Tea Kolar-Jurkovšek, Valery J. Vuks, Dunja Aljinović, Michael Hautmann, Andrzej Kaim & Bogdan Jurkovšek

OLENEKIAN (EARLY TRI AS SIC) FOS SIL AS SEM BLAGE

FROM EAST ERN JULIAN ALPS (SLOVENIA)

Tea KOLAR-JURKOVŠEK

, Valery J. VUKS

, Dunja ALJINOVIÆ

, Mi chael HAUTMANN

,

Andrzej KAIM

& Bogdan JURKOVŠEK

1

Geo log i cal Sur vey of Slovenia, Dimièeva ulica 14, 1 000 Ljubljana, Slovenia; e-mail: tea.kolar@geo-zs.si,

bogdan.jurkovsek@geo-zs.si

2

Fed eral State Uni tary En ter prise “A.P. Karpinsky Rus sian Geo log i cal Re search In sti tute” Sredny pr. 74,

199106 St.-Pe ters burg, Rus sia; e-mail Valery_Vuks@vsegei.ru

3

Uni ver sity of Zagreb, Fac ulty of Min ing, Ge ol ogy and Pe tro leum En gi neer ing, Pierottijeva 6, 10 000 Zagreb, Croatia;

e-mail: dunja.aljinovic@rgn.hr

4

Paläontologisches Institut und Mu seum, Karl Schmid-Strasse 4, 8006 Zürich, Swit zer land;

e-mail mi chael.hautmann@pim.uzh.ch

5

Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, PL-00-818 Warszawa, Po land;

e-mail: kaim@twarda.pan.pl

Kolar-Jurkovšek, T., Vuks, V. J., Aljinoviæ, D., Hautmann, M., Kaim, A.& Jurkovšek, B. 2013. Olenekian (Early Tri as sic) fos sil as sem blage from east ern Julian Alps (Slovenia). Annales Societatis Geologorum Poloniae, 83: 213–227.

Ab stract: New palaeontological and sedimentological data from the Lower Tri as sic strata of the east ern Julian Alps in Slovenia are pre sented., They are un usual for the Early Tri as sic of the Alps in rep re sent ing a rel a tively deeper, un re stricted ma rine (mid-ramp) set ting. There are two ba sic microfacies types in the sec tion in ves ti gated (types A and B), which are or ga nized as cou plets with coarsegrained tempestitic de pos its (microfacies A), over -lain by lam i nated or bioturbated lime mudstones and/or marls (microfacies B), fre quently con tain ing ammo-noids. This pat tern is in ter preted as storm de po si tion with oc ca sional win now ing of bot tom sed i ments and the for ma tion of coarse-grained skel e tal de pos its (lags), fol lowed by the slow set tling of sus pended par ti cles, when the storm waned, in ad di tion to back ground de po si tion. Dom i nantly lime mud de po si tion and the pres ence of ammo-noids in di cate de po si tion on a more dis tal, deeper ramp with an un re stricted con nec tion to the open sea. In tense re - work ing of bot tom skel e tal-rich sed i ment and ac cu mu la tion of storm lags sug gest de po si tion above the storm wave base, pos si bly in a wide low-en ergy mid-ramp en vi ron ment. Fau nas from such set tings have been re ported rel a tively rarely from the Early Tri as sic of the Alps. The macrofauna con tains ammonoids, bi valves and gas tro -pods, whereas the micro fauna is rep re sented by foraminifer tests and cono dont el e ments; rare fish re mains also oc cur. In the foraminifer as sem blages, spe cies of Ammodiscus, Hoyenella, Glomospirella dom i nated, cor re spon-d ing to the wispon-de spreaspon-d “Glomospira-Glomospirella” foraminifer com mu nity, with some miliolispon-ds anspon-d nospon-dosa- nodosa-riids. The cono dont fauna is char ac ter ized by Triassospathodus hungaricus (Kozur et Mostler), in di cat ing an early Spathian (Olenekian) age. The fos sil as sem blage high lights the wide dis tri bu tion of Early Tri as sic taxa in the Tethys and fa cil i tates its world wide cor re la tion. Its rel a tively low di ver sity by com par i son with shal low ma rine set tings is in ter preted as an evo lu tion ary prox i mal-dis tal trend in the wake of the end-Perm ian mass ex tinc tion. Re-diversification first oc curred in nearshore set tings and ex panded into deeper/dis tal ma rine en vi ron ments through geo log i cal time.

Key words: Olenekian, Early Tri as sic, foraminifers, bivaves, gas tro pods, cono donts, sedimentology, Julian Alps, Slovenia.

Manu script re ceived 4 August 2013, ac cepted 23 December 2013

IN TRO DUC TION

The end-Perm ian mass ex tinc tion event was piv otal in

the Phanerozoic his tory of ma rine life, ter mi nat ing the in

-cum bency of the “Palaeozoic Evo lu tion ary Fauna” that had

dom i nated life in the sea for nearly 250 mil lion years

(Sepkoski, 1984). In the wake of the ex tinc tion event, the eco

-log i cal struc ture of the pres ent-day bio sphere evolved from

the stock of the sur viv ing taxa, shap ing the “Mod ern Evo lu

-tion ary Fauna” that char ac ter izes ma rine eco sys tems un til

to day (Sepkoski, 1997). The re cov ery from the endPerm

-ian mass ex tinc tion is there fore cru cial for un der stand ing

how the pres ent-day bio sphere evolved. An in dis pens able

prereq ui site in this con text is a cen sus of the fau nas that

lived and evolved dur ing the Early Tri as sic, i.e., the ep och

that fol lowed the cri ses. Thus, col lect ing and in ter pret ing

new field data from the Early Tri as sic has been a ma jor re

-search agenda dur ing the past de cade (e.g., Brayard and Bu-

cher, 2008; Posenato, 2008; Kumagae and Nakazawa, 2009;

Kaim et al., 2010, 2013; Hautmann et al., 2011, 2013;

Brühwiler et al., 2012; Wasmer et al., 2012; Hofmann et al.,

2013a, b). How

ever, Early Tri

as

sic ben

thic fau

nas have

been de scribed mostly from rel a tively shal low ma rine sec

-tions, par tic u larly in Eu rope, where deeper shelf set tings are

rare. The sec tion in ves ti gated is also known as the lo ca tion,

where an iso lated temnospondyl bone (Lucas et al., 2008),

was found and there fore rep re sents an im por tant site of

Slo-venian nat u ral her i tage. In this study, new

sedimentologi-cal and palaeontologisedimentologi-cal data are pre sented from the

Spa-thian (late Early Tri as sic) sec tion in the Julian Alps

(Slove-nia). They con

trib

ute to a better un

der

stand

ing of how

diversity and fau nal com po si tion changed along depth gra

-di ents dur ing re--di ver si fi ca tion from the great est cri sis in

the his tory of life.

GEO LOG I CAL SET TING

The study area is in the Julian Alps, NW Slovenia

(Fig. 1). Struc tur ally they form the east ern most con tin u a tion

of the South ern Alps, where the South Al pine and the

Di-naric struc

tures now over

lap (Placer, 1999). The larg

est

overthrust unit in the Slovenian part of the Julian Alps is

rep

re

sented by the Julian Alps Overthrust (Jurkovšek,

1987a, b) or the Julian Nappe, which is com posed of suc

-ces sions, rang ing from Early Tri as sic to Cre ta ceous strata,

though a ma jor part of it is made up of Late Tri as sic car bon

ates. Early Tri as sic strata are scat tered in nar row dis con

-nected belts or patches in the wider Julian Alps area. They

usu ally oc cur within smaller tec tonic slices (Figs 2, 3).

