Geol. Paläont. Mitt. Innsbruck, ISSN 0378-6870, Bd. 19, S. 165-199

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MIDDLE TRIASSIC CONODONTS FROM THE SOUTHERN KARAWANKENMOUNTAINS (SOUTHERN ALPS) AND THEIR STRATIGRAPHIC IMPORTANCE

Heinz KOZUR, Karl KRAINER and Helfried MOSTLER

With 3 Figures and 4 Plates

Abstract:The conodont fauna of south alpine Middle Triassic pelagic limestones (Loibl Formation and Buchenstein Formation) ofthe Karawanken Mountains in Carinthia, southern Austria, is described and the systematics and stratigraphie importanceof Late IUyrian and Fassanian gondolellid conodonts are discussed.The investigated conodont fauna contains the following new taxa: Neogondolella cornuta ladinica n. subsp., Neogondo-lella aldae n. sp., Neogondolella aldae aldae n. subsp., Neogondolella aldae posterolonga n. subsp., Neogondolella ?postpridaensis n. sp., Paragondolella ? pridaensis posteroacuta n. subsp. and Budurovignathus gabriellae n. sp.The stratigraphie evaluation of the conodonts supports the priority of the position of the Anisian-Ladinian boundary at thebase of the Reitziites reitzi - Zone, where a distinct change of all stratigraphically important microfossil groups is ob-served.The oldest investigated sediments are red fissure fillings within the uppermost part of the Late Anisian platformcarbonates of the Contrin Formation, containing conodonts probably indicating latest Illyrian age. The conodont fauna ofthe Loibl Formation points to Fassanian age. Sediments of the Buchenstein Formation range in age from the Fassanian tothe Late Langobardian {Budurovignathus mungoensis - Zone).

Zusammenfassung:In der vorliegenden Arbeit wird die Conodontenfauna mitteltriadischer pelagischer Kalke (Loibl Formation und Buchen-stein Formation) aus dem Südalpin der Karawanken in Kärnten (Südösterreich) beschrieben sowie die Systematik undstratigraphische Bedeutung illyrischer und fassanischer gondolellider Conodonten diskutiert.Die untersuchte Conodontenfauna enthält folgende neue Taxa: Neogondolella cornuta ladinica n. subsp., Neogondolellaaldae n. sp., Neogondolella aldae aldae n. subsp., Neogondolella aldae posterolonga n. subsp., Neogondolella 1 postpri-daensis n. sp., Paragondolella ? pridaensis posteroacuta n. subsp. und Budurovignathus gabriellae n. sp.Die stratigraphische Auswertung der Conodonten bekräftigt die Position der Anis-Ladin-Grenze an der Basis der Reit-ziites reitzi - Zone, die auch der Priorität entspricht. An dieser Grenze kann eine deutliche Änderung aller stratigraphischwichtigen Mikrofossilien beobachtet werden.Die ältesten untersuchten Sedimente sind Rotkalke in Form von Spaltenfül-lungen im obersten Teil der oberanisischen Plattformkarbonate der Contrin Formation, deren Conodonten auf oberstesIllyr hinweisen.Die Conodontenfauna der Loibl Formation ist in das Fassan einzustufen. Sedimente der BuchensteinFormation reichen altersmäßig vom Fassan bis in das obere Langobard {Budurovignathus mungoensis - Zone).

1. Introduction

Although the investigation of the Triassic se-quence in the Karawanken Mountains alreadystarted during the last century (MOJSISOVICS1871, TELLER 1887, 1898 for example), untilrecently much confusion existed concerningMiddle Triassic stratigraphy in the southalpinepart of the Karawanken Mountains (see BAUER

1980, 1984, BAUER et al. 1983). Especially theposition of the Anisian-Ladinian boundary andthe age of the Middle Triassic basin sequencewas unclear due to the lack of detailed paleonto-logical and biostratigraphical investigations.

BAUER (1980, 1984) reported ammonitesfrom red pelagic limestones of the Loibl Forma-tion: Kellnerites sp. from the peak of the ZellerPrapotnik mountain and an Early Ladinian am-

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Fig. 1: Location map of the study area. 1 - 7: investigated outcrops (1 Weiße Wand, 2 Zelenitza forest road, 3 Kraßniggraben sec-tion, 4 Zeller Prapotnik, 5 Obojniggraben section, 6 Zimpaserkogel west, 7 Zimpaserkogel northwest).

monite fauna from outcrops along the Zelenitzaforest road (see also geol. map of BAUER 1981,1985).

KRAINER & MOSTLER (1992) describedsiliceous sponge spicules from red nodular lime-stones (Weiße Wand Member) of the Loibl For-mation and dated these limestones as Early Fas-sanain based on the radiolarian- and conodontfauna (Spongosilicarmiger italicus -Zone; Para-gondolella trammeri praetrammeri -Zone). Theradiolarian fauna will be described in a separatepaper (KRAINER & MOSTLER in prep.).

Rich radiolarian faunas from the BuchensteinFormation of the central Karawanken Moun-tains indicate an age ranging from Late Fassa-nian to Late Langobardian (MOSTLER &KRAINER 1994).

The investigation of the stratigraphically im-portant conodont fauna is of decisive impor-tance for the final definition of the Anisian-La-dinian boundary. In contrast to further opinions,the Middle Triassic gondollellid conodonts

show their strongest changes at the base of theReitziites reitzi - Zone which corresponds to thebase of the Ladinian according to the more than100 year old priority. At or near this boundary amajor change in the stratigraphically importantmicrofossils can be observed.

In the present paper the conodont fauna fromthe Middle Triassic basin sequence of the centralKarawanken Mountains is described. For thestratigraphie evaluation of the investigated co-nodont fauna it is also necessary to discuss thesystematics of the Illyrian and Fassanian gondo-lellid conodonts and the position of the Anisian-Ladinian boundary.

2. Location

The conodont faunas described and discussedin this paper are derived from red and grey pe-lagic limestones of the Loibl Formation andfrom nodular cherty limestones in the middle

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and upper part of the Buchenstein Formation ofthe south alpine Triassic seuqence of the Karaw-anken Mountains in southern Carinthia (Aus-tria), near the Austrian/Slovenian border.

Investigated outcrops of the Loibl Formationare situated in the central Karawanken Moun-tains, between Zeil Pfarre/Koschuta in the eastand Märchenwiese/Weiße Wand in the west:Zeller Prapotnik, Zelenitza forest road west ofLoibl and Weiße Wand (the locations are shownon fig. 1).

Two sequences of nodular limestones of theBuchenstein Formation, exposed along forestroads on the northern and western side of theZimpaserkogel southwest of Eisenkappel alsoprovided a relatively rich and interesting conod-ont fauna (location see fig. 1).

3. Geological setting and stratigraphy

The southern Karawanken Mountains arepart of the Southern Alps and separated fromthe Eastern Alps (Northern Karawanken Moun-tains) by the Periadriatic Line, which runsthrough the Karawanken Mountains in an E-W-direction.

The southern Karawanken Mountains arecomposed of a Variscan basement (Late Silurianto Middle Carboniferous) and a postvariscancover sequence which starts with sediments ofthe Late Carboniferous Auernig Group. ThePermian is represented by sediments of the Rat-tendorf Group, Trogkofel Group, Tarvis Brec-cia, Gröden and Bellerophon Formation.

The Triassic sequence is more than 2000mthick and predominantly composed of shallowwater and pelagic carbonates. The Middle Trias-sic sequence is shown on Fig 2.

The Triassic sequence begins with the southalpine Werfen Formation, which is developed ina very similar faciès as in the Dolomites, and isoverlain by several hundred m thick, well bed-ded and sometimes evaporitic platform carbo-nates corresponding to the Lower Sari Forma-tion of the Dolomites.

SCHLERNDOLOMITE

BUCHENSTEINFORMATION

PIETRA VERDE

MARLYLIMESTONES

MARLS

SANDSTONES

CONGLOMERATES

VOLCANICS

LOIBL FM.

CONTRIN

FORMATION

Fig. 2: Composite section of the south alpine Middle Triassic(Ladinian) basin sequence in the central Karawanken Moun-tains.

