectroscopy is a routinely used technique to study
small-scale mineralogical properties in fish otoliths (e.g.
Gauldie et al. 1997; Tzeng et al. 2007).
Fig. 1. Geographical and geological setting of the study site. Star
indicates the location of the studied Doba outcrop. Major geological
units (after Cziczer et al. 2009): 1 – pre-Late Miocene basement,
2 – sublittoral deposits of Lake Pannon, 3 – littoral to deltaic de-
posits of Lake Pannon and overlying fluvial sediments, 4 – Upper
Miocene—Pliocene volcanics. Inset map shows the location (black
square) on a map of Europe.
Fig. 2. The studied Pannonian sagittal otoliths, Umbrina cf. subcir-
rhosa Schubert, 1902. Top row shows the internal view with the
characteristic sulcus acusticus. The length of each specimen is indi-
cated. The bottom row shows the polished sections under reflected
light. Tip of the arrow points to the spot of the SEM image in
Fig. 3b. Rectangles help comparison with the CL images in Fig. 3a.
MINERALOGY OF PANNONIAN SCIAENID OTOLITHS: PALEOECOLOGICAL IMPLICATIONS
GEOLOGICA CARPATHICA, 2012, 63, 2, 175—178
Results and discussion
Macroscopic and microscopic incremental structures
Darker and lighter bands were clearly observed on each
polished section independently of the direction of the section-
ing plane (Fig. 2). These coupled bands were described of the
recent sciaenid otoliths to indicate an annual increment (annu-
lus) (Arneri et al. 1998; La Mesa et al. 2008; Engin & Seyan
2009). The annual periodicity of annulus deposition was con-
firmed by the edge analysis in recent sciaenids, indicating that
translucent and opaque zones are formed once a year. The
opaque zone was laid down in summer, between June and July
while the dark/translucent zone was found to be deposited
during the rest of the year, starting at the end of summer
(September) (La Mesa et al. 2008). The ontogenic age of the
specimens can be assigned on the basis of counts of dark/
translucent winter bands (Arneri et al. 1998). The estimated
age of UC1 is 17, of UC2 is 7, while of UC3 is 6 years.
The columnar crystal fabric was also perfectly preserved
(Fig. 3b) indicating the lack of any diagenetic impact on
Estimated somatic parameters
A formalized experimental relationship has been pub-
lished recently describing ontogenic age vs body length and
otolith maximum length vs body weight relationships for
brown meagre (Sciaena umbra Linnaeus, 1758) (La Mesa et
al. 2008), a possible present-day relative to the Pannonian
Umbrina subcirrhosa. The obtained equations are thought to
be robust as they rely on 532 specimens of brown meagre
captured over a period of 11 years. Using equations obtained
on this modern dataset, somatic parameters of the Pannonian
fish specimens can be estimated as 46 cm/1470 g for UC1;
42 cm/1130 g for UC2 and 41 cm/1030 g for UC3.
It is necessary to emphasize that these estimated somatic pa-
rameters must be treated with caution. For instances the large
difference in the size of the water bodies (i.e. Modern Adriatic
Sea and Late Miocene Lake Pannon) and the lack of knowl-
edge about the food web complexity of the paleoenvironment
might introduce significant and hardly quantifiable bias.
Preserved mineralogical composition
Otolith cores showed green, yellowish green luminescence.
The intensity vanished towards the rim while faint blue lumi-
nescence was observed at the outer sectors (Fig. 3a). It is inter-
esting to note that seasonal increments showed different CL
characteristics within the annulus. Greenish and weak blue lu-
minescence bands corresponded to the opaque (summer) and
dark/translucent (winter) zones, respectively.
minerals have characteristic CL colours caused by
the substitution of Mn
: orange(-red) for calcite and
green to yellow for aragonite (Marshall 1988). ~ 10—20 ppm
concentration is necessary for visible Mn
minescence in biogenic calcite and the same or less in arago-
nite (Barbin 2000). In the case of pure carbonates or
carbonates with minor traces of substituting element, a weak
blue (so-called intrinsic) CL is detected for both polymorphs.
The observed CL features, consequently, indicate arago-
nitic mineralogy in the otolith cores but the results are still
not conclusive in the outer part. Raman microspectrometric
analyses helped to clarify the ambiguity. The Raman spectra
Fig. 3. Structural evidence
for the lack of recrystalliza-
tion. a – Cathodolumines-
cence (CL) images. White
dots mark the spots for Ra-
(annotation corresponds to
the legend in c). Rectangles
help to orientate the CL im-
age in the polished section
of Fig. 2. b – Radially ar-
ranged columnar aragonite
crystals in UC3. c – Ra-
man spectra from UC1.
Vertical dashed lines indi-
cate the characteristic Ra-
man bands of aragonite.
correspond to the arago-
nite phase (Urmos et al.
1991; Gauldie et al.
1997) proving that the
Pannonian otolith un-
doubtedly contained ara-
gonite crystals at each
analysed spot (Fig. 3c).
As all the fossil otolith
KERN, KÁZMÉR, BOSNAKOFF, VÁCZI, BAJNÓCZI and KATONA
GEOLOGICA CARPATHICA, 2012, 63, 2, 175—178
material provided only the characteristic Raman bands of ara-
gonite, regardless of the CL colour found in the otolith zone, it
is proven that the original aragonitic mineralogy has been pre-
served also in the parts with weak blue luminescence.
Incidentally, alternating greenish and weak blue CL zones
corresponding to opaque and translucent incremental zones
of annuli indicate that Mn
concentration drops below the
~10—20 ppm level in the increment laid down during the
Macroscopic and microscopic observations of the three
studied Pannonian (Late Miocene) sciaenid otoliths offered
clear evidence for the preservation of the genuine structural
characteristics. Ontogenetic ages were assigned for the three
specimens as 16, 7 and 6 years, respectively, by counting the
annuli of sagittae. Corresponding somatic parameters of the fish
specimens can be estimated as 46 cm/1470 g; 42 cm/1130 g
and 41 cm/1030 g using experimental physiological relation-
ships of the brown meager (Sciaenia umbra Linnaeus, 1758) a
modern relative of the studied Pannonian species.
Results from CL and Raman microspectrometric analyses
proved the preservation of the original aragonitic mineralo-
gy. The excellent preservation of these fossils invites further
microchemical studies to reconstruct the past physicochemi-
cal environment. As earlier geochemical/paleoecological
studies concentrated exclusively on benthic organisms, these
otoliths might provide the first geochemical record from
Pannonian nektonic organisms related directly to the open
water marine/lacustrine conditions free from any bias due to
the benthic environment.
Acknowledgments: The European Union and the European
Social Fund have provided financial support to the Project
under the Grant agreement No. TÁMOP 4.2.1./B-09/KMR-
2010-0003. The research has been supported by Baross Gá-
bor Fund REG-KM-09-1-2009-0044 and OTKA K67583
grants. The authors thank Ágnes Takács for her help with
sample preparation. Rostislav Brzobohatý and Norman
Halden are acknowledged for their constructive reviews.
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