Olenekian (Early Triassic) fossil assemblage from eastern Julian Alps (Slovenia)

Annales Societatis Geologorum Poloniae (2013), vol. 83: 213–227.

OLENEKIAN (EARLY TRI AS SIC) FOS SIL AS SEM BLAGEFROM EAST ERN JULIAN ALPS (SLOVENIA)

Tea KOLAR-JURKOVŠEK1, Valery J. VUKS2, Dunja ALJINOVIÆ3, Mi chael HAUTMANN4,Andrzej KAIM5 & Bogdan JURKOVŠEK1

1 Geo log i cal Sur vey of Slovenia, Dimièeva ulica 14, 1 000 Ljubljana, Slovenia; e-mail: [email protected],[email protected]

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 [email protected]

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: [email protected]

4 Paläontologisches Institut und Mu seum, Karl Schmid-Strasse 4, 8006 Zürich, Swit zer land;e-mail mi [email protected]

5 Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, PL-00-818 Warszawa, Po land;e-mail: [email protected]

Kolar-Jurkovšek, T., Vuks, V. J., Aljinoviæ, D., Hautmann, M., Kaim, A. & Jurkovšek, B. 2013. Olenekian (EarlyTri 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 JulianAlps 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 tivelydeeper, 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 coarse-grained 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 thefor 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 stormwave 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 portedrel 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 alsooc cur. In the foraminifer as sem blages, spe cies of Ammodiscus, Hoyenella, Glomospirella dom i nated, cor re spon-d ing to the wide spread “Glomospira-Glomospirella” foraminifer com mu nity, with some miliolids and 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 theTethys 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 rineset 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 mentsthrough 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 inthe Phanerozoic his tory of ma rine life, ter mi nat ing the in -cum bency of the “Palaeozoic Evo lu tion ary Fauna” that haddom i nated life in the sea for nearly 250 mil lion years (Sep-koski, 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 fromthe 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 tilto day (Sepkoski, 1997). The re cov ery from the end-Perm -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 ableprereq ui site in this con text is a cen sus of the fau nas thatlived and evolved dur ing the Early Tri as sic, i.e., the ep ochthat fol lowed the cri ses. Thus, col lect ing and in ter pret ingnew 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 havebeen 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 palaeontological 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 howdiversity 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 inthe 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 isrep 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. Theyusu 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 ofthese strata is in tec tonic con tact with the Norian–RhaetianDachstein 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). Thicklime mudstones and marls and thin beds en riched in coarse-grained 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 sistsmainly of foraminifer as sem blages in as so ci a tion with rarecono dont el e ments and fish re mains.

Li thol ogy

The Studorski preval sec tion mainly con sists of thickmudstones (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

214 T. KOLAR-JURKOVŠEK ET AL.

Fig. 1. Lo ca tion of Lower Tri as sic strata at Studorski preval in Julian Alps (Slovenia)

is con cen trated at the base and grad u ally passes up ward into mudstones/marls. Thus, the microfacies in the Studorskipreval sec tion cor re spond to two microfacies types: mi cro-fa cies type A, fossiliferous float stone/packstones, and mi -crofacies type B, dense or lam i nated lime mudstones thatmay pass grad u ally into marls.

Centi metre- to decimetre-thick fossiliferous float stone/packstone beds (A) have dis tinc tive sharp lower ero sionalbound aries. The float stone/packstone interbeds gradatio-nally 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 oflarge 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 ofpeloidal par ti cles and some micrite (Fig. 4E). Whole valvesare 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 sparrycal cite. Some shel ter pores, at the con cave side of bi valveshells, are also infilled by sparite.