The area in ves ti gated is lo cated in the up per most tec

-tonic slice of the Julian Alps Overthrust, SE of Triglav Mt.

(2863 m), the high est peak in Slovenia. The lower part of

these strata is in tec tonic con tact with the Norian–Rhaetian

Dachstein Lime stone, whereas up wards they pass con tin u

-ously into Anisian lime stones and dolomites (Jurkovšek,

1987a; Celarc and Kolar-Jurkovšek, 2008).

The Studorski preval sec tion is strongly tectonized (co

-or di nates: N46°21'14.46'', E13°52'55.80''; WGS 84). Thick

lime mudstones and marls and thin beds en riched in

coarsegrained bioclastic ma te rial, pre dom i nate. The fauna is char

-ac ter ized by Early Tri as sic mol lusks, dom i nated by bi valves

and gas tro pods. The micro fauna is di ver si fied, but con sists

mainly of foraminifer as sem blages in as so ci a tion with rare

cono dont el e ments and fish re mains.

Li thol ogy

The Studorski preval sec tion mainly con sists of thick

mudstones (Fig. 4A) and marls (Fig. 4B), and thin beds, en

-riched in coarse-grained bioclastic ma te rial. These

litholo-gies form units, in which coarse-grained bioclastic ma te rial

is con cen trated at the base and grad u ally passes up ward into

mudstones/marls. Thus, the microfacies in the Studorski

preval sec tion cor re spond to two microfacies types: mi

crofa cies type A, fossiliferous float stone/packstones, and mi

-crofacies type B, dense or lam i nated lime mudstones that

may pass grad u ally into marls.

Centi metre- to decimetre-thick fossiliferous float stone/

packstone beds (A) have dis tinc tive sharp lower ero sional

bound

aries. The float

stone/packstone interbeds

gradationally change up ward to much thicker mudstone/marl in ter

-vals, which lo cally were in ten sively re worked by or gan isms

(Fig. 4C).

Float stones/packstones of microfacies type A con sist of

coarse (usu ally > 2 mm) well pre served skel e tal de tri tus

(Fig. 4F). Among the skel e tal frag ments bi valves, gas tro

pod, and ammonoid de tri tus pre dom i nate. Echinoderm os si

-cles and foraminifer tests are rare. In the float

stone/pack-stone a bi modal sorted biofabric is found, which con sists of

large bioclasts and micritic ma trix. Very fre quently, the ma

-trix is non-ho mog e nous and con tains a high pro por tion of

peloidal par ti cles and some micrite (Fig. 4E). Whole valves

are dom i nantly ori ented par al lel to the bed ding sur face, very

of ten in a con

vex-up po si

tion. Com pletely pre served bi

-valve or gas tro pod shells are geopetally infilled with sparry

cal cite. Some shel ter pores, at the con cave side of bi valve

shells, are also infilled by sparite.

Microfacies type B con sists of dense, lam i nated or

bioturbated mudstones and/or marls. Sparse, well pre served fos

-sils can be dis persed in the mudstones (Fig. 4D). The

mud-stones con sist of ho mog e nous micritic mud or micrite-rich

laminae. Dis tur bance of the lam i na tion is due to re work ing

by infaunal or gan isms. Lime mud is slightly recrystallized. In

both microfacies types euhedral py rite crys tals and large ce

-lest ite crys tals (de ter mined by EDS anal y sis) are pres ent.

MA TE RIAL AND METH ODS

The pres ent study is based on field work, car ried out in

2011 at Studorski preval in the Julian Alps. Ap prox i mately

10 m of the Lower Tri as sic strata was sam pled just be low

the con tact with the Anisian do lo mite. Nine sam ples (SP

1–9) were col lected for ex am i na tion from the Lower Tri as

-sic strata; sam ple SP 1 is the low est and the sam ple SP 9 is

the high est. Rock sam ples with an av er age weight of 4 kg

were pro cessed for cono dont study, us ing stan dard lab o ra

-tory tech niques. Sev eral thin sec tions were made for the

study of foraminifers as well as for petrographic pur poses.

Lab o ra tory prep a ra tion was car ried out at the Geo log i cal

Sur vey of Slovenia (Geološki zavod Slovenije) where all of

the micropalaeontological ma te rial is stored and in ven to ried

un der re pos i tory num bers 4429, 4548–4555, 4901–4904

and ab bre vi ated GeoZS. The col lec tion of macrofossil spec

-i mens: Jurkovšek Paleontolog-ical Col lec t-ion, Kamn-ica 27,

Dol pri Ljubljani, Slovenia, has been reg is tered with the

Nat u ral His tory Mu seum of Slovenia, Ljubljana and ab bre

vi ated as BJ. The de ter mi na tions of cono dont el e ments pre

-sented here and the SEM/EDS anal y ses were car ried out,

us ing the JEOL JSM 6490LV Scan ning Elec tron Mi cro

-scope at the Geo log i cal Sur vey of Slovenia.

Fig. 2. Gen er al ized strati graphic sec tion of Tri as sic strata in Julian Alps. Thick dark bar in di cates sec tion stud ied in tec tonic slice at Studorski preval

Fig. 3. Palaeogeographic map for Early Tri as sic with po si tion of the Julian Alps marked (star), mod i fied from Scotese (2001)

Fig. 4. Microfacies types from Early Tri as sic of Studorski preval, Slovenia. Ham mer length 26.5 cm (A) and 29 cm (B, C). A. Lam i -nated, thin-bed ded mudstones. B. Ho mog e nous marl in ter val. C. Bioturbated lime mudstone. D. Rare skel e tal frag ments in dom i nantly muddy sed i ment – microfacies type B. E. Float stone (microfacies type A) with coarse skel e tal de tri tus and inhomogeous ma trix con tain -ing high pro por tion of pelloidal par ti cles and some micrite. F. Float stone of microfacies type A con sists pre dom i nantly of bi valves and gas tro pod de tri tus. Note con vex-up po si tion of most valves

PALAEOENVIRONMENT

The Studorski preval sec tion can be com pared to the

Werfen For ma tion of the South ern Alps that rep re sents de

-po

si

tion on a storm-in

flu

enced ramp (Brandner et al.,

2012). Sed i men ta tion of the dom i nantly fine-grained mi cro-

fa cies type B can be in ter preted as slow de po si tion in a

deeper ma rine en vi ron ment be low the fair-weather wave

base, which cor re sponds to the dis tal part of a ramp. The

pres ence of ammonoid shells in di cates a good con nec tion

with the open un re stricted sea and also im plies de po si tion in

a deeper-ramp en vi ron ment. Nev er the less, the interlayering

with lime stone of microfacies type A sug gests that slow de

-po si tion of fine-grained sed i ments (microfacies type B) was

pe ri od i cally in ter rupted by the de po si tion of coarse skel e tal

de

tri

tus dur

ing high-en

ergy events (microfacies type A).

The sharp and ero sive lower bound aries of microfacies type

A beds im ply the abrupt com mence ment of sed i men ta tion.

