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Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72 – 81

First record of Hindeodus–Isarcicella population

in Lower Triassic of Slovenia
Tea Kolar-Jurkovšek ⁎, Bogdan Jurkovšek
Geological Survey of Slovenia, Dimičeva 14, SI-1000 Ljubljana, Slovenia
Accepted 30 November 2006


This study documents the Permian–Triassic Boundary in the Žiri area of western Slovenia. A paleontological study of the
Lukač section, northwest of Žiri, resulted in the recovery of a variety of microfossils. This report documents the conodont fauna
recovered from the section. This is the first report of the Hindeodus–Isarcicella population in Slovenia and it provides important
new biostratigraphic data for the definition of the Permian–Triassic Boundary. Hindeodus parvus has been recovered, and its first
appearance in the lowermost Werfen Formation indicates the base of the Triassic Period. Based on the conodont taxa H. parvus,
Hindeodus typicalis, Hindeodus sp., Isarcicella isarcica, Isarcicella lobata, Isarcicella staeschei, Isarcicella turgida, and Isarci-
cella sp. A, at least three faunas can be distinguished, and they make possible a worldwide correlation.
© 2007 Elsevier B.V. All rights reserved.

Keywords: Conodonts; Hindeodus–Isarcicella; Permian–Triassic Boundary; Lower Triassic; Slovenia

1. Introduction Masore area, west of Žiri. Later, Dolenec et al. (1999)

documented a negative shift of organic carbon (Corg) in
The Permian–Triassic interval of wider Žiri area was these strata.
the scope of numerous studies. However, it has not been The purpose of our study was to document the PTB
dealt with paleontologically in detail by means of interval by means of conodonts. A paleontological study
conodonts. Often the boundary was lithologically of the lowermost part of the Werfen Formation in the
defined within the dolomite interval overlying the Lukač section, 3.5 km northwest of Žiri (Fig. 1), was
limestone of the Žažar Formation (= Bellerophon carried out and 7.5 m of the section (Fig. 2 – shown as
Formation, Grad and Ogorelec, 1980; Mlakar and the shaded part – and Fig. 3) was examined in detail. In
Placer, 2000). Based on the foraminifera Earlandia the first phase of our study we collected 16 samples for
tintinniformis (Mišik) and the annelid Spirorbis phlyc- conodont analyses (Table 1). All except one were
taena Brönnimann and Zaninetti, Buser (1986) inter- productive, yielding a diverse fauna of conodonts,
preted a Permian–Triassic Boundary (PTB) in the ostracods, foraminifers, small bivalves, gastropods, etc.
In this paper we document the lowermost Triassic
⁎ Corresponding author. Tel.: +386 1 2809739; fax: +386 1
conodont faunas, the first report of the Hindeodus–
Isarcicella population from western Slovenia. The
E-mail addresses: conodont Hindeodus parvus (Kozur and Pjatakova)
(T. Kolar-Jurkovšek), (B. Jurkovšek). has been identified in our material and the first
0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.
T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81 73

Fig. 1. Location map of the studied section in western Slovenia.