Microfacies type B con sists of dense, lam i nated or bio-turbated 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-richlaminae. Dis tur bance of the lam i na tion is due to re work ingby 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 in2011 at Studorski preval in the Julian Alps. Ap prox i mately10 m of the Lower Tri as sic strata was sam pled just be lowthe con tact with the Anisian do lo mite. Nine sam ples (SP1–9) 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 isthe high est. Rock sam ples with an av er age weight of 4 kgwere pro cessed for cono dont study, us ing stan dard lab o ra -tory tech niques. Sev eral thin sec tions were made for thestudy 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 calSur vey of Slovenia (Geološki zavod Slovenije) where all ofthe micropalaeontological ma te rial is stored and in ven to ried un der re pos i tory num bers 4429, 4548–4555, 4901–4904and ab bre vi ated GeoZS. The col lec tion of macrofossil spec -i mens: Jurkovšek Paleontological Col lec tion, Kamnica 27,Dol pri Ljubljani, Slovenia, has been reg is tered with theNat 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.

OLENEKIAN (EARLY TRI AS SIC) FOS SIL AS SEM BLAGE, SLOVENIA 215

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

Fig. 3. Palaeogeographic map for Early Tri as sic with po si tionof 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 nantlymuddy 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 andgas 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 theWerfen 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 crofa cies type B can be in ter preted as slow de po si tion in adeeper ma rine en vi ron ment be low the fair-weather wavebase, which cor re sponds to the dis tal part of a ramp. Thepres ence of ammonoid shells in di cates a good con nec tionwith 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 interlayeringwith 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 talde tri tus dur ing high-en ergy events (microfacies type A).The sharp and ero sive lower bound aries of microfacies typeA 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/pack-stones and the up ward-in 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 areover lain by lam i nated or bioturbated cal car e ous mudstonesand/or marls (microfacies type B), their de po si tion re flectsshort 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 tratedshells 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 storm-sus -pended par ti cles or by ex hum ing pre vi ously bur ied shells byrip 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 trixof microfacies type A are in ter preted as the re sult of stormevents, 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 offoraminifera in di cates de tri tus de rived by storms from theshal 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 tionun der short-term high-en 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 theces sa tion of the storm and a re turn to nor mal low-en ergyramp sed i men ta tion be low the fair-weather wave base. Thepres ence of py rite also in di cates low-en ergy, and pos si blypartly an aer o bic con di tions, whereas the gen e sis of ce lest iteis con sid ered as diagenetic in deep-sea sed i ments, as explai-ned by Baker and Bloomer (1988) al though it also has beende scribed from shal low ma rine set tings (Hautmann, 1997).Storms are fre quently re corded both in the in ner and outershelf/ramp in the Induan/Olenekian sed i ments of the widerDinaric 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-rampzone (the zone be tween the fair- and storm- weather wavebases), as de fined by Burchette and Wright (1992).

There are only a few out crops of Lower Tri as sic stratain 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 ityin their de vel op ment and macrofossil con tent (Natiria costataand 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 tailedgeo 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 dontsam ples (SP 1–9) was ex am ined in the pres ent study. Theas 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 offoraminifera 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 forforaminifera 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.

ForaminiferaThe foraminifer clas si fi ca tions of Loeblich and Tappan

(1988) and Mikhalevich (2000) are com bined and usedhere. The foraminifer de scrip tions are short, be cause theseforaminifers are mainly well-known and the pres er va tion ofthese forms is not par tic u larly good.

Ge nus Ammodiscus Reuss, 1862Type spe cies: Ammodiscus infimus Bornemann, 1874

Ammodiscus? parapriscus Ho, 1959Fig. 5A–D

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


Ta ble 1Dis tri bu tion of microfossils in sam ples

from Studorski preval

SampleTaxa

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. praedensaUroševiæ

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 (5–10spec 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 61free spec i mens from the sam ple SP 1.De scrip tion: The test is cir cu lar, dis coid; pe riph ery rounded andcon cave on both sides. The spher i cal proloculus is fol lowed by aclosely 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 ofthe 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 well-pre 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 thege nus. The pres er va tion of the wall of the spec i mens stud ied is notgood, be cause they are recrystallized and in some cases pyritized.Strati graphic and geo graphic oc cur rence: The spe cies Ammo-dis 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 Lowerand 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, 1945Type spe cies: Glomospira umbilicata Cushman

& Wa ters, 1927

Glomospirella facilis Ho, 1959Fig. 5E, F

*1959 Glomospirella facilis n. sp. – Ho, p. 414, pl. 6, figs8–12.