The well-pre served skel e tal de tri tus in the float

stones/packstones and the up wardin creas ing pro por tion of ma trix in di

-cate de po si tion dur ing storms, in a zone above the

storm-weather wave base. As the beds of microfacies type A are

over lain by lam i nated or bioturbated cal car e ous mudstones

and/or marls (microfacies type B), their de po si tion re flects

short storm events in a deeper low-en ergy ramp en vi ron ment,

where the de po si tion of lime mud pre vails. The in tense bot

-tom-shear con

di

tions dur

ing the storm peak con

cen

trated

shells of liv ing and dead or gan isms from the sea bot tom, ei

ther by bury ing them un der a sud den in flux of stormsus

-pended par ti cles or by ex hum ing pre vi ously bur ied shells by

rip ping up the un der ly ing weakly con sol i dated sed i ments and

form ing skel e tal con cen tra tions (lag de pos its). There fore,

well-pre served fos sils within a fine micritic/peloidal ma trix

of microfacies type A are in ter preted as the re sult of storm

events, which pre serve whole fos sils by bury ing and pro tect

-ing them from nor mal de struc tive pro cesses. The pres ence of

foraminifera in di cates de tri tus de rived by storms from the

shal low prox i mal ramp. Valves ori ented par al lel to the bed

-ding plane in a con vex-up po si tion also in di cate de po si tion

un der shortterm highen ergy con di tions. Fur ther more, win

now ing and sus pen sion of fine sed i ment re sulted in the de po

-si tion of the up ward-fin ing units above coarse lag de pos its.

As the storm waned, more mud was de pos ited from

suspen-ded ma te rial, re sult ing in fin ing-up ward grad ing. In tense

co-lonization by or gan isms and bioturbation could re flect the

ces sa tion of the storm and a re turn to nor mal low-en ergy

ramp sed i men ta tion be low the fair-weather wave base. The

pres ence of py rite also in di cates low-en ergy, and pos si bly

partly an aer o bic con di tions, whereas the gen e sis of ce lest ite

is con sid ered as diagenetic in deep-sea sed i ments, as

explai-ned by Baker and Bloomer (1988) al though it also has been

de scribed from shal low ma rine set tings (Hautmann, 1997).

Storms are fre quently re corded both in the in ner and outer

shelf/ramp in the Induan/Olenekian sed

i

ments of the wider

Dinaric re gion (Aljinoviæ, 1995; Aljinoviæ et al., 2006, 2011),

while this in ves ti ga tion points to de po si tion in wide mid-ramp

zone (the zone be tween the fair- and storm- weather wave

bases), as de fined by Burchette and Wright (1992).

There are only a few out crops of Lower Tri as sic strata

in the Julian Alps and they oc cur in small tec toni cally con

-fined ar eas (e.g., Mavrinc, Lipanca). There is some sim i lar ity

in their de vel op ment and macrofossil con tent (Natiria costata

and badly-pre served ammonoids). Cono dont and other mi

cro-palae on tol ogic anal y ses were neg a tive and there fore a de tailed

geo log i cal com par i son of these strata had not been possible.

PALAE ON TOL OGY

Micro fauna

The microfossil ma te rial re cov ered from nine cono dont

sam ples (SP 1–9) was ex am ined in the pres ent study. The

as so ci a tions con sisted of cono dont el e ments, rare fish re

-mains, as well as some recrystallized free spec

i

mens of

foraminifera from two sam ples (SP 1 and SP 5). In ad di tion,

sev eral thin sec tions were pre pared from each sam ple for

foraminifera study. The cono donts re cov ered are white with

CAI = 1 (Ep stein et al., 1977).

The list of microfossils de ter mined is pre sented in Ta ble 1.

Foraminifera

The foraminifer clas si fi ca tions of Loeblich and Tappan

(1988) and Mikhalevich (2000) are com

bined and used

here. The foraminifer de scrip tions are short, be cause these

foraminifers are mainly well-known and the pres er va tion of

these forms is not par tic u larly good.

Ge nus Ammodiscus Reuss, 1862

Type spe cies: Ammodiscus infimus Bornemann, 1874

Ammodiscus? parapriscus Ho, 1959

Fig. 5A–D

*1959 Ammodiscus parapriscus n. sp. – Ho: p. 408, pl. 2, figs 3–6.

Ta ble 1

Dis tri bu tion of microfossils in sam ples

from Studorski preval

Sample Taxa SP1 SP2 SP3 SP4 SP5 SP6 SP7 SP8 SP9 Ammodiscus? parapriscus Ho va a va a va va r r a Glomospirella facilis Ho va r r va a Glomospirella shengi Ho va vr a a Glomospirella sp. a r r a Pilammina? cf. praedensa Uroševiæ vr Arenovidalina? sp. vr Meandrospira? sp. vr Agathammina? sp. vr vr

Hoyenella sinensis (Ho) va a va a a a r

Dentalina splendida Schleifer vr vr vr vr

Nodosaria? spp. a vr r r vr r

Triasospathodus hungaricus

(Kozur & Mostler) a va r

fish teeth r r

Vr – very rare (1 spec i men), r – rare (2–4 spec i mens), a – abun dant (5–10 spec i mens), va – very abun dant (more than 11 spec i mens)

1976 Ammodiscus parapriscus Ho – Zaninetti, p. 89, pl. 2, figs 1–2.

partim 1983 Ammodiscus parapriscus Ho – Salaj et al., p. 61, pl. 1, ?non fig. 2, pl. 6, figs 3–5, ?non fig. 6, non 7–8. 1992 Ammodiscus parapriscus Ho – Trifonova, p. 9, pl.

1, figs 3–5, pl. 6, fig. 10.

?non1995 Cornuspira? paraprisca (Ho) – Rettori, p. 102, pl. 19, fig. 6.

Ma te rial: 75 sec tions in sam ples SP 1, 2, 3, 4, 5, 6, 7, 8, 9 and 61 free spec i mens from the sam ple SP 1.

De scrip tion: The test is cir cu lar, dis coid; pe riph ery rounded and con cave on both sides. The spher i cal proloculus is fol lowed by a closely planispiral en rolled un di vided tu bu lar sec ond cham ber, grad u ally in creas ing in di am e ter. The num ber of whorls is 3–5. The wall is ag glu ti nated and com pact. The ap er ture is at the end of the tube.

Re marks: This spe cies was as signed to Cornuspira with a ques -tion mark by Rettori (1994), but the ma te rial of the pres ent study is not wellpre served. There fore, it is not pos si ble to sup port his con -clu sion, but the ques tion mark is re tained here for def i ni tion of the ge nus. The pres er va tion of the wall of the spec i mens stud ied is not good, be cause they are recrystallized and in some cases pyritized. Strati graphic and geo graphic oc cur rence: The spe cies Ammodis cus? parapriscus was first found in the Lower and Mid dle Tri -as sic of China (Ho, 1959). This spe cies oc curs also in the Lower and Mid dle Tri as sic (Anisian) of the Dinarides, Al ba nia, Hun gary, Bul garia, Greece, Tur key, Iran, China (Zaninetti, 1976; Pantiæ-Prodanoviæ and Radoševiæ, 1977a; Dager, 1978; Oravecz-Schef-fer, 1987; Pirdeni, 1988; Salaj et al., 1988; Trifonova, 1992; Rettori, 1995) and in the Anisian and Rhaetian of the West Carpa-thians (Salaj et al., 1983).

Ge nus Glomospirella Plummer, 1945

Type spe cies: Glomospira umbilicata Cushman

& Wa ters, 1927

Glomospirella facilis Ho, 1959

Fig. 5E, F

*1959 Glomospirella facilis n. sp. – Ho, p. 414, pl. 6, figs 8–12.

1976 Glomospirella facilis Ho – Zaninetti, p. 95, pl. 2, fig. 18.

partim 1983 Glomospirella facilis Ho – Salaj et al., p. 63, pl. 2, figs 9–12, figs 14–15, ?non figs 13, 16.