Above there follow dark grey, well bedded,bioturbated marly limestones ('Wurstelkalke')and intercalated marls, up to some tens of mthick, and about 30 m thick, thin bedded lami-nated dolomites.

At the Weiße Wand (location see fig. 1) thesedolomites are overlain at places by an approxi-mately 7 m thick sequence of red marls andintercalated limestonebeds (probably equiva-lents of the Lower Peres Formation of the Dolo-mites).

The red marls and laminated dolomites areoverlain by massive, light grey limestones (Con-

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dark marls,siltstones

-ZG3O

-ZG 2 green marls en

nodularlimestone

green marls

-ZG1 (Retra Verde)

- 9- 8

nodular chertylimestone

nodularlimestone

ammonites

r-r-n

chertylimestone

- 2

+-_t±f / +\ !±_+|- Z1 tufflayer

Fig. 3: Measured sections through parts of the Buchenstein Formation exposed along the forest road on the northwestern side (sec-tion A) and western side (section B) of the Zimpaserkogel SW Eisenkappel, and location map (see also fig. 1).

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trin Formation) with a maximum thickness ofabout 50 m (Late Anisian "algal reefs"), form-ing prominent cliffs like Motschiwa, Kosmatica,Weiße Wand and Heilige Wand.

At the base of the Ladinian (Fassanian), ex-tensional tectonic movements caused break-down of the Anisian carbonate platforms andformation of basins by rapid subsidence of dis-tinct blocks, which were filled with different ty-pes of basin sediments and volcanic rocks.

These tectonic processes are indicated by fis-sures within the uppermost algal reef limestones(Contrin Formation) filled with red limestones,and red pelagic limestones (Loibl Formation)which at places overlie the massive shallow wa-ter limestones of the Contrin Formation.

Due to the rich microfauna (sponge spiculae,holothurian remains, radiolarians and conod-onts) the pelagic limestones of the Loibl Forma-tion are of Early Fassanian age (Spongosilicar-miger italiens - Zone; Paragondolella trammeripraetrammeri - Zone; KRAINER & MOSTLER1992, KRAINER & MOSTLER in prep.).

Two types of red pelagic limestones can bedistinguished:a) A very thin sequence of nodular cherty lime-stones (bioclastic wackestones rich in spongespiculae and radiolarians; Weiße Wand Mem-ber), interpreted as the basin faciès, andb) massive to indistinctly bedded red limestonescontaining abundant echinoderm fragments,representing the slope faciès. Thin layers ofpyroclastic material (tuffs) are intercalated.

Grey limestones are composed of wacke-stones containing radiolarians, spiculae andshell fragments. Echinoderm remains, foramini-fers, gastropods and ostracods are also present.

Red limestones are represented by wacke-stones-packstones rich in echinoderm fragmentsor radiolarians, and less frequently by grain-stones. The grainstones are composed mainly ofechinoderm remains. Carbonate lithoclasts de-rived from shallow water carbonates are alsopresent. Nodular red limestones (radiolarianwackestones) also contain some ammonites.

The fauna of the nodular cherty limestones(Weiße Wand Member) points to water depths

of approximately 200m. The sponge spiculaefauna is very similar to that of the VicentinianAlps (Recoaro, Southern Alps), but differs sig-nificantly from that of the Northern LimestoneAlps (Eastern Alps).

A detailed description of the Loibl Formationwill be presented in a separate paper.

The tectonic event at the Anisian/Ladinianboundary lasted for a very short time and was apreliminary stage to the intra-Ladinian tectonicevent which resulted in the formation of basinsfilled with sediments of the Buchenstein Forma-tion and associated deposits.

Contrin and Loibl Formation are overlain byvolcanic rocks (agglomerates, tuffs, lavabrecciasand lavas) of andesitic to dacitic compositionand a maximum thickness of about 100m (seeOBENHOLZNER 1985).

The basin sequence above these volcanicrocks starts with clastic sediments (polymictconglomerates, sandstones, siltstones and mud-stones) with thicknesses ranging between a fewmeters and 45 m. The conglomerates containmany volcanic clasts derived from the underly-ing volcanic rocks and a few red imestone clastsderived from the Loibl Formation. The sand-stones are also rich in reworked volcanic materi-al.

These clastic sediments are equivalents of theUggowitz Breccia and are interpreted to be for-med by submarine mass flows (debris flows,turbidites).

The clastic sequence upward grades intomudstones, which in the lower part containintercalations of fine-grained conglomerate,sandstone and siltstone layers. Carbonate con-tent increases upward and up to several m thicknodular marly limestones are frequently inter-bedded. In the Kraßniggraben east of the Loibl-paß this sequence reaches a thickness of 150 m.

With a relatively sharp boundary sediments ofthe Buchenstein Formation overlie this muddysequence.

In the Kraßniggraben the Buchenstein Forma-tion is completely exposed and about 40 m

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thick. The sequence starts with 2.5 m thick, bed-ded limestones with thin tuffaceous intercala-tions, overlain by 6 m thick, coarse grained crys-tal tuffs (Pietra Verde), and thin bedded, nodulargrey limestones (bioclastic wackestones rich inradiolarians and filaments), partly with chertnodules, thin chert layers and thin marl and gre-en tuffaceous intercalations.

In the upper part red and green nodular lime-stones with up to 10 cm thick marl intercalationsare developed.

In the Kraßniggraben section the BuchensteinFormation is overlain by thin bedded, dark greyplaty limestones with dark marl intercalations.

West of the Loiblpaß, on the northeasternflank of the Bielschitza (location see fig. 1), theuppermost Buchenstein Formation is composedof cm to dm bedded calcarenites and fine grai-ned breccias (turbiditic shallow water detritusderived from the prograding Schiern platform).This faciès is overlain by massive carbonates(reef-facies) of the Schiern Dolomite.

In the Obojniggraben (location see fig. 1) theBuchenstein Formation (about 20 m thick) iscomposed of well bedded grey nodular lime-stones with up to several cm thick dark marlyintercalations and a few thin Pietra Verde lay-ers. Cherty limestones are rare. On top of theBuchenstein Formation thin bedded dark greyplaty limestones are developed.

Of interest are outcrops of the BuchensteinFormation along a forest road on the northwest-ern and western side of the Zimpaserkogel onthe eastern side of the Obojniggraben (fig. 3).

On the western side (ca. 1050m) a 7 m thicksequence of grey and light red colored nodularlimestones, indistinctly thick bedded, in the up-per part thinn bedded with thin shale partings isexposed above a coarse grained crystal tuff (Pie-tra Verde). Samples from this section containeda rich conodont fauna.

On the northwestern side of the Zimpaserko-gel (ca. 1160 m) along the forest road the upper-most part of the Buchenstein Formation is ex-posed: grey, well bedded nodular limestoneswithout chert, bed thickness ranges between 5and 30 cm. Thin green tuffaceous intercalations

and marls occur. The sequence is overlain bydark grey, partly laminated marls and siltstoneswith small scale current ripples (? equivalents ofthe Wengen Formation). From the nodular lime-stones of this section conodonts were obtained.

Conodont faunas desribed in this paper arederived from (locations see fig.l and 3):a) Red and grey pelagic limestones of the LoiblFormation from following localities:- Weiße Wand (Weiße Wand Member)- Zelenitza forest road- Zeller Prapotnikb) from the middle and upper part of the Bu-chenstein Formation, exposed along forest roadson the western and northwestern side of theZimpaserkogel.

Samples from other localities (Obojniggrabensection, Kraßniggraben section, Bielschitza sec-tion, Rotschitzagraben and Rauscherbach-Gra-ben) did not contain determinable conodonts, al-thogh more than 100 samples have been investi-gated.

4. Systematic part

Most of the stratigraphically important Illyr-ian and Fassanian conodonts belong to the genusNeogondolella BENDER & STOPPEL, 1965emend. KOZUR, 1990 and ParagondolellaMOSHER, 1968 emend. KOZUR, 1990. Thesesmooth platform conodonts need a revision. Afirst revision of the N. constricta-N. momber-gensis group by NICORA & KOVÄCS (1984)has failed. These authors have included intoNeogondolella constricta several well definedand partly even not related species and subspe-cies, such as Neogondolella cornuta BUDU-ROV & STEFANOV, 1973, N balkanica BU-DUROV & STEFANOV, 1975, Neogondolellamombergensis prava KOZUR. Moreover, theyplaced the early juvenile forms of all Illyrianand Ladinian Neogondolella species into N.constricta.