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 freespec i mens from the sam ples SP 1 and 5.De scrip tion: The test is small, the equa to rial sec tion rounded, theax ial sec tion plain or con cave on the both sides. The spher i calproloculus 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 iswider than oth ers and a lit tle an gu lar in cross-sec tion. The wall isag glu ti nated, thin, and smooth. The ap er ture is at the end of thetube.Re marks: The char ac ter is tic fea tures of this spe cies are a roundform 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 abovemen tioned fea tures of Glomospirella facilis dif fer from those ofGlomospirella 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 insome 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 dleTriassic of China (Ho, 1959). Later this spe cies was re ported fromthe 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, 1959Fig. 5G, H

*1959 Glomospirella shengi n. sp. – Ho: p. 413, pl. 5, figs20–25.

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 freespec 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 calproloculus 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 theend of the tube.Re marks: The char ac ter is tic fea tures of this spe cies are the ovalform in the equa to rial sec tion and the con vex or plain form in theax ial sec tion, the streptospiral stage of coil ing is wider in the ax ialsec tion than the last whorl. The above men tioned fea tures ofGlomospirella shengi are dif fer ent from those of Glomospirellafacilis. 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). Laterthe 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, 1965Type spe cies: Pilammina densa Pantiæ, 1965

Pilammina? cf. praedensa Uroševiæ, 1988Fig. 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 around proloculus and an other inseptate, tu bu lar chamberlet. Theini tial wind ing cov ers two and a half to three close coils. The lastwhorl 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. There forethere 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 andthere is no con fi dence that this spe cies be longs to Pilammina andthis is true for the stud ied spec i men too.Strati graphic and geo graphic oc cur rence: Spe cies Pilamminapraedensa was first de scribed from the Lower Tri as sic of theDinarides (Uroševiæ, 1988) and af ter it was found in the up per partof Olenekian of Greece and the Lower Tri as sic of China (Rettori,1995).

Ge nus Hoyenella Rettori, 1994Type spe cies: Glomospira sinensis Ho, 1959

Hoyenella sinensis (Ho, 1959)Fig. 5M–O

*1959 Glomospira sinensis n. sp. – Ho, p. 410, pl. 3, figs16–34.

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, figs1–10.

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 1-6, 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 inshape, 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 thespe cies stud ied, be cause of the very sim i lar struc ture of the testand in this point, the de ci sion of Rettori (1994) is fol lowed. Glo-mospirella sp. (Zaninetti and Brönnimann, 1975) is placed intosyn 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 thespec 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 ofChina (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; Efi-mova, 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, 1826Type 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 ovalcham bers is from 4 to 8. They usu ally in crease very grad u ally insize and are, com par a tively loosely jointed in all cham bers. Thesu 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 ovalprolocolus and other cham bers are oval and elon gated. The holo-

type 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 thereis prolocolus pres ent in all spec i mens.Strati graphic and geo graphic oc cur rence: Spe cies Dentalinasplendida was first de scribed from the lower part of the Olenekianof the North Si be ria (Gerke, 1961). Later this spe cies was found inthe 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 lowerpart 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., Hoyenellasinensis (Ho), Dentalina splendida Schleifer, Nodosaria? spp. (Fig.5A–R). Spe cies of the gen era Ammodiscus, Glomospirella and Ho-yenella 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 somemiliolids and nodosariids. The foraminifers are very small and notwell-pre 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, fromthe Alps to China. Sev eral au thors have in di cated that there are“Glomospira-Glomospirella” 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 Studorskipreval lo cal ity has some spe cies com mon with the foraminifer as -sem blages from the Lower Tri as sic of China (He, 1993), theOlenekian of the Cau ca sus area (Efimova, 1991), the Lower Tri as -sic and Anisian of Bul garia (Trifonova, 1992) and Hun gary (Ora-vecz-Scheffer, 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 somemiliolids and nodosariids are known mainly from the Lower Tri as -sic (Dimitrijeviæ et al., 1968; Pantiæ, 1970; Pantiæ-Prodanoviæ andRadoševiæ, 1977b; Pisa et al., 1979; Resch, 1979; Broglio Loriga etal., 1990). Ear lier in west ern Slovenia, the “Glomospira-Glomospirella”foraminifer as sem blages with some nodosariids were re portedfrom 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 theforaminifer as sem blages men tioned above, be cause they have acom 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. InTri as sic time, the rep re sen ta tives of Meandrospira are typ i cal forthe Early Tri as sic and the Anisian.It is rea son able to con clude that the foraminifer as sem blage fromthe 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 blyre stricted to the up per part of the Early Tri as sic.