1992 Glomospirella facilis Ho – Trifonova, p. 18, pl. 2, figs 13, 15–16, pl. 6, figs 13–14, 26.

Ma te rial: 21 sec tions in the sam ples SP 1, 3, 4, 5, 6, and 43 free spec i mens from the sam ples SP 1 and 5.

De scrip tion: The test is small, the equa to rial sec tion rounded, the ax ial sec tion plain or con cave on the both sides. The spher i cal proloculus is fol lowed by a streptospirally en rolled un di vided tu -bu lar sec ond cham ber and the last 2–3 whorls be come planispi-rally coiled and abruptly in creas ing in di am e ter. The last whorl is wider than oth ers and a lit tle an gu lar in cross-sec tion. The wall is ag glu ti nated, thin, and smooth. The ap er ture is at the end of the tube.

Re marks: The char ac ter is tic fea tures of this spe cies are a round form in the equa to rial sec tion and a con cave form in the ax ial sec -tion, with a very small streptospiral stage of coil ing. The above men tioned fea tures of Glomospirella facilis dif fer from those of Glomospirella shengi. The pres er va tion of the wall of the spec i -mens stud ied is not good, be cause they are recrystallized and in some cases pyritized.

Strati graphic and geo graphic oc cur rence: The spe cies

Glomo-spirella facilis was first de scribed from the Lower and Mid dle Triassic of China (Ho, 1959). Later this spe cies was re ported from the Lower and Mid dle Tri as sic (Anisian) of the Alps, Dinarides, Al ba nia, Hun gary, Bul garia, Cau ca sus, Iran (Zaninetti, 1976; Pan- tiæ-Prodanoviæ and Radoševiæ, 1977b; Pisa et al., 1979; Oravecz-Scheffer, 1987; Pirdeni, 1988; Efimova, 1991; Trifonova, 1992; Rettori, 1995) and in the Up per Tri as sic of the West Carpathians (Salaj et al., 1983).

Glomospirella shengi Ho, 1959

Fig. 5G, H

*1959 Glomospirella shengi n. sp. – Ho: p. 413, pl. 5, figs 20–25.

1976 Glomospirella shengi Ho – Zaninetti, p. 104, pl. 2, figs 14–16.

partim 1983 Glomospirella shengi Ho – Salaj et al., p. 65, pl 3, figs 1–10, 13-16, non 11–12, pl. 5, figs 10–11, 15–16, ?non figs 12–14

partim 1992 Glomospirella shengi Ho – Trifonova, p. 18, pl. 2, fig. 4, pl. 6, figs 20, 27, ? non fig. 28.

Ma te rial: 17 sec tions in the sam ples SP 1, 3, 5, 6, and 30 free spec i mens from the sam ples SP 1 and 5.

De scrip tion: The test is small, equa to rial sec tion oval, ax ial sec -tion mainly plain or con vex on the both sides. The spher i cal proloculus is fol lowed by a streptospirally en rolled un di vided tu -bu lar sec ond cham ber and the last 2–3 whorls be come planispi-rally coiled and grad u ally in creas ing in di am e ter. The streptospiral stage of coil ing is wider in the ax ial sec tion than the last whorl. The wall is ag glu ti nated, thin, and smooth. The ap er ture is at the end of the tube.

Re marks: The char ac ter is tic fea tures of this spe cies are the oval form in the equa to rial sec tion and the con vex or plain form in the ax ial sec tion, the streptospiral stage of coil ing is wider in the ax ial sec tion than the last whorl. The above men tioned fea tures of Glomospirella shengi are dif fer ent from those of Glomospirella facilis. The pres er va tion of the wall of the spec i mens is not good, ow ing to recrystallization or pyritization.

Strati graphic and geo graphic oc cur rence: This spe cies was first found in the Lower and Mid dle Tri as sic of China (Ho 1959). Later the spe cies was de scribed from Lower and Mid dle Tri as sic (Ani-sian) of the Dinarides, Hun gary, Bul garia, Tur key, Cau ca sus (Pan- tiæ, 1970; Uroševiæ and Jelièiæ, 1973–1974; Zaninetti, 1976; Dager, 1978; Oravecz-Scheffer, 1987; Efimova, 1991; Trifonova, 1992; Rettori, 1995) and from the Rhaetian of the West Carpa-thians (Salaj et al., 1983).

Ge nus Pilammina Pantiæ, 1965

Type spe cies: Pilammina densa Pantiæ, 1965

Pilammina? cf. praedensa Uroševiæ, 1988

Fig. 5I

*1988 Pilammina praedensa n. sp. – Uroševiæ: p. 372, pl. 1, figs 1–6.

partim 1995 Pilammina praedensa Uroševiæ – Rettori, p. 56, pl. 3, figs 1–3, 5, ?non fig. 4.

Ma te rial: One sec tion in the sam ple SP 6.

De scrip tion: The test is free. It is close-coiled, con sist ing of a round proloculus and an other inseptate, tu bu lar chamberlet. The ini tial wind ing cov ers two and a half to three close coils. The last whorl is much wider than first whorls.

Re marks: The sec tion stud ied is sim i lar to the holotype. The pres -er va tion is not good, be cause the test is not com plete. Th-ere fore there is no pos si bil ity to de scribe the in ner struc ture in de tail. Usu

-ally the pres er va tion of the wall of this spe cies is not good and there is no con fi dence that this spe cies be longs to Pilammina and this is true for the stud ied spec i men too.

Strati graphic and geo graphic oc cur rence: Spe cies Pilammina praedensa was first de scribed from the Lower Tri as sic of the Dinarides (Uroševiæ, 1988) and af ter it was found in the up per part of Olenekian of Greece and the Lower Tri as sic of China (Rettori, 1995).

Ge nus Hoyenella Rettori, 1994

Type spe cies: Glomospira sinensis Ho, 1959

Hoyenella sinensis (Ho, 1959)

Fig. 5M–O

*1959 Glomospira sinensis n. sp. – Ho, p. 410, pl. 3, figs 16–34.

1959 Glomospira sinensis var. rara, n. sp. n. var. – Ho, p. Fig. 5. Foraminifera from Early Tri as sic of Studorski preval, Slovenia. Scale bar 200 µm (A–O), 50 µm (P, R). A–D. Ammodiscus? parapriscus Ho, 1959, A. SP 7, equa to rial sec tion. B. SP. 1, equa to rial sec tion. C. SP 6, equa to rial sec tion. D. SP 3, ax ial sec tion. E, F. Glomospirella facilis Ho, 1959, E. SP 6, F. SP 1, subequatorial sec tions. G, H. Glomospirella shengi Ho, 1959, G. SP 6, H. SP 6, subequatorial sec tions. I. Pilammina? cf. P. praedensa Uroševiæ, 1988, SP 6, oblique sec tion. J. Arenovidalina? sp., SP 3, oblique sec tion. K. Meandrospira? sp., SP 9, subequatorial sec tion. L. Agathammina? sp., SP 2, subtransverse sec tion. M–O. Hoyenella sinensis (Ho, 1959), M. SP 1, subaxial sec tion. N. SP 2, ax ial sec tion. O. SP 3, subaxial sec tion. P, R. Dentalina splendida Schleifer, 1961, P. SP 2. R. SP 3, sublongitudinal sec tions

410, pl. 4, figs 1–2.

1972 Glomospirella elbursorum n. sp. – Brönnimann, Zaninetti, Bozorgnia and Huber, p. 9, pl. 3, figs 1–10.

partim 1974 Glomospirella elbursorum Brönnimann, Zaninetti, Bozorgnia and Huber – Baud et al., pl. 30, figs 4–6, 8–11, non figs 7, 12, pl. 31, figs 1–6, 8–10, non fig. 7. partim 1975 Glomospirella sp. – Zaninetti and Brönnimann, pl.