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KOVÀCS used for this publication a samplefrom the German Basin given by KOZUR to-gether with conodont descriptions for the neverpublished conodont catalagous of Middle andUpper Triassic platform conodonts. Because hehas not contacted KOZUR about the fossil con-tent of this sample before using it for publica-tion, he has not known that this sample yieldedonly conodonts of the N. mombergensis group(the sample has been derived from the type lo-cality of N. mombergensis). Whereas someforms from this sample have been correctly pla-ced into N. mombergensis, a part of juvenileforms have been placed into N. constricta andmisinterpreted as medium ontogenetic stage(NICORA & KOVÀCS, 1984, PL 8, fig. 4).

Identical late juvenile Neogondolella fromanother sample have been placed partly into N.constricta, partly into N. mombergensis (e.g.,NICORA & KOVÀCS, 1984, pi. 7, figs. 12,13). On the other hand, none of the M momber-gensis from Nevada, figured by NICORA &KOVÀCS, 1984) belong to this species.

Because NICORA & KOVÀCS (1984) pla-ced even juvenile forms of typical N. momber-gensis and of stratigraphically still younger Neo-gondolella species into N. constricta, the strati-graphic value of the Illyrian and Ladinian Neo-gondolella, for the first time shown by KOZUR(1968) and BUDUROV & STEFANOV (1973a) have been totally mask. As an logic conse-quence, RITTER (1989) has then all M constric-ta and N. mombergensis, figured by NICORA &KOVÀCS (1984), placed in a single species (ac-cording the priority N. mombergensis).

KOVACS et al. (1990) subdivided their Nconstricta s.l. into several morphotypes of dif-ferent stratigraphie ranges. However, in the co-nodont form taxonomy all species with differentranges are morphotypes. Therefore the termmorphotype should be only used for intraspecif-ic form groups of the same stratigraphie range oras informative term before establishing a formtaxon. Most of the morphotype gamma sensuKOVÀCS et al. (1990) belong to N. mesotrias-sica (KOZUR & MOSTLER, 1982).

The Illyrian and Ladinian Neogondolella spe-cies will be revised by KOZUR & MOSTLER(in prep.). The preliminary results are used inthe present paper to make some remarks on spe-cies that are present in our material or near relat-ed to them. Moreover, some new taxa are de-scribed.

Unfortunately, by technical mistakes much ofthe conodonts have been crashed during stick-ing. However, using the data of the revision ofthe Middle Triassic gondolellids (KOZUR &MOSTLER, in prep.), also broken specimensconsisting only of the posterior half or eventhird of an adult form can be determined.

Neogondolella mombergensis(TATGE, 1956)

Remarks: Typical adult representatives of thisspecies display high anterior and posterior denti-cles without prominent cusp, or the penultimatetooth is the cusp. The middle part of the carina islow and mostly fused to a smooth to wavy ridge.The basal cavity is somewhat foreward-shiftedwith respect to the keel end. Early juvenile sta-ges are of constricta type (the penultimate denti-cle is the cusp, the last denticle is smaller thanthe cusp) and have been therefore erroneouslyplaced into this species by NICORA &KOVÀCS (1984). Late juvenile stages are ofmodified mombergensis type (or media type, alldenticles have about the same size). Mediumontogenetic and subadult stages have already amombergensis type of carina, but the denticlesof the lower middle carina are still separated, atleast at their tips.Occurrence: Upper P. trinodosus - Zone andFassanian. Dominant in the German Basin. Pre-sent also in the Northern Tethys (e.g. Beckovsection of Slovakia). Rare in the Southern Teth-ys. So far not known from North America. AllNevadian uppermost Anisian and Ladinian adultforms figured as N. mombergensis mombergen-sis by NICORA & KOVÀCS (1984) representother species (mostly N. aldae n. sp.).

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Neogondolella constrictaMOSHER & CLARK, 1965(PL 4, figs. 12-15, 17, 20)

Remarks: The holotype of N. constricta is ajuvenile form. All Illyrian and Fassanian Neo-gondolella species display an early juvenile on-togenetic constricta stage. Unfortunately, so farno adult specimens have been figured from thestratum typicum of N constricta (sample FH-3by MOSHER & CLARK, 1965 from the Para-ceratites clarkei beds of the Rotelliforme Zone).Even NICORA & KOVÂCS (1984), who re-vised the N. constricta group, did not figure anyadult specimen from the stratum typicum andeven not from the P. clarkei beds, but only fromyounger beds. They reported, however, the pres-ence of adult forms in the P. clarkei beds thatcorrespond to their opinion to the cornutamorphotype and to the balkanica morphotype.The presence of N. balkanica s. str., a Ladinianguide form, in the lower part of the Illyrian Par-aceratites beds is not likely. However, it is wellpossible that two Neogondolella species, separa-ble only in adult stages occur together with thejuvenile N constricta in the Paraceratites clar-kei beds. In this case, N. constricta has to beabandoned, because all Illyrian and LadinianNeogondolella species have at least an early on-togenetic constricta stage. In some of them,even the late juvenile forms are of constricta ty-pe. If in the type material two different speciesoccur together with juvenile forms attributed tothe type constricta, then it is not clear, to whichof the two species belong the juvenile forms.

In our material from the lower part of Para-ceratites beds of the Eurasian Tethys all neogon-dolellid juvenile platform conodonts, but alsomedium ontogenetic stages are of N. constrictatype. They are accompanied by two differentspecies of adult Neogondolella that may beidentical with the adult forms mentioned but notfigured by NICORA & KO VACS and errone-ously attributed to the cornuta and balkanicamorphotypes. The dominating form (A) has acarina very similar to that of N mombergensiswith high denticles in the anterior and posterior

part and a fused low middle part. The cusp, ifpresent, is terminal, fused with the posteriorplatform margin and not prominent. The onlydifference to N mombergensis is the position ofthe basal cavity that is situated nearly at the ter-minal end of the keel even in adult forms. In Nmombergensis, the basal cavity is somewhatforeward-shifted in adults and lies subterminalwith respect the keel end. This form (A) couldcorrespond to the cornuta morphotype sensu NI-CORA & KOVÂCS (1984) of the P. clarkeibeds. The second, rare form (B) has a distinctcusp, surrounded by a brim. It is similar to theparatypes of Neogondolella basisymmetricaBUDUROV & STEFANOV, 1973, that are,however, not identical with the holotype (seeunder remarks to TV. aldae n. sp.). This form (B)could be identical with the balkanica morpho-type sensu NICORA & KOVÂCS (1984) thatthey reported, but not figured from the P. clarkeibeds.

Both discussed adult forms correspond also intheir medium ontogenetic stages to N. constric-ta. By this they are clearly distinguished from TV.cornuta and N. balkanica that have both a slen-der medium ontogenetic stage with a carina ofmombergensis type (denticles of the same heightor somewhat lower in the middle part, in thisstage no distinct main cusp is present). For thisreason neither N. cornuta nor N. balkanica arejunior synonyms of N. constricta as assumed byNICORA & KOVÂCS (1984).

At present, we regard the mombergensis-\ïkeadult forms as true adults of N. constricta, be-cause the adults with distinct cusp, surroundedby a brim, are rare (seemingly also in the typematerial of TV. constricta the so-called balkanicamorphotype is rarer than the so-called cornutamorphotype). In this restricted scope N. con-stricta does not reach above the Illyrian Para-ceratites beds both of North America and theEurasiatic Tethys. Perhaps it is even restricted tothe lower part of the Paraceratites beds. How-ever, an occurrence in the lower Illyrian Para-gondolella bifurcata - Zone or even in the Pelso-nian cannot be excluded, because also there the

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Neogondolella species have a constricta juve-nile stage.