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

Or chard (2005).

Ge nus Triassospathodus Kozur, 1998 in Kozur, Mostlerand Krainer (1998)

Type spe cies: Spathognathodus homeri Bender, 1970


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 P-el 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 inlength 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 mentshave an ex panded basal cav ity. The ramiform el e ments are mostly frag mented and they do notper mit re con struc tion. Some of them are de ter mined as ?M, ?S1and S3 el e ments.Re marks: Triassospathodus can be eas ily dis tin guished fromNeospathodus Mosher, 1968, which has a ter mi nally up ward-di -rected lower mar gin of the basal cav ity (Mosher, 1968). On theother hand, there ex ists a morphologic sim i lar ity in the P el e mentsof Triassospathodus Kozur, 1998 (in Kozur et al., 1998) andNovispathodus Or chard, 2005 (Or chard, 2005); the lat ter has ashorter and more wid ened basal cav ity and it has also a dif fer entap 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. hunga-ricus was first de scribed from the Tirolites beds of Felsõrs in Hun -gary (Kozur and Mostler, 1970). Triassospathodus is an im por tantLower 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 thebasal Spathian equiv a lent to the Tirolites cassianus ammonoidZone. Ac cord ing to H. Kozur (pers. comm., 2012), in the shal lowwest ern Tethys the T. hungaricus fauna lies within the lowerSpathian, 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 andwith 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 Dina-rides, 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-knownfor 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 Spathianage.

Bi valves

Ge nus Bakevellia King, 1848Type spe cies: Avicula antiqua Graf zu Münster in

Goldfuss, 1836 [non A. antiqua Defrance] = Aviculabinneyi 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, figs3a–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 withcon 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. Beakspointed, 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 thereare slight dif fer ences in the out line by com par i son with the sin gleright 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 notpro ject above the hinge line, whereas in the ma te rial de scribed, itdoes. 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 sionand 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 salmar 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 theintraspecific 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 ofspec 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 tralvalve mar gins, i.e. the hinge mar gin lies in a dor sal po si tion, moreor less par al lel to the sub strate. It is un likely that this was the lifepo 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 entedmore 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 thetop of a tempestitic bed, as in di cated by hummocky cross-strat i fi -ca tion, a pos si ble ex pla na tion is that the an i mals were ex ca vatedand 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 Studorskipreval material might thus be an unnatural dying position of thisspecies.

Ge nus Eumorphotis Bittner, 1901Type 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 shelllayer (BJ 2766), ques tion able three abraded left valves (BJ 2701).De scrip tion: Very large Eumorphotis with about 70 ra dial ribsnear 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, butpos te rior wing rel a tively elon gated.Re marks: The larg est spec i men is about 8 cm long and more than8 cm high. This is to the knowl edge of the au thors the larg estEumorphotis 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 isob served on the pos te rior wing. There fore, it is as sumed that thehigher 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 chieflyin 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 ofthe disc. Al though the pres ent au thors cur rently treat both spe ciessep a rately, they wish to note that the dif fer ence in or na men ta tionmight turn out to be a pres er va tion ef fect or as ly ing within therange of intraspecific vari a tion. In this case, E. kittlii would havepri or ity over E. gronensis and spe cies as sign ment of the spec i menwould change ac cord ingly (see also Broglio Loriga and Mirabella, 1986).