36, figs 1, 3–6, 10–12, non fig. 9. 1994 Hoyenella sinensis Ho – Rettori, p. 342.

partim 1995 Hoyenella gr. sinensis Ho – Rettori, p. 126, pl. 24, figs 1–15, pl. 25, figs 1–3, 5, 8, non figs 4, 6–7, 9, pl. 26, figs 1–3, 6–8, 10–12, 14, non 4–5, 9, 13, 15–17, pl. 27, figs 5–7, 10, non 1–4, 8–9, 11.

Ma te rial: 29 sec tions in the sam ples SP 16, 9 and 10 free spec i -mens from the sam ples SP 1 and SP 5.

De scrip tion: The test is free and small, mainly el lip soi dal in shape, the spher i cal proloculus is fol lowed by an un di vided tu bu -lar sec ond cham ber, grad u ally in creas ing in di am e ter. In coil ing, early whorls are miliolid-like, later ones are short with 1–2 plani-spi ral whorls. The wall is dark and com pact in trans mit ted light. The ap er ture is sim ple, ter mi nal.

Re marks: Glomospirella elbursorum Brönnimann, Zaninetti, Bo-zorgnia and Huber (1972) could be placed into syn on ymy with the spe cies stud ied, be cause of the very sim i lar struc ture of the test and in this point, the de ci sion of Rettori (1994) is fol lowed. Glo-mospirella sp. (Zaninetti and Brönnimann, 1975) is placed into syn on ymy with the spe cies stud ied, ac cord ing to the mor pho log i -cal fea tures of this spe cies. The pres er va tion of the wall of the spec i mens stud ied is not good ow ing to recrystallization or pyri-tization.

Strati graphic and geo graphic oc cur rence: The spe cies Hoye-nella sinensis was first found in the Lower and Mid dle Tri as sic of China (Ho, 1959). More over, it is known from the Lower and Mid -dle Tri as sic (Anisian) of the Alps, Dinarides, Al ba nia, Hun gary, Bul garia, Greece, Tur key, Cau ca sus, Iran, Pa ki stan, Himalaya (Dimitrijeviæ et al., 1968; Baud et al., 1974; Zaninetti and Brön-nimann, 1975; Zaninetti, 1976; Pantiæ-Prodanoviæ and Radoševiæ, 1977a, b; Dager, 1978; Pisa et al., 1979; Kristan-Tollmann, 1984; Oravecz-Scheffer, 1987; Pirdeni, 1988; Salaj et al., 1988; Efimova, 1991; Trifonova, 1992; Rettori, 1995) and in the Up per Tri -as sic of the West Carpathians and Bul garia (Salaj et al., 1983; Trifonova, 1992).

Ge nus Dentalina Risso, 1826

Type spe cies: Nodosaria (Dentaline) cuvieri

(d’Orbigny, 1826)

Dentalina splendida Schleifer, 1961 in Gerke (1961)

Fig. 5P, R

*1961 Dentalina splendida n. sp. – Schleifer in Gerke: p. 233, pl. 29, figs 5–6.

1991 Dentalina splendida Schleifer – Efimova, pl. 1, fig. 7. 1994 Dentalina splendida Schleifer – Trifonova, p. 37, pl. 4,

figs 7, 11–12, 18.

Ma te rial: Four sec tions in the sam ples SP 1, SP 2, SP 3 and SP 8. De scrip tion: The small test is al most rec ti lin ear or slightly cur-ved. The proloculus is usu ally oval. The num ber of elon gated oval cham bers is from 4 to 8. They usu ally in crease very grad u ally in size and are, com par a tively loosely jointed in all cham bers. The su tures are de pressed. The wall is very thin. The ap er ture is sim -ple, ter mi nal.

Re marks: The typ i cal fea ture of this spe cies is usu ally the oval prolocolus and other cham bers are oval and elon gated. The

holotype is bro ken, with out a proloculus. The pres er va tion of the spec i -mens is not good, be cause they are mainly recrystallized, but there is prolocolus pres ent in all spec i mens.

Strati graphic and geo graphic oc cur rence: Spe cies Dentalina splendida was first de scribed from the lower part of the Olenekian of the North Si be ria (Gerke, 1961). Later this spe cies was found in the Olenekian of the North Cau ca sus and Precaucasus (Efimova, 1991), and in the up per part of the Lower Tri as sic and in the lower part of the Anisian of Bul garia (Trifonova, 1994).

Re marks on foraminifer fauna. The foraminifer assemblage con -sists of Ammodiscus? parapriscus Ho, Glomospirella facilis Ho, Gl. shengi Ho, Gl. sp., Pilammina? cf. P. praedensa Uroševiæ, Areno-vidalina? sp., Meandrospira? sp., Agathammina? sp., Hoyenella sinensis (Ho), Dentalina splendida Schleifer, Nodosaria? spp. (Fig. 5A–R). Spe cies of the gen era Ammodiscus, Glomospirella and Hoyenella are dom i nant. An over view on the foraminifer taxa de ter -mined is given in Ta ble 1.

The main fea ture of the tax o nomic com po si tion of this fora-minifer as sem blage from the Studorski preval lo cal ity in the Julian Alps of west ern Slovenia is the dom i nance of prim i tive ag glu ti -nated foraminifers (Ammodiscus and Glomospirella) and some miliolids and nodosariids. The foraminifers are very small and not wellpre served. The ge neric com po si tion of this foraminifer as -sem blage is mainly sim i lar to the com po si tion of the Early Tri as sic and Anisian as sem blages from dif fer ent parts of the Tethys, from the Alps to China. Sev eral au thors have in di cated that there are “GlomospiraGlomospirella” foraminifer com mu ni ties at dif fer -ent lev els of the Lower Tri as sic and the lower Anisian in the Te-thyan Realm (Zaninetti, 1976; Salaj et al., 1983; Trifonova, 1992; Vuks, 2007). The foraminifer as sem blage from the Studorski preval lo cal ity has some spe cies com mon with the forami nifer as -sem blages from the Lower Tri as sic of China (He, 1993), the Olenekian of the Cau ca sus area (Efimova, 1991), the Lower Tri as -sic and Anisian of Bul garia (Trifonova, 1992) and Hun gary (OraveczScheffer, 1987), and it can be cor re lated with foraminifer as -sem blages from these ar eas. In the Alps and the Dinarides the “Glomospira-Glomospirella” foraminifer com mu ni ties with some miliolids and nodosariids are known mainly from the Lower Tri as -sic (Dimitrijeviæ et al., 1968; Pantiæ, 1970; Pantiæ-Prodanoviæ and Radoševiæ, 1977b; Pisa et al., 1979; Resch, 1979; Broglio Loriga et al., 1990).

Ear lier in west ern Slovenia, the “Glomospira-Glomospirella” foraminifer as sem blages with some nodosariids were re ported from the Lower Tri as sic and Anisian (Ramovš, 1972). The fora-minifer as sem blage from west ern Slovenia is mainly sim i lar to the foraminifer as sem blages men tioned above, be cause they have a com mon ge neric com po si tion. In east ern Slovenia “Glomospira-Glomospirella” foraminifer as sem blages are known from the Ani-sian (Anièiæ and Dozet, 2000).