Neogondolella bacaloviBUDUROV & STEFANOV, 1973

Neogondolella transita(KOZUR & MOSTLER, 1971)

Remarks: The central morphotype of this spe-cies displays a nearly symmetrical, but mostlyslightly sigmoidally bent platform. The holotypebelongs to this morphotype (posterior end onlyslightly curved). In other forms the posterior endis more or less strongly curved toward the left orthe right. Between all these forms gradual transi-tions exist throughout the entire stratigraphierange. BUDUROV & STEFANOV (1973 a) es-tablished for the same species the taxon N. ex-centrica. As holotype an eccentric form (posteri-or end curved toward right) has been chosen, butalso the central morphotype of N. transita wasincluded into N excentrica that is a junior syno-nym of N. transita.Occurrence: Typical N. transita with obliqueand pointed posterior end of the keel occurs inthe Nevadites - Zone and in the lower E. curionii- Zone. Often different forms with asymmetricposterior end (a feature that is common in manyNeogondolella species) have been placed into N.'excentrica'. However, in these forms the keelend is only oblique, but blunt or rounded, neverpointed. Such forms with distinctly foreward-shifted basal cavity with respect to the keel endare common in the Reitziites reitzi - Zone. Theyare here listed as N sp. aff. transita. They areseemingly the forerunner of N. transita. In theIllyrian and even in the Pelsonian forms withsimilar eccentric posterior end occur that displaya nearly terminal basal cavity with respect to thekeel end. They are eccentric morphotypes of dif-ferent Illyrian Neogondolella that have beenpartly placed into N. 'excentrica ' as well. Noneof these forms is directly related to N. transita(including N. excentrica).

Remarks: This relatively small, slender form iseasily recognizable by the distinctly protrudingmargin of the basal cavity and the strongly fore-ward-shifted basal cavity both in respect to theplatform and keel end. Already in juvenileforms this foreward-shifting of the basal cavityis recognizable, but it is not so strong in juvenileforms and not present in earliest juvenile stages.Characteristic is also the upward bent of the pos-terior platform after the basal cavity. The keel isalso distinct in this upward bent part. BUDU-ROV & STEFANOV (1973 a) figured onlysymmetrical or slightly asymmetrical forms (tothe latter ones belong the holotype). However,there are also strongly asymmetric forms, oftenstrongly constricted on one or two side of theupward bent part of the platform. Such formsthat are connected with N. bacalovi by all transi-tions and occur also in the same stratigraphielevel, have been described as N. huckriedei BU-DUROV & STEFANOV, 1975. The typical bac-alovi features of the lower side of N. huckriedeiare especially well figured by BUDUROV(1976, PL 1). This species is here regarded asjunior synonym of N. bacalovi BUDUROV &STEFANOV, 1973.

N transita KOZUR & MOSTLER, 1971 issimilar, but the platform is broader and the unitis mostly larger. Especially the part after the ba-sal cavity is broader and shorter and not so dis-tinctly upward bent. By this, there is no strongprotrusion of the basal cavity margin.

Neogondolella cornutaBUDUROV & STEFANOV, 1973

(PL 3, Figs. 2,5, 8, 11, 14, 17;PL 4, Figs. 16, 18,19,21)

Remarks: As already shown by BUDUROV &STEFANOV (1973 a), N. cornuta has a mediumontogenetic stage with mombergensis type ofthe carina. By this it is clearly distinguished

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from N. constricta in the here used narrow sco-pe. Only the early juvenile stages are of con-stricta type.

Typical Neogondolella balkanica BUDU-ROV & STEFANOV, 1975 are distinguished byan erect main cusp, surrounded by a distinctplatform brim. These forms are restricted to theLadinian. BUDUROV & STEFANOV (1975)and later papers placed all rather slender formswith erect main cusp into N. balkanica, but asholotype a specimen was chosen that displays adistinct platform brim behind the erect cusp.Forms with erect cusp fused with the posteriorplatform end are gradually and transitionallyconnected with such forms of N. cornuta thatdisplay a posteriorly inclined cusp, likewise fu-sed with the platform end. However, such formsbecome more frequent in the upper range of cor-nuta. May be that they represent an independentsubspecies, transitional to N. balkanica. Onlysuch forms with terminal erect cusp we place in-to N. balkanica, in which the cusp is not moredistinctly fused with the posterioir platformmargin. All these forms have a distinctly for-ward-shifted basal cavity and are therefore onlysimilar to N. cornuta ladinica n. subsp.

Advanced N. cornuta (N. cornuta ladinica,description below) from the Fassanian have adistinctly foreward-shifted basal cavity with re-spect to the keel. The posterior platform part be-hind the basal cavity is somewhat upward bentand the keel portion behind the basal cavity isdistinct on this part as well.Occurrence: N. cornuta cornuta is very com-mon in the (upper) Paraceratites faunas of theIllyrian. Fassanian forms have been derived of-ten from condensed beds (like from glauconite-bearing limestones of the Karwendel). In uncon-densed beds N. cornuta cornuta is successivelyreplaced by N. cornuta ladinica during the earlyFassanian. All late Fassanian N. cornuta belongto N. cornuta ladinica.

Neogondolella longaBUDUROV & STEFANOV, 1973

(PL 2, Figs. 18, 21, 22; PL 4, Figs. 3-5)

Remarks: The holotype of N. longa is a me-dium ontogenetic stage to subadult form withmombergensis type of carina. The separation ofmedium ontogenetic to subadult stages of N.longa and N. balkanica is difficult. The differ-ences are gradual, because also N. balkanicadisplays a slender medium to subadult ontoge-netic stage with mombergensis type of carinawithout pronounced main cusp that cannot bewell separated from specimens of the same on-togenetic stage of N. longa. In the present paperall very long, very slender medium ontogeneticto subadult stages with mombergensis type ofcarina and somewhat foreward-shifted basalcavity are attributed to N. longa. The adults ofN. longa are long, but not so slender as the me-dium ontogenetic to subadult stages. The cusp isterminal, not pronounced, erect or posteriorlyinclined.

Both N. balkanica and N. longa are typicalrepresentatives of the Southern Tethyan LowerLadinian beds, where they have the same range.In the northern Tethys, these species are mostlyrare.

N. pseudolonga KOVÄCS, KOZUR &MIETTO, 1980 is a somewhat earlier juvenilestage (late juvenile to early medium ontogeneticstage) than N. longa and therefore a junior syno-nym of this species. In this ontogenetic stage thecarina has still the constricta type with the sec-ond last denticle as cusp. In contrast to N. con-stricta already in this ontogenetic stage the basalcavity is somewhat foreward-shifted and not ter-minal with respect to the keel end as in the sameontogenetic stage in N. constricta. Moreover,the unit is longer in this ontogenetic stage com-pared with the relatively short juvenile to me-dium ontogenetic stages in N. constricta. In stillearlier juvenile stages the unit is shorter and thebasal cavity is not foreward-shifted. Such formsare inseparable from juvenile N. constricta.Occurrence: Reitziites reitzi - Zone to Eoprot-rachyceras curionii - Zone. Common in the

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Southern Tethys, in the R. reitzi - Zone some-times rare. Mostly rare in the Fassanian of theNorthern Tethys.

Neogondolella balkanicaBUDUROV & STEFANOV, 1975(P1.1, Fig. 22; PL 2, Figs. 3, 6, 9;

PL 3, Figs. 3, 6, 9, 27-29)

Remarks: N. balkanica is restricted to ratherslender forms with platform brim behind the er-ect cusp, as discussed under N. cornuta. N. su-hodolica BUDUROV & STEFANOV, 1973 hasa shorter and broader platform. According toBUDUROV & STEFANOV (1975) N. suhodol-ica has evolved from N. balkanica, but no exactrange data has been given and both forms havebeen derived from different localities from thesame stratigraphic level (excentrica - Zone ac-cording to BUDUROV & STEFANOV, 1973 a,1975).

N. balkanica evolved at the base of the Reit-ziites reitzi - Zone from advanced N. cornutawith erect terminal cusp by development of aplatform brim or at least by separation of thecusp from the posterior platform margin. In theR. reitzi - Zone the brim is always very narrowor even missing, but the cusp is not fused withthe platform end in N. balkanica. This is a gooddistinction against advanced N. cornuta.

N. longa BUDUROV & STEFANOV, 1973 isalso very similar, but more slender. However,the holotype is a subadult form with momber-gensis type of carina, in which the cusp is notyet pronounced. All N. balkanica display a lon-ga stage during their ontogenesis, but this doesnot exclude that N. longa is an independent spe-cies. Adults of N. longa are long and slender anddisplay a rather small, terminal, inclined or erectcusp.Occurrence: Reitziites reitzi - Zone to Eoprot-rachyceras curionii - Zone. Common in theSouthern Tethys, sometimes rare in the R. reitzi -Zone. Mostly rare in the Fassanian of the North-ern Tethys.