appearance datum (FAD) of this taxon has been ap- ciation, among which prevail the alga Gymnocodium
proved to define the base of the Triassic system (Yin bellerophontis (Rothplatz), frequent bellerophons, and
et al., 2001). locally rich brachiopods.
Upper Permian strata are continuously overlain by a
2. Geological setting Lower Triassic succession. In many cases, in proximity
to the PTB these strata are represented by dolomitic
In western Slovenia three large geotectonic units facies (Buser et al., 1986). In the wider Žiri area, the
are recognized: the Southern Alps; the Inner Dinarides; lowermost Triassic is represented by a unit known as the
and the Outer Dinarides (Buser, 2003). The investigated “Streaky Limestone Member” (Slovenian: “pasnati
section of the interval located in the Žiri area is part apnenec”) of the Werfen Formation (Mlakar, 2002)
of the Outer Dinarides. The area is composed of that is up to 40-m thick. This unit also contains some
Carboniferous, Permian, and Triassic rocks. Carbonif- intercalations of siltstone. The lowermost part of the
erous strata are represented by clastic sediments with Werfen Formation was the subject of our study and it
prevailing black shale, sandstones and conglomerate. contains diverse microfauna, of which the conodonts are
The upper part of this sequence may be Lower Perm- dealt with in this study.
ian in age, but that has not yet been demonstrated
paleontologically. 3. Conodont faunas
The Middle Permian strata rest discordantly on the
Carboniferous succession. They are composed of silt- The recovered associations are marked by a Hin-
stones, sandstones, and conglomerates, lithologies that are deodus–Isarcicella-dominated population. It is charac-
similar to those found in the Val Gardena Formation. In teristic for shallow water habitats, and that is in
Žirovski vrh Hill region, located 4.5 km northeast of Žiri, accordance with the lithofacies of the sediment. A
these strata are characterized by uranium mineralization complete absence of gondolellids has been noted.
(Budkovič, 1980; Skaberne, 2003). Conodonts of the PTB interval have been intensively
The Upper Permian fossiliferous limestones in studied in many parts of the world due to great
Slovenia are known as the Žažar Formation, a unit stratigraphic use of the Hindeodus–Isarcicella species,
that is synonymous with the Bellerophon Formation of parallel to the pelagic gondolellids whose importance in
the Carnian Alps and Dolomites (Ramovš, 1958; biostratigraphy has been demonstrated and is well
Skaberne and Ogorelec, 2003). The Upper Permian known. H. parvus, representative of the Hindeodus
strata are characterized by limestone and dolomite typicalis group, was favored and later verified to define
development with a typical Late Permian fossil asso- the base of the systemic boundary (Yin, 1993; Jin et al.,
74 T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81

Fig. 2. Lithostratigraphic succession of Paleozoic and Lower Triassic beds in Žiri area with position of investigated interval (shaded).

1996; Yin et al., 2001). The decision was based upon shore, shallower regions. Kozur (1996) made a remark
world-wide distribution and very high ecological that due to its very high ecologic tolerance – beyond the
tolerance of the species. tolerance of other conodonts – the H. typicalis group,
Similar conodont faunas lacking abundant gondolel- occurring in many shallow water deposits, is best suited
lids have been widely reported from the Southern Alps for making world-wide comparison.
(Huckriede, 1958; Staesche, 1964; Farabegoli et al.,
1986; Perri and Andraghetti, 1987; Perri, 1991; Kozur, 3.1. Conodont faunas of the Lukač section
1996; Nicora and Perri, 1999; Perri and Farabegoli,
2003). The facies control of the P–T conodont faunas For this study the lower 7.5 m of the “Streaky
was intensively studied in the last decade (Orchard, Limestone Member” was examined in detail (Fig. 2). A
1996; Kozur, 1996; Orchard and Krystyn, 1998; Nicoll total of 16 samples, mainly composites, were collected
et al., 2002; Kozur, 2005). The relative abundance of (Fig. 3) and all but one (sample LU 06) produced cono-
Hindeodus and Neogondolella was observed in an donts (Table 1, Plate 1). In addition to conodonts they
onshore–offshore relationship: gondolellids are more yielded various microfauna, including ostracods, tiny
common in offshore, deeper water marine environ- bivalves, and gastropods. The conodont fauna is marked
ments, whereas Hindeodus flourished mostly in near- by Pa elements of Hindeodus and Isarcicella species,
T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81 75

Fig. 3. Studied P–T interval of the Lukač section near Žiri; 1 — limestone, 2 — dolomite, 3 — silty limestone, 4 — calcareous shale, 5 — covered,
6 — clayey shale lamina, 7 — ooid.
76 T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81