In only one thin-sec tion a Meandrospira-like spec i men has been de tected, but the pres er va tion of the in ter nal struc ture is poor. In Tri as sic time, the rep re sen ta tives of Meandrospira are typ i cal for the Early Tri as sic and the Anisian.

It is rea son able to con clude that the foraminifer as sem blage from the Studorski preval lo cal ity of the Julian Alps in west ern Slovenia oc curred in Early Tri as sic–Anisian time, or that it was prob a bly re stricted to the up per part of the Early Tri as sic.

Cono donts

The cono dont tax on omy fol lows the clas si fi ca tion of

Or chard (2005).

Ge nus Triassospathodus Kozur, 1998 in Kozur, Mostler

and Krainer (1998)

Triassospathodus hungaricus (Kozur and Mostler, 1970)

Fig. 6A–G

*1970 Spathognathodus hungaricus n. sp. – Kozur and Mos-tler: p. 453, pl 4, figs 8–11.

Ma te rial: Five Pel e ments and eleven ramiform el e ments (sam -ples SP 1, 3, 5) of mod er ate pres er va tion.

De scrip tion: The spathodid P-el e ments ex am ined are sim i lar in length and bear three denticles. The basal cav ity is wide and open. How ever, part of the pos te rior mar gin in the il lus trated spec i men is bro ken in the aboral view (Fig. 6D). It is ob vi ous that the el e ments have an ex panded basal cav ity.

The ramiform el e ments are mostly frag mented and they do not per mit re con struc tion. Some of them are de ter mined as ?M, ?S1 and S3 el e ments.

Re marks: Triassospathodus can be eas ily dis tin guished from Neospathodus Mosher, 1968, which has a ter mi nally up warddi -rected lower mar gin of the basal cav ity (Mosher, 1968). On the other hand, there ex ists a morphologic sim i lar ity in the P el e ments of Triassospathodus Kozur, 1998 (in Kozur et al., 1998) and Novispathodus Or chard, 2005 (Or chard, 2005); the lat ter has a shorter and more wid ened basal cav ity and it has also a dif fer ent ap pa ra tus com po si tion (Heinz Kozur, pers. comm., 2012). From the Spathian strata of the Thaynes Group in Ne vada, Lucas and Or chard (2007) re ported el e ments that are mor pho log i -cally close or iden ti cal with “Neospathodus” cf. hungaricus. Strati graphic and geo graphic oc cur rence: Spe cies T. hungaricus was first de scribed from the Tirolites beds of Felsõrs in Hun -gary (Kozur and Mostler, 1970). Triassospathodus is an im por tant Lower Tri as sic ge nus that com prises the ma jor ity of Up per Olene-kian (Spathian) in dex spe cies (Kozur et al., 1998).

In an in te grated ammonoid and cono dont zonation of the Tri as sic, Kozur (2003) pre sented T. hungaricus as the marker taxon for the basal Spathian equiv a lent to the Tirolites cassianus ammonoid Zone. Ac cord ing to H. Kozur (pers. comm., 2012), in the shal low west ern Tethys the T. hungaricus fauna lies within the lower Spathian, where the fauna with Icriospathodus collinsoni is mis-sing.

There are sev eral re cords of Early Tri as sic cono donts from Slo-venia, but most of them are older than Spathian (see Kolar-Jur-kovšek et al., 2011). Re cov ery of the spathodid Spathian fauna has been doc u mented only from the bor der land of the Ljubljana de -pres sion in cen tral Slovenia (Dozet and Kolar-Jurkovšek, 2007). In Slovenia, a cono dont fauna, high in spathodid el e ments and with few denticles, is abun dant in the Spathian of Slovenia (Idrija-Žiri and Krško ar eas; un pub lished data). From the Ex ter nal Dinarides, the Muæ sec tion in Croatia was pro posed as a stan dard sec -tion for the Eu ro pean Up per Scythian. This sec -tion is well-known for its rich macrofauna and also con tains Triassospathodus trian-gularis (Herak et al., 1983).

Macrofauna

The macrofauna col lected con tains bi valves and gas tro

-pods, as well as a few badly pre served spec i mens of

ammo-noids of the ge nus Tirolites that in di cate an Early Spathian

age.

Bi valves

Ge nus Bakevellia King, 1848

Type spe cies: Avicula antiqua Graf zu Münster in

Goldfuss, 1836 [non A. antiqua Defrance] = Avicula

binneyi Brown, 1841, by orig i nal des ig na tion

Bakevellia cf. incurvata (Lepsius, 1878)

Fig. 7A–C

*1878 Gervillia incurvata sp. n. – Lepsius: p. 353, pl. 1, figs 3a–b.

1908 Gervillia incurvata Lepsius – Wittenburg, p. 31, pl. 4, fig. 7.

1923 Gervillia incurvata Lepsius – Diener, p. 91 (cum syn.). Ma te rial: One well-pre served spec i men with con joined valves Fig. 6. Cono donts from Early Tri as sic of Studorski preval, Slovenia. A–G. Triassospathodus hungaricus (Kozur and Mostler, 1970). A–D. P1 el e ment (up per, lower, lat eral views), SP 1. E–G. ramiform el e ments (?M, ?S1, S3 el e ments), SP 3

(BJ 2766), one left valve and one spec i men with con joined valves (BJ 2763), ca. 40 mod er ately pre served spec i mens mostly with con joined valves (BJ 2764).

De scrip tion: Rel a tively slen der, pos te ri orly elon gated Bakeve-llia, with left and right valve more or less equally con vex. Beaks pointed, pro ject ing above hinge mar gin in both valves.

Re marks: This spe cies is un usual for its ge nus in hav ing more or

less equally con vex left and right valves. On this ba sis, the ma te -rial is ten ta tively as signed to Bakevellia incurvata, al though there are slight dif fer ences in the out line by com par i son with the sin gle right valve, fig ured by Lepsius (1878, pl 1, figs 3a, b). Lepsius (1878, p. 353) in di cated that the beak of the right valve did not pro ject above the hinge line, whereas in the ma te rial de scribed, it does. An other dif fer ence to the de scrip tion by Lepsius (1878) is Fig. 7. Bi valves from Early Tri as sic of Studorski preval, Slovenia. Scale bar 1 cm. All spec i mens black ened with graph ite emul sion and coated with am mo nium chlo ride. A–C. Bakevellia cf. incurvata (Lepsius, 1878). A. shell ac cu mu la tion of mostly con joined valves, BJ 2764. B. Spec i men with con joined valves, an te rior shell end to right, BJ 2766. Note ori en ta tion with valves slightly opened and dor sal mar gin up side. See text for ex pla na tion. C. Left valve, BJ 2763. D. Avichlamys sp., one left valve (in di vid ual on right) and two valves with un known side ori en ta tion, BJ 2765. E. Eumorphotis cf. gronensis (Wittenburg, 1908), left valve, BJ 2766

the ap par ent lack of a shell sulcus at the ven tral mar gin, but the intraspecific vari abil ity of this fea ture is un clear.

A pe cu liar ity of the pres ent ma te rial is an un usual ori en ta tion of spec i mens that have both valves con joined. Fre quently, these spe-cimens have their valves opened up to 30° and rest on the ven tral valve mar gins, i.e. the hinge mar gin lies in a dor sal po si tion, more or less par al lel to the sub strate. It is un likely that this was the life po si tion of the an i mal, be cause con struc tional mor phol ogy (elon -gated shape, long hinge axis, sim i lar con vex ity of valves) in di cates life as an endobyssate mud sticker with the hinge axis ori ented more or less ver ti cally to the sed i ment sur face (Seilacher, 1984, text-fig. 7; Mus ter, 1995, p. 98). Be cause the fauna oc curs at the top of a tempestitic bed, as in di cated by hummocky crossstrat i fi -ca tion, a pos si ble ex pla na tion is that the an i mals were ex -ca vated and dis placed by storm waves and then smoth ered by the re set tling silt-clay sus pen sion. The ob served ori en ta tion in the Studorski preval material might thus be an unnatural dying position of this species.