Neogondolella mesotriassica(KOZUR & MOSTLER, 1982)

(PL 2, ? Figs. 12, 15, 20; PL 4, Figs. 7-9)

Remarks: The type series of this species aremedium to subadult ontogenetic stages. Earlyjuvenile forms are of constricta type. Already inmedium ontogenetic stages the basal cavity isforeward-shifted with respect to the keel end.The platform end in these forms is typically up-turned to a narrow ridge. In adult forms on oneor both side denticles evolve on this ridge. Theseforms have been often erroneously placed into P.bifurcata BUDUROV & STEFANOV, fromwhich they are clearly distinguished by the verylow middle part of the carina.

Most specimens of morphotype gamma of N.constricta sensu KOVÄCS et al. (1990) belongto this species.Occurrence: Very characteristic form of theFassanian that starts at the base of the R. reitzi -Zone. Transitional forms to N. cornuta occur inthe uppermost P. trinodosus - Zone (Asseretoce-ras camunum horizon).

Paragondolella alpina(KOZUR & MOSTLER, 1982)

(? PL 2, Figs. 2, 5, 8)

Remarks: This very characteristic Fassanianguide form is easily recognizably by its free bla-de and the relatively high denticles throughoutthe entire carina. In juvenile, medium and sub-adult ontogenetic stages the platform is narrowand the difference to the free blade is then not sopronounced.

KOVÄCS (1983) described a 'Gondolella'szaboi from the same stratigraphic level as P. al-pina. As an adult form he figured a subadultspecimen with narrow platform. The holotype isa late juvenile to medium ontogenetic stage. Allthe figured specimens by KOVÄCS (1983) areidentical with late juvenile to subadult ontoge-netic stages of P. alpina and therefore 'G.' sza-boi is a junior synonym of this species.

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KOVÂCS (1983) did not discuss the relationsbetween 'G. ' szaboi and P. alpina, but comparedit only with P. trammeri (KOZUR). KOVÂCS(in GAETANI, 1993) recognized already theabove synonymy and mentioned 'G. ' szaboinow as 'G. 'alpina szaboi. There is no reason toseparate the two contemporaneous forms assubspecies. The platform width is not only de-pendent from the ontogenetic stage, but alsosomewhat variable in adult form. KOZUR &MOSTLER (1982) have figured broader speci-mens, in which the free blade is especially wellrecognizable. As holotype a specimen with rel-atively narrow platform was selected that is on-ly a little broader than the platform of the onlyadult (or subadult) specimen, figured byKOVÂCS (1983) and in the diagnosis of P. al-pina by KOZUR & MOSTLER (1982) theplatform was described as narrow to moderate-ly wide.

Occurrence: P. alpina is a characteristic formof the P. reitzi -Zone and the lower Nevadites -Zone. Its first occurrence is at the base of the P.reitzi - Zone. Transitional forms to the P. hanbu-logi group occur in the uppermost P. trinodosus- Zone (Asseretoceras camunum horizon). Theyhave not yet a free blade and represent mostprobably a new species. In any case they cannotbe placed into P. alpina.

Few specimens are very little arched in lateralview (PI. 2, Figs. 2, 5, 8). They may represent anindependent species near related to P. alpina.However, so far only juvenile forms of this typeare knowri\ For the moment, they are determinedas P. cf. alpina (KOZUR & MOSTLER).

Paragondolella excelsa MOSHER, 1968(PI. 4, Fig. 6)

Remarks: Adults of typical P. excelsa, conspe-cific with the holotype, display a very high cari-na decreasing uniformly to a posterior nodelikedenticle (cusp). There is a wide platform brimbehind the last denticle. Such forms appear atthe base of the Reitzi - Zone. In the Illyrian and

Fassanian there is a similar form with very highanterior carina, that has in adult stages (like inall juvenile P. excelsa) one or two, rarely threesmall denticles behind the cusp that reaches toor nearly to the posterior margin. These formswill be described in a separate paper as new sub-species of P. excelsa.

There is still a third species of the P. excelsagroup with very high anterior carina, but in con-trast to P. excelsa, the carina becomes not muchlower toward the posterior end and reaches untilthe posterior platform margin. Such forms havebeen determined by SASHIDA et al. (1993) as'Neogondolella' bulgarica (BUDUROV &STEFANOV) and therefore regarded as Pelso-nian in age. However, the accompanyingradiolarians are clearly Ladinian. This new spe-cies is common in our material from fissure fill-ings of pelagic limestones in the Weiße Wandsection (PI. 1, Figs. 1-5, 11-19). Unfortunately,the few complete preserved specimens have be-en crashed during the sticking of the material.Therefore this species will be described in a lat-er paper.

Paragondolella trammeri KOZUR, 1972(PI. 2, figs. 19, 23, 24; PL 3, Figs. 10,13, 16)

Remarks: Paragondolella trammeri is one ofthe most important Ladinian conodont guideforms, easily recognizable both in juvenileforms and adults, common both in the Southernan Northern Tethys, in open pelagic and insomewhat restricted pelagic deposits. There areonly minor changes in the P. trammeri popula-tions during the Ladinian. The holotype is a latejuvenile to medium ontogenetic stage of an ad-vance P. trammeri from the Fassanian-Lango-bardian boundary, figured and described in KO-ZUR & MOCK (1972). An adult form from thesame sample has been figured as Gondolellatrammeri trammeri in KOZUR & MOCK(1972). These adults are 500-600 |Jm long, in theLongobardian still a little shorter. The keel ishigh and narrow. The basal cavity is with respect

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to the keel wide (almost as wide as the widestpart of the keel).

Primitive forms of P. trammeri from the R. re-itzi - Zone, described as P trammeri praetram-meri (KOZUR & MOSTLER, 1982) are verysimilar to advanced P. trammeri and can be onlyseparated in adults. They are somewhat larger(600-700 Jim), the keel is broader and shallowerand the basal cavity is distinctly narrower (oftenonly as half as wide as the surrounding keel).Moreover, often the platform is parallel-sidedand the anterior decrease of the platform widthbegins only before the midlength. However, thelatter feature cannot be used for separation ofthe two subspecies, because also in some adultsof P. trammeri praetrammeri the anterior de-crease of the platform width begins after themidlength and in some advanced P. trammeritrammeri the anterior decrease of the platformwidth starts only in front of the midlength. Theadults of medium advanced forms from the up-per R. reitzi - Zone and from the Nevadites - Zo-ne are as large as the primitive P. trammeri prae-trammeri, but the keel is already relatively nar-row and the basal cavity is only a little narrowerthan the widest part of the platform. These formsare transitional between P. trammeri praetram-meri and advanced P. trammeri.

Already medium ontogenetic stages of P.trammeri trammeri and P. trammeri praetram-meri with terminal cusp, but still relatively nar-row platform are not more definitely separable,because also in medium ontogenetic stages of P.trammeri praetrammeri the keel is narrow andhigh and the basal cavity is nearly as wide as thewidest part of the keel. Juvenile forms of ad-vanced and primitive P. trammeri are insepara-ble. In these forms the cusp is in penultimate po-sition and a small denticle is present behind thecusp. As typical for all juvenile Paragondolella,the keel is narrow and high, the basal cavity liesterminal with respect to the keel end that flaredaround the basal cavity.