Table 1
Numerical distribution of conodont taxa in samples from the Lukač section
07 06 05 04 03 02 01 1A 1B 2 3 4 5 6 7 8 Sample number taxa
1 Hindeodus sp.
1 1 12 Hindeodus parvus
1 Hindeodus cf. typicalis
10 1 2 5 Isarcicella sp. A
1 2 1 Isarcicella turgida
1 1 1 Isarcicella lobata
3 9 9 1 Isarcicella staeschei
2 4 Isarcicella isarcica
16 1 2 1 8 8 1 Isarcicella sp.
1 2 1 16 1 1 1 12 2 5 2 Ramiforms

but other element types are very rare. The following taxa section, marked only by the presence of H. parvus, can
were identified: H. parvus; H. typicalis (Sweet); Hin- be distinguished as Fauna 1.
deodus sp.; Isarcicella sp. A; Isarcicella lobata Perri The following conodont fauna, Fauna 2, develops
and Farabegoli; Isarcicella turgida (Kozur, Mostler and with the first record of the genus Isarcicella in the
Rahimi-Yazd); Isarcicella isarcica (Huckriede); and sample LU 02. The first occurrence of Isarcicella in the
Isarcicella staeschei Dai and Zhang. Through the section is marked by the co-occurrence of Hindeodus
section a gradual morphological change among isarci- species. The predominant isarcicellid forms are char-
cellids was observed. This has enabled the recognition acterized by a thickened cup, but without lateral orna-
of at least three distinct faunas. In addition, Hadrodon- mentation such as nodes and denticles. The specimens
tina is present in the uppermost samples. share some similarities with I. prisca and I. turgida, but
The lowest conodont recovered in the section was a they both reveal a more thickened inner side of the cup
single specimen of Hindeodus from sample LU 07, and thus form an asymmetric outline of the lower side.
located approximately 20 cm above dolomite (Fig. 3); that Most specimens of this interval display a subsymme-
is, below the first occurrence of H. parvus. The specimen trical cup and have denticles that are unequal in size and
markedly differs from all other hindeodid specimens typically unevenly aligned. Therefore, these specimens
found in the higher stratigraphic levels in that it is an are separated here as Isarcicella sp. A. They are close to
elongate and thin form. Unfortunately, it is not completely I. lobata. The latter species is herein recognized for its
preserved and the posteriormost part is lacking so that the specimens with conspicuous development of the lobe
profile cannot be fully reconstructed. The specimen and the lower view that shows a strong asymmetric
shows a close resemblance to H. pisai Perri and outline. However, in Isarcicella sp. A the lower side is
Farabegoli, described from the Dolomites where it ranges nearly symmetrically expanded on both sides of the cup.
from the uppermost Permian to the lowermost Triassic Moreover, the two species also differ in alignment of
(uppermost Upper praeparvus–lower parvus zones, Perri denticles as viewed from upper side. In I. lobata, the
and Farabegoli, 2003). series of denticles is aligned with bending in the middle
The species H. parvus first occurs in the sample LU part, and sometimes sigmoidal bending can be observed
05 and is present up to the sample LU 02. The highest (see Perri and Farabegoli, 2003); in Isarcicella sp. A this
occurrence of the species is accompanied by H. typicalis line of denticles is straight, apart from the two small
and Isarcicella sp. A. The lower part of the investigated denticles posterior to the cusp.

Plate 1. Lowermost Triassic (Lower Induan) conodonts from the Lukač section, western Slovenia.

1. Hindeodus parvus (Kozur and Pjatakova), sample LU 05 (GeoZS 3804).

2–3. Isarcicella sp. A, samples LU 1A (GeoZS 3791), LU 4 (GeoZS 3795).
4. Isarcicella lobata Perri and Farabegoli, sample LU 2 (GeoZS 3793).
5. Isarcicella turgida (Kozur, Mostler and Rahimi-Yazd), sample LU 1A (GeoZS 3791).
6–9. Isarcicella staeschei Dai and Zhang, samples LU 4 (GeoZS 3795), LU 7 (GeoZS 3798), LU 2 (GeoZS 3793).
10–11. Isarcicella isarcica (Staesche), samples LU 4 (GeoZS 3795), LU 7 (GeoZS 3798) Scale bar 50 μm.All studied and illustrated material is
deposited with Geološki zavod Slovenije (Geological Survey of Slovenia) and abbreviated GeoZS.
T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81 77
78 T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81