Ge nus Eumorphotis Bittner, 1901

Type spe cies: Pseudomonotis telleri Bittner,

1898 by sub se quent des ig na tion (Cossmann, 1902)

Eumorphotis cf. gronensis (Wittenburg, 1908)

Fig. 7E

*1908 Mysidioptera gronensis sp. n. – Wittenburg: p. 31, pl. 4, fig. 4.

Ma te rial: One left valve with a partly pre served ex ter nal shell layer (BJ 2766), ques tion able three abraded left valves (BJ 2701). De scrip tion: Very large Eumorphotis with about 70 ra dial ribs near the shell mar gin. Ribs nodose, in creas ing in num ber by in ter -ca la tion dur ing on tog eny. Out line of shell typ i -cal of the ge nus, but pos te rior wing rel a tively elon gated.

Re marks: The larg est spec i men is about 8 cm long and more than 8 cm high. This is to the knowl edge of the au thors the larg est Eumorphotis spec i men, re ported so far from the Early Tri as sic. Al though the type ma te rial of E. gronensis (Wittenburg, 1908) is in com plete, the spec i men in the pres ent study is ten ta tively as -signed to that spe cies on the ba sis of the sim i lar or na men ta tion. The shell ex te rior of the spec i men is dor sally eroded, but an in -crease in the num ber of ribs by in ter ca la tion of ad di tional ribs is ob served on the pos te rior wing. There fore, it is as sumed that the higher num ber of ribs in this spec i men (ca. 70) by com par i son with the holotype of E. gronensis (28) could be re lated to its larger size. Spe cies E. kittlii Bittner, 1901 dif fers from E. gronensis chiefly in its stron ger de vel op ment of the ten to twelve most pos te rior ra -dial ribs, which cover the pos te rior wing and the ad ja cent part of the disc. Al though the pres ent au thors cur rently treat both spe cies sep a rately, they wish to note that the dif fer ence in or na men ta tion might turn out to be a pres er va tion ef fect or as ly ing within the range of intraspecific vari a tion. In this case, E. kittlii would have pri or ity over E. gronensis and spe cies as sign ment of the spec i men would change ac cord ingly (see also Broglio Loriga and Mirabella, 1986).

Ge nus Aviclamys Allasinaz, 1972

Type spe cies: Pecten csopakensis Frech, 1905

by orig i nal des ig na tion

Aviclamys? sp.

Fig. 7D

Ma te rial: One com plete left valve and two in com plete valves (BJ 2765).

De scrip tion: Disc nearly cir cu lar, cov ered with about ten strong first-or der ra dial ribs plus the same num ber of in ter ca lated weaker sec ond-or der ribs. The pos te rior au ri cle is with out a si nus. Re marks: The state of pres er va tion does not al low spe cific de ter mi na tion of this taxon. How ever, the cir cu lar disk, trun cated pos -te rior au ri cle and style of or na ment are typ i cal of Avichlamys. Pecten vajolettensis Wittenburg, 1908, sub se quently re ferred to Avichlamys by Allasinaz (1972, p. 225), has a sim i lar or na ment. How ever, the pres ence of an elon gated an te rior au ri cle, as fig ured by Wittenburg (1908, pl. 1, fig. 9), can not be ver i fied in the ma te -rial de scribed.

Gas tro pods

Ge nus Natiria de Koninck, 1881

Type spe cies: Natiria lyrata Phillips, 1836 by monotypy

(de Koninck, 1881, p. 5)

?Natiria costata (Münster, 1841)

Fig. 8A, B

*1841 Naticella costata sp. n. – Münster: p. 101, pl. 10, fig. 14. 1892 Naticella costata Münster – Kittl, p. 67.

1967 Naticella costata Münst. – Leonardi, pl. 20, figs 15–16. 1985 Natiria costata (Münster) – Neri and Posenato, p. 91. 2002 Natiria costata (Münster) – Hips and Pelikán, p. 356,

fig. 8A.

2005 Natiria costata (Münster) – Nützel, p. 440, fig. 6. Type lo cal ity: Most likely Groneshof, South Tirol, It aly; Werfen For ma tion, Olenekian, Early Tri as sic (com pare Kittl, 1892: 67). Ma te rial: Sin gle rock piece, com posed of gas tro pod co quina from Studorski preval. The as so ci a tion is monospecific, the shells are recrystallized and mostly weath ered out from the vis i ble sur faces. Sam ple BJ 2480.

Re marks: In spite of the mass oc cur rences of this gas tro pod in the Werfen For ma tion, its tax o nomic – both ge neric and spe cific – sta -tus re mains un clear. The type spe cies of Natiria was de scribed from the Visean (Lower Car bon if er ous) of York shire (Phillips, 1836), while the spe cies Natiria costata was de fined by Münster (1841) in a mono graph on gas tro pods from the Carnian (Up per Fig. 8. Neritimorph gas tro pod ?Natiria costata (Münster, 1841) from Early Tri as sic of Studorski preval, Slovenia, BJ 2480. A. Gen eral view of rock frag ment with nu mer ous poorly pre served in di vid u als. B. Close-up of the spec i men ar rowed on A, show ing mode of pres er va tion

Tri as sic) St. Cassian For ma tion. It seems, how ever, that the type se ries of N. costata came from a lo cal ity of the Werfen For ma tion, rather that of the Cassian Fm. Kittl (1892, p. 67) sug gested that the ma te rial of Münster (1841) could have come from an out crop of the Werfen Fm at Groneshof, where it oc curs abun dantly. Tak ing into ac count ap prox i mately 100 mil lion years and the PT ex tinc -tion event be tween the oc cur rences of N. lyrata and N. costata, it is doubt ful that both forms are con ge neric (see also dis cus sion in Yin and Yochelson, 1983, p. 523). Un for tu nately, de spite the ubiq uity of the ma te rial from the Lower Tri as sic, no protoconch of N. costata has been dis cov ered yet.

The ma te rial re cov ered at Studorski preval is poorly pre served and does not con trib ute any new mor pho log i cal in for ma tion to the ex ist ing de scrip tions. In fact the iden ti fi ca tion it self is poorly con sti tuted and only con di tional, on the ba sis of gross shell mor phol -ogy. Nev er the less, no other gas tro pod of this mor phol ogy and size is known from the Werfen For ma tion and there fore the iden ti fi ca tion seems to be plau si ble and it is yet an other piece of data, con -firm ing the wide dis tri bu tion of this gas tro pod in the Early Tri as sic Tethys.

Strati graphic and geo graphic oc cur rence: Natiria costata along with Werfenella rectecostata are the most typ i cal gas tro pods of the Early Tri as sic Werfen For ma tion (e.g. Nützel, 2005). These two spe cies oc cur in Eu rope, co in cid ing roughly with out crops of the up per Werfen For ma tion or its equiv a lents. Oc cur rences in clude in It aly (e.g., Leonardi, 1967; Neri and Posenato, 1985, Nützel, 2005), Hun gary (Broglio Loriga, 1990; Hips and Pelikán, 2002), Aus tria (Diener, 1926; Kutassy, 1940), Croatia (Prlj-Šimiæ, 2006), and Slovenia (Pavšiè, 2003; this pa per). Re cently, poorly pre -served Natiria cf. costata has also been re ported from the Spathian of Utah, USA (Hofmann et al., 2013a, b).