The transition from primitive to advanced P.trammeri is gradual and does not mark any dis-tinct conodont boundary. This is well document-ed by the fact that KRYSTYN (1983) and

KOVÂCS et al. (1990) failed to separate theseforms despite the fact that they believe that thedevelopment of typical (advanced) P. trammerifrom more primitive representatives is a first or-der conodont event (see also chapter "Strati-graphic evaluation of the conodont faunas").KRYSTYN (1983, p. 257) rejected P. trammeripraetrammeri and regarded it as typical P. tram-meri. He was so sure in his determination that hewrote that the conodont fauna with P. trammeripraetrammeri (= typical 'G. ' trammeri sensuKRYSTYN, 1983) is „...ganz sicher für die Nev-adites - Zone und keineswegs für die Parakell-nerites -Zone charakteristisch...". However, ac-cording to a personal communication KRYS-TYN has now restudied the ammonoid fauna ofthe stratum typicum of P. trammeri praetram-meri and he could now in contrast to his abovestatement confirm that this fauna belongs to hisKellnerites fauna (= R. reitzi - Zone) as pointedout by KOZUR & MOSTLER (1982). After thisconfirmation of the age he finds now also differ-ences between P. trammeri praetrammeri and P.trammeri trammeri. In the discussion on the An-isian/Ladinian boundary field workshop he ex-plained repeatedly that he had misinterpreted P.trammeri praetrammeri because KOZUR &MOSTLER (1982) have only figured adultspecimens and no juvenile form. This explana-tion is strange, because juvenile forms of all P.trammeri (both very primitive and advancedforms) are inseparable. Moreover, in the paperKOZUR & MOSTLER (1982) not only adultforms, but also a juvenile P. trammeri praetram-meri has been figured. What we can see fromKRYSTYN's statement is the fact that the prim-itive P. trammeri are so similar to P. trammeritrammeri (advanced P. trammeri) that KRY-STYN could not separate these forms as long hebelieved that the ammonoid fauna belongs to hisNevadites fauna. The later separation (afterknowing that the ammonoid data by KOZUR &MOSTLER are correct) can be only evaluatedas the attempt to maintain his conodont boun-dary at the base of the Nevadites fauna.

KOVÄCS et al. (1990) wrote: 'KOZUR &MOSTLER (1982) also introduced a new spe-

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cies to describe the forerunner of G. trammeri;however, the holotype of their G. praetrammeri(see MOSTLER & KOZUR, 1982, PL 5, Figs.5a-b), is a true G. trammeri specimen withstraight upper edge of the carina. Consequently,'G. praetrammeri' KOZUR & MOSTLER,1982, should be treated as a junior synonym ofG. trammeri KOZUR, 1972.' Superficial evalu-ation of papers and often intentionally wrongquotation are characteristic for papers ofKOVÄCS or KOVÀCS et co-authors, if he hasmade the quotations. Thus, KOZUR & MOST-LER (1982) have not introduced a new speciesGondolella praetrammeri, but a new subspeciesGondolella trammeri praetrammeri. The holo-type was not figured in a paper MOSTLER &KOZUR, 1982, PI. 5, Fig. 5, but in KOZUR &MOSTLER, 1982, PI. 1, Fig. 5. In species rank,P. trammeri praetrammeri is, of course, a P.trammeri KOZUR, 1972 as pointed out by KO-ZUR & MOSTLER (1982) and therefore it gi-ves no new data to write that 'Gondolella'(trammeri) praetrammeri is a 'G. ' trammeri.Moreover, KOZUR & MOSTLER (1982) havenot separated P. trammeri praetrammeri from P.trammeri trammeri by having no straight upperedge of the carina, but by the above mentioneddifferences in the keel, basal cavity and size ofthe unit. These differences have not been dis-cussed by KOVÄCS et al. (1990).

The position of 'Gondolella' eotrammeriKRYSTYN (1983) is not yet clear. The figuredadult forms are rather large (around 800 jamlong). The holotype has a flat platform and alsoin the anterior part a rather high carina. Platformshape and development of the keel and basalcavity are identical with P. trammeri praetram-meri. The adult paratype is not so flat and the ca-rina in the anterior part is rather low. This formcorresponds to P. trammeri praetrammeri. Alsothe other paratypes, juvenile to medium ontoge-netic stages cannot be separated from P. tram-meri praetrammeri. Thus, the holotype is eitheran extreme form or, more probably, an indepen-dent species with transitional character betweenP. hanbulogi SUDAR & BUDUROV and P.trammeri.

Unfortunately, in contrast to the opinion ofKRYSTYN (1983), the Epidaurus section is un-suitable for exact correlation between the Mid-dle Triassic ammonoid and conodont succes-sion, because the sediments are strongly con-densed. Therefore the age of P. eotriassica s.str.(restricted here to the holotype) is unknown.Similar forms are so far unknown from the Alpsand Western Carpathians.

In the following some new taxa will be de-scribed.

Genus NeogondolellaBENDER & STOPPEL, 1965

Type species: Gondolella mombergensisTATGE, 1956

Neogondolella cornuta ladinica n. subsp.(PI. 3, Figs. 2, 5, 8, 11, 14, 17; PI. 4, Fig. 21)

Derivatio nominis: According to the strati-graphic occurrenceHolotype: The specimen figured on PI. 3, Figs.2, 5, 8, rep.-no. KKM 1993 111-17Locus typicus: Zelenitza forest road W Loibl-pass, Southern Karawanken Mts.Stratum typicum: Red pelagic limestones ofthe Loibl Formation, sample ZE 6a.Material: More than 100 specimens, mostlyfrom the here not described Karwendel se-quence.Diagnosis: A slender N. cornuta with relativelylong, somewhat upward bent posterior platformbehind the distinctly foreward-shifted basal cav-ity. Keel behind the basal cavity distinct on thelower side of the upward bent platform portion.Description: Unit in primitive forms large, inadvanced forms medium sized. Platform long,slender, in large part nearly parallel-sided, withrounded posterior margin. Carina low, the 13-18denticles are highest in the anterior third and inthe posterior part. Middle part of carina lowestand more or less fused. Cusp distinct, fused with

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the posterior platform margin, inclined posteri-orly to nearly erect.

Platform in lateral view arched, relativelylong posterior part behind the basal cavity up-ward bent. Keel broad with distinct basal fur-row. Basal cavity distinctly foreward-shifted,with two pits, connected by a furrow. The keelcontinues on the lower surface of the upwardbent part.

The fore ward-shifting of the basal cavity canbe already observed in medium ontogenetic sta-ges, in which the continuation of the keel behindthe basal cavity reaches somewhat on the lowersurface of the upward bent part. These mediumontogenetic stages have a mombergensis type ofcarina with higher denticles in the anterior andposterior part of the carina and without distinctmain cusp. Early ontogenetic stages are of con-stricta type, like in all Upper Anisian and Ladin-ian Neogondolella species.Occurrence: Fassanian. In the R. reitzi - Zoneprimitive, large forms occur, in the E. curionii -Zone advanced, smaller forms are present.Remarks: N. cornuta ladinica n. subsp. hasevolved from N. cornuta cornuta at the base ofthe R. reitzi - Zone. In N. cornuta cornuta theupward bent posterior part of the platform isshort and the continuatibn of the keel behind thebasal cavity does not continue on the lower sideof the upward bent part of the platform. Primi-tive representatives of N cornuta ladinica fromthe R. reitzi - Zone are still as large as N. cornu-ta cornuta, but the upward bent part of the plat-form behind the basal cavity is already long andthe keel continues on the lower surface of thispart. Advanced specimens from the upper Fas-sanian are smaller, but otherwise identical. Thetrend to size reduction can be observed in all La-dinian Neogondolella lines.

Medium ontogenetic stages of N. mesotriassi-ca (KOZUR & MOSTLER, 1982) are some-times similar to the same ontogenetioc stages ofN. cornuta ladinica, especially specimens withrounded posterior end without v-shaped inci-sion. However, the posterior end of the platformis always strongly and steeply upturned in formof a narrow sharp ridge in medium ontogenetic

stages of N. mesotriassica, whereas the posteriorplatform end is rather low in N. cornuta ladini-ca.

Neogondolella aldae n. sp.

1965 Gondolella constricta n. sp. pars -MOSHER & CLARK, p. 560, PI. 65, onlyPL 65, Figs. ? 14, ? 15, ? 19, 22.

1965 Gondolella mombergensis TATGE, pars -MOSHER & CLARK, p. 560, PL 65, onlyFigs. 20, ? 23, ? 26, ? 27, 29.

1965 Gondolella navicula HUCKRIEDE -MOSHER & CLARK, p. 560-561, PL 66,figs. 10, 17-21, non ! Figs. 14, 16.

1973 a Neogondolella basisymmetrica sp. n.,pars - BUDUROV & STEFANOV, p.837-838, PL 3, Figs. 4-8, non ! Figs. 1-3(holotype).