The third conodont fauna, Fauna 3, is marked by the In the Italian Dolomites the Werfen Formation can be
presence of Isarcicella elements with lateral ornamen- divided in eight members based on lithostratigraphic
tation that make its first appearance in the sample LU 2 subdivision and the recognition of transgressive and
(Fig. 3). The species I. lobata and I. staeschei first regressive events (Broglio Loriga et al., 1983, 1986).
appear simultaneously, but I. staeschei is the dominant Recent conodont biostratigraphic study across the P–T
species. boundary interval clarified morphological trends in the
The present concept of discriminating highly evolved genera Hindeodus and Isarcicella. It resulted in
isarcicellids is based on the presence of lateral nodes or recognition of the praeparvus zone, the youngest
denticles on one lateral side (I. staeschei) or on both Permian conodont zone immediately above the Beller-
sides (I. isarcica) of the inflated cup. The wide ophon Formation, and the following succeeding parvus,
intraspecific variability of these species was observed lobata, staeschei, and isarcica zones in the Werfen
by Perri (1991), and it can also be demonstrated in the Formation (Perri and Farabegoli, 2003). That was
material from this study. The recovered elements differ possibly due to the medium-to high-sedimentation rate
in their length to height ratio. The lateral lobe may bear of the sequences across the PTB. In this zonation a rapid
weak nodes or strong denticles, varying from one to four evolution of isarcicellids is noted: an introduction of two
in number. In our material the representatives of forms with a swelling cup producing lobes on one or
I. staeschei and I. lobata first appear in the same bed both sides (I. lobata, I. inflata Perri and Farabegoli)
(sample LU 2). However, in the Dolomites a different marking the lower limit of the lobata zone followed by
FAD of these species was demonstrated. Based on these ornamentation of the cup in the youngest isarcicellids
data, two conodont zones were separated in Dolomites: (I. staeschei, I. isarcica), based on gradual evolution of
the lobata zone and the staeschei zone (Perri and the two taxa. Correspondingly, staeschei and isarcica
Farabegoli, 2003). zones have been discriminated.
According to Kozur (1996), the entire isarcica zone In Austria, the Gartenkofel-1 core of the Carnic Alps,
is characterized by dominant isarcicellids with one or comprising mainly dolomitic carbonates of the Upper
two denticles on one side, in association with the Permian Bellerophon Formation and the Lower Triassic
elements displaying denticles on both sides of the cup. Werfen Formation underwent a multidisciplinary study
The first appearance of the forms with only one denticle (Holser and Schöulaub, 1991). Conodont examination
present on the lateral side of the blade is extremely rare. provided diverse Hindeodus– Isarcicella faunas
As they occur already in the parvus zone, Kozur (1996) (Assemblages A–D), and the youngest assemblage
is of the opinion, that only forms with one denticle (Assemblage E) characterized by H. aequabilis Staesche
(monodenticulate forms) on one side should be used as a (originally described as Ellisonia aequabilis, Schön-
separate taxon at the subspecific level (I. isarcica laub, 1991). The lowermost association, derived from
staeschei). In this paper the two forms are ascribed as the lowermost Tesero Horizon, was correlated to the
I. staeschei and I. isarcica. Otoceras woodwardi zone.
The youngest isarcicellid occurrences are accompa- In Slovenia, Lower Triassic strata were the scope of
nied by Hadrodontina aequabilis Staesche that con- many studies, but conodont faunas were mostly obtained
tinues to be present (as mono fauna), and are also above from its higher part. All recovered assemblages are
the last occurrence of Isarcicella. H. aequabilis is the characterized by the Pachycladina obliqua Staesche
marker of the next zone, the H. aequabilis zone. As this apparatus, Hadrodontina sp. and/or Furnishius triserra-
form does not belong to the Hindeodus–Isarcicella tus Clark, Parachirognathus ethingtoni Clark, and Fo-
phylogenetic lineage, it is not a matter of discussion here. liella gardenae (Staesche). Occasionally, elements of
Ellisonia can be present. The conodont faunas were
3.2. Comparison with conodont faunas of other areas marked by the typical shallow water genera Foliella,
Hadrodontina, Pachycladina, and Parachirognathus.
The conodont fauna, as part of an extensive study of Conodont collections from the Slovenian part of the
the P–T interval, was investigated in many regions of the Outer Dinarides (sensu Placer, 1999) have been well
world. In comparing the obtained conodont faunas from documented and can be compared well with other
Slovenia, correlation with the faunas of the Southern equivalent collections hitherto studied in other parts of
Alps is most suitable due to its geographic proximity as Slovenia (Kolar-Jurkovšek, 1990; Kolar-Jurkovšek and
well as its very detailed conodont biozonation. However, Jurkovšek, 1995; Kolar-Jurkovšek, 1996; Kolar-Jurkov-
there are also strong similarities to the Early Triassic šek and Jurkovšek, 1996). Faunal lists of these Smithian
faunas of the Meishan section of China. localities are very similar (Kolar-Jurkovšek and
T. Kolar-Jurkovšek, B. Jurkovšek / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 72–81 79