DIS CUS SION AND CON CLU SIONS

This pa per pres ents de tailed doc u men ta tion and in ter

-pre ta tion of sedimentological data on a re mark able Early

Tri as sic fos sil site in the Julian Alps of Slovenia. The re port

on the palaeontological ma

te

rial, col

lected at Studorski

preval, con firms the wide dis tri bu tion of mol lusk and

fora-minifer taxa in the Early Tri as sic Tethys. The re cov ery of

the ear li est Spathian cono dont fauna with

Triassospatho-dus hungaricus in di cates an Olenekian age of the fauna and

is an im por tant cor re la tive el e ment for com par i son with co

-eval fau nas in the Dinarid area and world wide.

The sedimentological data pre sented here in di cate that

the strata of Studorski preval in the Julian Alps of west ern

Slovenia were de pos ited in a more dis tal, storm-in flu enced

ramp. A ramp set ting was also men tioned for the de po si tion

of the Werfen For ma tion in the South ern Alps (Brandner et

al., 2012). Two types of beds were rec og nized, des ig nated

here as microfacies types A and B. These al ter nate ver ti cally

and form a sed i men tary ar chi tec ture that con sists of

coarse-grained skel e tal lag de pos its of microfacies A, over lain by

the lam

i

nated or bioturbated mudstones and/or marls of

microfacies B. Units with such or ga ni za tion im ply the oc ca

-sional win now ing of bot tom sed i ments dur ing storm peaks

and the de po si tion of coarse-grained skel e tal ma te rial as the

storms waned, fol lowed by the slow set tling out of

fine-grained ma te rial dur ing in ter vals with out storms.

Ammo-noids en

coun

tered in microfacies B im

ply a bar

rier-free

con nec tion of the depositional en vi ron ment with the open

sea, i.e. de po si tion in a more dis tal, deeper-ramp set ting.

Nu mer ous trace fos sils in the mudstones and marls of mi

-crofa cies B type beds sug gest de po si tion in a low-en ergy

deeper part of the ramp. A lowen ergy, poorly aer ated en vi

ron ment is also in di cated by the pres ence of py rite and ce

lest ite. Pe ri odic re work ing of the bot tom skel e talrich sed i

-ment and the sed i men ta tion of storm beds suggest

deposi-tion in a zone above the storm wave base, possibly in a wide

low-energy mid-ramp environment.

Foraminifer as sem blages mainly con sist of the prim i

tive ag glu ti nated foraminifers, typ i cal for the Lower Tri as

sic and the Anisian. The foraminifer as sem blage is rep re

-sented by Ammodiscus? parapriscus Ho, Glomospirella

fa-cilis Ho, Gl. shengi Ho, Gl. sp., Pilammina? cf. praedensa

Uroseviæ, Arenovidalina? sp., Meandrospira? sp.,

Agatha-mmina? sp., Hoyenella sinensis (Ho), Dentalina splendida

Schleifer and Nodosaria? spp. The main fea ture of the tax o

-nomic com po si tion of this foraminifer as sem blage is the

dom i nance of prim i tive ag glu ti nated foraminifers

(Ammo-dis cus, Glomospirella, Hoyenella), as so ci ated with some

miliolids and nodosariids. It can be as signed to the time

range from Early Tri as sic to Anisian

“Glomospira-Glomo-spirella” foraminifer com mu nity. An Early Tri as sic age is

more plau si ble.

The mol lusk fauna in cludes taxa com pa ra ble to those of

the Tirolites beds of the (Up per) Werfen Fm. Gas tro pods

form monospecific coquinas that are most likely of the

shells of Natiria costata, a typ i cal faunistic el e ment of the

Tethyan Early Tri as sic. The bi valve fauna in cludes three ge-

nera that dif fer in strati graphi cal range and geo graph ical

dis tri bu tion. Bakevellia is a longlived ge nus, known world

-wide from the Perm ian up to the Cre ta ceous . Eumorphotis

was cos mo pol i tan in the Early Tri as sic and reached its acme

in the Olenekian, al though the ge nus might have per sisted

lo cally into the Mid dle Tri as sic. In con trast, Avichlamys

was re stricted to the Olenekian and is cur rently known only

from the west ern Tethys. The bi valve spe cies in clude

Bake-vellia cf. incurvata, Eumorphotis cf. gronensis and

Aviclamys sp. One of the spec i mens of Eumorphotis dis plays un

-usu ally large di men sions, seem ingly con sti tut ing a re cord

for the larg est size of this ge nus, so far re ported from the

Early Tri as sic. The un usual po si tion of ar tic u lated

Bakevellia spec i mens is in ac cor dance with stormevent de po si

tion, in ferred from sedimentological ob ser va tions. By com

-par i son with other re cently de scribed bi valve fau nas from

the Spathian (Neri and Posenato, 1985; Wasmer et al.,

2012; Hautmann et al., 2013) and even from the

Griesba-chian (Kumage and Nakazawa, 2009; Hautmann et al.,

2011), the bi valve di ver sity is low in the Studorski preval

sec tion, and no infaunal taxa are pres ent. How ever, the nor

-mal to large growth sizes of the spec i mens in di cate that this

low di ver sity is not re lated to en vi ron men tal stress. Rather,

it may be re lated to the rel a tively greater wa ter depth, pro

-vided that the spec i mens have not been trans ported over

very large dis tances, which is un likely given the low level

of frag men ta tion. Miller (1988) dem on strated an evo lu tion

ary on shoreoff shore trend in the Palaeozoic his tory of bi

valves, in which taxa mainly evolved in shal low ma rine set

tings and ex panded later into deeper/dis tal set tings. This on

shoreoff shore or prox i maldis tal trend can be eas ily ap

-plied to the weakly in clined ramp sit u a tion, as de picted for

the site in ves ti gated. The low bi valve di ver sity in the rel a

-tively deeper dis tal set ting de scribed here may thus re flect

an evo lu tion ary de lay of re-di ver si fi ca tion in dis tal set tings,

com pared to the more di verse fau nas in shal lower set tings

of the same age (e.g., Neri and Posenato, 1985; Hautmann et

al., 2013). The data in di cate a gen eral evo lu tion ary trend

that is on shore-off shore (see also Jablonski et al., 1983),

seldom considered in analyses of the timing of biotic

reco-very after mass extinctions.

Ac knowl edge ments

The au thors thank the Triglav Na tional Park (Triglavski naro- dni park) for per mis sion to col lect sam ples at Studorski preval (no. 35607-1/2011-2 dated 9th June 2011). Com mu ni ca tion with Heinz Kozur on the cono dont fauna is ap pre ci ated. Alex Nützel (Münich, Ger many) is ac knowl edged for the dis cus sion on N. costata. The study was partly sup ported by the Slovenian Re search Agency (Programme Num ber P1-0011 and Pro ject Num ber J1-9465) and Cro atian Min is try of Sci ence (Pro ject Num ber 195-0000000-3202). The re search of AK was sup ported by the Al ex an der von Humboldt Foun da tion Re turn Fel low ship and the In sti tute of Paleobiology of the Pol ish Acad emy of Sci ences, War saw. The au thors thank two anon y mous re view ers and ed i tor for care ful read -ing and thought ful com ments. This is a con tri bu tion to IGCP-Pro ject 572 (“Re cov ery of eco sys tems af ter the Perm ian–Tri as sic mass ex tinc tion”).

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