1979 Neogondolella basisymmetrica BUDU-ROV & STEFANOV - BUDUROVGANEV & STEFANOV, PL 1, Figs. 3, 4.

1984 Gondolella constricta MOSHER &CLARK, 1965, pars - NICORA &KOVÀCS, p. 144-148, only the speci-mens on PL 7, Fig. 11; PL 8, Fig. 3; PL 10,fig. 1.

1984 Gondolella mombergensis mombergensisTATGE, 1965 - NICORA & KOVÀCS,p. 149, 150, PI 7, Figs. 3, 15; PL 8, Figs.6-12; PL 9, Figs. 7, ? 8, 9, 10, 12-14; PL10, Figs. 3, ? 5; non! PL 7, Figs. 2, 5, 7,13; PL 9, Fig. 11.

1984 Gondolella mombergensis longa (BUDU-ROV & STEFANOV, 1973) - NICORA& KOVÀCS, p. 150, PL 10, Figs. 2, 4,6-10.

Derivatio nominis: In honour of Prof. Dr. AidaNicora, Milano.Holotypus: The specimen, figured by NICORA& KOVÀCS, 1984, PL 10, Fig. 9 as Gondolellamombergensis longa BUDUROV & STEFA-NOV

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Locus typicus: Site A, Fossil Hill, HumboldtRange, Nevada (see NICORA & KOVÀCS,1984).Stratum typicum: Sample N 49, middle PridaFormation, Nevadites humboldtensis beds oflower Occidentalis - Zone (see NICORA &KOVÀCS, 1984).Material: From the Alps (KOZUR & MOS-TLER, in prep.) 12 specimens are known. Theholotype is therefore chosen from the Nevadianmaterial, where this form is dominating.Diagnosis: Platform moderately broad to slen-der, also in adult stages mostly with distinct con-striction near the posterior end, widest in orsomewhat before the midlength. Platform partafter the constriction short and broad or long andslender. Widest before or in the midlength. Cari-na in adults low, highest in the anterior third, inthe middle and partly also in the posterior partfused to a ridge. Cusp distinct, terminal, not fu-sed with the posterior platform margin, withoutor rarely with narrow platform brim behind thecusp. Basal cavity somewhat foreward-shiftedwith respect to the keel.Description: Adults moderately large to large.Platform moderately broad to slender. Posteriorpart also in adult forms constricted. Constrictionsometimes only on the lower surface visible, ingeneral distinct. Platform part after the constric-tion short and broad or long and slender. Plat-form end rounded. Platform widest before or inthe midlength. Toward the posterior platformend the width decreases only very few or re-mains nearly constant. Toward the anterior endthe platform width decreases more rapid. Lateralplatform margins moderately wide and moder-ately upturned, with honeycomb microsculpture.Furrow on both sides of the carina smooth andrelatively deep. Carina in adults in the middlepart and most of the posterior part fused to a lowridge. The penultimate denticle and the terminalcusp are mostly free, but sometimes also the pe-nultimate denticle is included into the fused rid-ge that rarely reaches even the cusp that is poste-riorly inclined or nearly erect. Denticles in theanterior part of the carina highest and in theirupper part always free. Distinct cusp always ter-

minal, not fused with the posterior platformmargin. Platform brim behind the cusp missingor very narrow. Keel narrow with distinct basalfurrow. Basal cavity somewhat to distinctlyforeward-shifted with respect to the keel end.Two pits, connected by a furrow are present inthe basal cavity.

Early juvenile stages are of constricta typeand not separable from early juvenile stages ofother late Illyrian and Fassanian Neogondolellaspecies. Late juvenile forms are of mombergen-sis type (denticles all of the same size or on themiddle part a little lower, no distinct main cusp).Already in this ontogenetic stage the basal cav-ity is a little foreward-shifted (see NICORA &KOVÀCS, 1984, PI. 7, Fig. 11, late juvenile N.aldae, erroneously placed into N. constricta thathas in this ontogenetic stage always a terminalbasal cavity). Medium ontogenetic stages havealready a terminal cusp, but the denticles are notyet fused (well recognizable in the specimen fig-ured by MOSHER & CLARK, 1965, PI. 66,Figs. 10, 17, 18 and assigned to Gondolella nav-icula HUCKRIEDE). Until this ontogenetic sta-ge the constriction is very prominent. Subadultforms are already very similar to adult ones, butonly in the middle part of the carina the denti-cles are fused to a ridge.

Occurrence: Fassanian {Meeki- to Subasperum- Zone) of Nevada. Dominant form in the UpperMeeki- and Occidentalis - Zone, rare in the Sub-asperum - Zone. Rare in the Fassanian of the Al-ps and of Bulgaria.Remarks: Nearly all forms, figured by NICO-RA & KOVÀCS (1984) as 'Gondolella' mom-bergensis from Nevada belong to N. aldae n. sp.that can be subdivided into two subspecies (seebelow). Only the few figured juvenile formsfrom the Paraceratites vogdesi and P. crickibeds belong to N. cornuta or a similar form. Al-so the assumed adult form from these beds (NI-CORA & KOVÀCS, 1984, PI. 9, Fig. 11) is a la-te juvenile to medium ontogenetic stage of aNeogondolella of the mombergensis group. N.aldae is therefore restricted to the Meeki-, Occi-dentalis- and lower Subasperum - Zone.

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Adults of N. mombergensis have mostly 3slender large teeth in the posterior part of the ca-rina. A distinct cusp is mostly not present. In so-me forms the penultimate teeth is the cusp (e.g.in N. mombergensis prava). Adult N. momber-gensis have in general no distinct posterior con-striction.

Some representatives of the N. mombergensisgroup s.l. from the German Basin display a dis-tinct terminal cusp. Among these forms Neo-gondolella haslachensis TATGE, 1956 is asmall form with very slender long, mostly separ-ate teeth. It is unrelated to N. aldae and an en-demic species restricted to the German Basin.Other forms (only from the Hauptmuschelkalkmo 1) belong to N. cornuta (differences to N. al-dae, see below). A few forms from the Haupt-muschelkalk mo 2 belong to N. balkanica (dif-ferences to N. aldae, see below). A further newspecies from the German Hauptmuschelkalk(only in the mo 2) with distinct terminal cuspdisplays slender long denticles that are not fusedto a ridge in the middle part of the carina in adultforms. Subordinately, forms similar to N. aldaeare present in the German Hauptmuschelkalk(mo 1 ß, mo 2).

In N. cornuta BUDUROV & STEFANOV,1973 the very prominent terminal cusp is alwaysfused with the posterior platform margin, theposterior constriction of the platform in adultforms is mostly missing. The lateral platformmargins are mostly over a longer distance par-allel.

N balkanica BUDUROV & STEFANOV,1975 displays an prominent, always erect cusp,surrounded by a distinct platform brim. In richerpopulations of N. aldae there are few specimenswith very narrow platform brim and nearly erectcusp (see NICORA & KOVÀCS, 1984, P. 9,Fig. 8) that are hardly separable from very prim-itive N balkanica. Either the intraspecific vari-ability of both species overlaps a little (moreprobably) or primitive N. balkanica are presentin the Nevadian material.

The paratypes of M basisymmetrica BUDU-ROV & STEFANOV and the specimen figuredby BUDUROV, GANEV & STEFANOV

(1979) as TV. basisymmetrica corresponds toslender morphotypes oïN. aldae aldae. Howev-er, the holotype of N. basisymmetrica (BUDU-ROV & STEFANOV, 1973 a, pi. 3, figs. 1-3) isa medium ontogenetic stage without a cusp,with upward bent posterior platform and dis-tinctly fore ward-shifted basal cavity. The lastdenticle is rather small. The holotype does notfit in the diagnosis of the species (last denticlesnot larger than the other ones, but distinctlysmaller!), but corresponds to the drawing byBUDUROV & STEFANOV (1973 a, fig. 5).Thus, the holotype was not chosen erroneously.It is very similar to N. bacalovi BUDUROV &STEFANOV, 1973, but the basal cavity is not sostrongly foreward-shifted and its margins arenot so strongly protruding as in N. bacalovi. Theholotype belongs to the N. bacalovi group, but ismore primitive. The other figured specimens ofN. basisymmetrica (BUDUROV & STEFA-NOV, 1973 a, pi. 3, figs. 4-8; BUDUROV, GA-NEV & STEFANOV, 1979, PI. 1. Figs. 3, 4) be-long clearly to an other species (N. aldae).