Jurkovšek, 2001). They were correlated to the Lower of our study it would be premature to make a conclusion
Smithian Zone 7 (Parachirognathus–Furnishius zone) on their development and to draw differences to the
of Sweet et al. (1971) and the Smithian obliqua zone of faunas of other areas. More data would probably answer
Perri (1991). the questions that appeared in this study; therefore,
In the vicinity of Tržič, in Karavanke, a biostrati- further examination of this section is proceeding. The
graphic study of the Upper Permian–Lower Triassic strata ongoing step of paleontological study will include
was conducted (Kolar-Jurkovšek and Jurkovšek, 1995). detailed descriptions of identified conodont taxa and
The Upper Permian complex consisting of monoto- establishment of a conodont biozonation. Additional
nous bedded dolomite was examined for microfossils. sampling will be focused towards bringing precision to
The samples were barren of conodonts, but an ostracod the lobata–staeschei zone, and defining the uppermost
fauna yielding representatives of the palaeocopid genus Permian level with a conodont fauna.
Hollinella was recognized. That study indicated the A detailed lithofacies analysis is part of an ongoing
possibility that in certain sections, the ostracod fauna study, and together with chronostratigraphic data it will
may serve as a valuable biostratigraphic tool, as shown by allow comparison with equivalent strata of adjacent
Kozur (1985) for beds of the same age and similar facies regions. A precise correlation of the studied Lower
in the Bükk Mts., Hungary. The stratigraphic importance Triassic beds in Slovenia with the Tesero and lower
of Palaeocopida and other ostracods across the P–T Mazzin members, the oldest lithostratigraphic units of
Boundary was emphasized by Kozur (1985) and the Werfen Formation in the Southern Alps, will be
Crasquin-Soleau et al. (2004). worked out.
In the Outer Dinarides of Croatia, a lithostratigraphic
and biostratigraphic study of the Lower Triassic shallow Acknowledgments
marine succession in Gorski Kotar region was undertaken
(Aljinović et al., 2006). Conodont faunas were recovered This study was financially supported by the Slove-
from all five studied sections. The occurrence of nian Research Agency (program number P1-0011 and
H. parvus in the Školski Brijeg section was reported project number J1-6665). The authors thank D. Alji-
from the lower part of the section represented by a basal nović (Zagreb) for her comments and providing valuable
dolomitized, oolitic bar facies (F-1, Aljinović et al., 2006). observations during preparation of this paper and Y.
Lower Triassic conodonts were also reported from some Isozaki (Tokyo) for reading an earlier draft of the
other sections of this unit in Croatia, but they belong to manuscript. R. Nicoll (Canberra) and H. Kozur (Buda-
Olenekian fauna, namely the obliqua and triangularis pest) are gratefully acknowledged for reviewing the
zones (Herak et al., 1983, Jelaska et al., 2003). manuscript and making helpful suggestions. The authors
are much obliged to M.J. Orchard (Vancouver), for
4. Concluding remarks editing and thoroughly reading the manuscript. Scanning
electron microscopy was carried out at Paleontološki
In order to define the PTB interval by means of inštitut ZRC SAZU (Ljubljana). The paper constitutes a
conodonts, a study of the Werfen Formation in Žiri area contribution to the IGCP-Project 467.
was undertaken. Examination of the microfauna obtained
from the first sampling in its lower part revealed References
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