Ge nus Aviclamys Allasinaz, 1972Type 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 (BJ2765).

De scrip tion: Disc nearly cir cu lar, cov ered with about ten strongfirst-or der ra dial ribs plus the same num ber of in ter ca lated weakersec 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 toAvichlamys 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 uredby 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, 1881Type 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; WerfenFor 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 arerecrystallized 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 theWerfen 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 scribedfrom 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 ingmode of pres er va tion

Tri as sic) St. Cassian For ma tion. It seems, how ever, that the typese 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 thema te rial of Münster (1841) could have come from an out crop ofthe Werfen Fm at Groneshof, where it oc curs abun dantly. Tak inginto ac count ap prox i mately 100 mil lion years and the P-T 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 uityof 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 servedand does not con trib ute any new mor pho log i cal in for ma tion to theex 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 sizeis 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 twospe cies oc cur in Eu rope, co in cid ing roughly with out crops of theup 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 EarlyTri as sic fos sil site in the Julian Alps of Slovenia. The re porton the palaeontological ma te rial, col lected at Studorskipreval, 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 ofthe ear li est Spathian cono dont fauna with Triassospatho-dus hungaricus in di cates an Olenekian age of the fauna andis 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 thatthe strata of Studorski preval in the Julian Alps of west ernSlovenia were de pos ited in a more dis tal, storm-in flu encedramp. A ramp set ting was also men tioned for the de po si tionof the Werfen For ma tion in the South ern Alps (Brandner etal., 2012). Two types of beds were rec og nized, des ig natedhere 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 bythe lam i nated or bioturbated mudstones and/or marls ofmicrofacies 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 peaksand the de po si tion of coarse-grained skel e tal ma te rial as thestorms 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-freecon nec tion of the depositional en vi ron ment with the opensea, 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 ergydeeper part of the ramp. A low-en 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 tal-rich 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. praedensaUroseviæ, Arenovidalina? sp., Meandrospira? sp., Agatha-mmina? sp., Hoyenella sinensis (Ho), Dentalina splendidaSchleifer and Nodosaria? spp. The main fea ture of the tax o -nomic com po si tion of this foraminifer as sem blage is thedom i nance of prim i tive ag glu ti nated foraminifers (Ammo-dis cus, Glomospirella, Hoyenella), as so ci ated with somemiliolids and nodosariids. It can be as signed to the timerange from Early Tri as sic to Anisian “Glomospira-Glomo-spirella” foraminifer com mu nity. An Early Tri as sic age ismore 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 podsform monospecific coquinas that are most likely of theshells of Natiria costata, a typ i cal faunistic el e ment of theTethyan Early Tri as sic. The bi valve fauna in cludes three genera that dif fer in strati graphi cal range and geo graph icaldis tri bu tion. Bakevellia is a long-lived ge nus, known world -wide from the Perm ian up to the Cre ta ceous . Eumorphotiswas 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 sistedlo cally into the Mid dle Tri as sic. In con trast, Avichlamyswas re stricted to the Olenekian and is cur rently known onlyfrom the west ern Tethys. The bi valve spe cies in clude Bake-vellia cf. incurvata, Eumorphotis cf. gronensis and Avicla-mys 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 cordfor the larg est size of this ge nus, so far re ported from theEarly Tri as sic. The un usual po si tion of ar tic u lated Bake-vellia spec i mens is in ac cor dance with storm-event 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 fromthe 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 prevalsec 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 thislow 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 oververy large dis tances, which is un likely given the low levelof frag men ta tion. Miller (1988) dem on strated an evo lu tion -ary on shore-off 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 -shore-off shore or prox i mal-dis 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 flectan 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 tingsof 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 trendthat 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. Thestudy was partly sup ported by the Slovenian Re search Agency(Programme Num ber P1-0011 and Pro ject Num ber J1-9465) andCro 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 vonHumboldt Foun da tion Re turn Fel low ship and the In sti tute of Pa-leobiology 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 sicmass ex tinc tion”).

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Olenekian (Early Triassic) fossil assemblage from eastern Julian Alps (Slovenia)

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