The differences to B. longa BUDUROV &STEFANOV, 1973 are discussed under N aldaeposterolonga n. subsp.

Neogondolella aldae aldae n. subsp.

1965 Gondolella constricta n. sp. pars -MOSHER & CLARK, p. 56O,P1. 65, onlyPI. 65, Figs. ? 14, ? 15, ? 19, 22.

1965 Gondolella mombergensis TATGE, pars -MOSHER & CLARK, p. 560, PL 65, onlyFigs. 20, ? 23, ? 26, ? 27, 29.

1965 Gondolella navicula HUCKRIEDE -MOSHER & CLARK, p. 560-561, PI. 66,figs. 10, 17-21, non ! Figs. 14, 16.

1973 a Neogondolella basisymmetrica sp. n.,pars - BUDUROV & STEFANOV, p.837-838, PI. 3, Figs. 4-8, non ! Figs. 1-3(holotype).

1979 Neogondolella basisymmetrica BUDU-ROV & STEFANOV - BUDUROV, GA-NEV & STEFANOV, PI. 1, Figs. 3, 4.

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1984 Gondoleila constricta MOSHER &CLARK, 1965, pars - NICORA &KOVÂCS, p. 144-148, only the speci-mens on PL 7, Fig. 11; PI. 10, fig. 1.

1984 Gondolella mombergensis mombergensisTATGE, 1965 - NICORA & KOVÀCS,p. 149, 150, PL 7, Figs. 3, 15; PL 8, Figs.7-12; PL 9, Figs. ? 8, 9, 12-14; PL 10,Figs. ? 3,5; non! PL 7, Figs. 2,5, 7,13; PL8, Fig. ? 6; PL 9, Figs. ? 7, ? 10, 11.

1984 Gondolella mombergensis longa (BUDU-ROV & STEFANOV, 1973),pars - NICO-RA & KOVÂCS, p. 150, PL 10, Figs. 2,8-10, non ! Figs. ?4, 6,7.

Derivatio nominis, holotypus and stratumtypicum: As for the species.Diagnosis: With the character of the species(description see above). Platform slender tomoderately wide. Posterior platform part afterthe constriction always short, comprising on thecarina only the cusp or the cusp and the penulti-mate tooth.Occurrence: Meeki- and Occidentalis - Zone ofNevada. Reitzi - Zone and Nevadites - Zone ofthe Alps and Bulgaria.Remarks: N. aldae posterolonga n. subsp. isdistinguished by a rather long, slender con-stricted part of the always slender to very slen-der platform. The narrow platform behind theconstriction comprises on the carina the maincusp and two further denticles. Already latejuvenile forms with equally sized denticles with-out terminal cusp and very narrow posteriorconstricted platform display a distinctly fore-ward-shifted basal cavity.

The differences between the two subspeciesare rather gradual. Few specimens, placed byNICORA & KOVÄCS (1984) into 'G.' mom-bergensis mombergensis belong rather to N. al-dae posterolonga n. subsp., but have clearlytransitional character to N. aldae aldae. Howev-er, the ranges of the two subspecies are different.N. aldae posterolonga starts later (upper Meeki-or lower Occidentalis - Zone) than N. aldae al-dae (lower Meeki - Zone, lower Reitzi - Zone)and ranges in younger beds (lower Subasperum

- Zone). The more important feature for separa-tion seem not to be the width of the platform, butthe width of the constricted posterior part of theplatform that is longer and narrower in N. aldaeposterolonga. Such forms have, of course, al-ways a slender to very slender platform, butslender platforms display also some N. aldae al-dae with short constricted posterior platformpart.

Neogondolella aldae posterolonga n. subsp.

Derivatio nominis: According to the long con-stricted posterior part of the platform.Holotypus: The specimen figured by NICORA& KOVÂCS, 1984, PL 10, Fig. 7 as N. mom-bergensis longa.Locus typicus: Site A, Fossil Hill, HumboldtRange, Nevada (see NICORA & KOVÂCS,1984).Stratum typicum: Sample N 50, middle part ofPrida Formation, lower Occidentalis- ZoneMaterial: Only few doubtful juvenile forms arepresent in our collections, therefore the holotypeis chosen from the Nevadian material.

1984 Gondolella constricta MOSHER &CLARK, 1965, pars - NICORA &KOVÂCS, p. 144-148, only the speci-mens on PL 8, Fig. 3.

1984 Gondolella mombergensis mombergensis,pars - NICORA & KOVÂCS, p. 149-150,only PL 8, Fig. ? 6; PL 9, Figs. ? 7, ? 10.

1984 Gondolella mombergensis longa (BUDU-ROV & STEFANOV, 1973), pars - NICO-RA & KOVÂCS, p. 150, only the speci-mens on PL 10, figs. ? 4, 6, 7.

Diagnosis: With the character of the species(description see under N. aldae). Platform veryslender to slender, widest before the mid-lengthof the unit, with long, narrow, constricted poste-rior part. Juvenile forms display a very narrow,constricted posterior part of the platform. Al-ready these late juvenile stages that have not yet

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a terminal main cusp, but denticles of the samesize throughout the carina (see NICORA &KOVÀCS, 1984, PL 3, Fig. 3, specimen errone-ously assigned to 'Gondolella ' constricta) havea distinctly fore ward-shifted basal cavity.Occurrence: ? Upper Meeki - Zone, Occiden-talis- to lower Subasperum - Zone of Nevada.Remarks: Differences to N. aldae aldae see un-der this subspecies.

Only a part of 'Gondolella' mombergensislonga sensu NICORA & KOVÂCS (1984) be-longs to N. aldae posterolonga n. subsp., theothers are slender N. aldae aldae with short con-stricted posterior platform portion.

N. longa BUDUROV & STEFANOV, 1973displays in the same ontogenetic stages a stilllonger, compared with the length narrower plat-form, widest distinctly behind the midlength.

Only adult forms have an indistinct terminalcusp.

Neogondolella ? postpridaensis n. sp.

1981 Gondolella pridaensis sp. n., pars - NICO-RA, KOZUR & MIETTO, p. 762-763,PI. 89, Figs. 5, 7.

Derivatio nominis: According to the strati-graphic occurrence after N. pridaensis s.str.Holotypus: The specimen figured by NICORA,KOZUR & MIETTO (1981), PI. 89, Figs. 5 a-c.Locus typicus: Section C, Fossil Hill, Hum-boldt Range, Nevada (locality data see SILBER-LING & NICHOLS (1982).Stratum typicum: Sample N 77, lower part ofEoprotrachyceras beds, lower part of UpperFassanianMaterial: Typical representatives of this speciesare so far only known from Nevada. Thereforethe holotype is chosen from this material.Diagnosis: Platform slender. Posterior endmostly pointed, partly narrowly rounded, margi-nally only slightly upturned. In advanced formsthe platform is mostly very slender and ends of-

ten before the last denticle. Outer part of plat-form surface with honeycomb microstructure.

Carina very high, highly fused, but the upperpart of the slender denticles (up to 14) is neverfused to a wavy line. The last denticle is fusedwith the platform margin. It is mostly smallerthan the following 1-2 denticles. The second orthird denticle is largest, but no distinct cusp ispresent. The other denticles of the carina havenearly the same size. Only the 2-3 anterior denti-cles are smaller. Sometimes the size of the denti-cles varies irregularly with a bigger denticle ne-ar the midlength of the carina. Lower surfacewith high, narrow keel. Loop elongated, posteri-or end narrowly rounded or even pointed. Thebasal furrow is broad and distinct. Behind the pitthat is situated at the beginning of the posteriorwidening of the keel a distinct short furrow con-tinues that is at its end not widened into a dis-tinct second pit, typically for Neogondolella.Occurrence: Middle and lower part of upperFassanian of North America.Remarks: Paragondolella ? pridaensis (NICO-RA; KOZUR & MIETTO, 1981) from the Illyr-ian of North America and Japan displays a dis-tinct cusp, the carina is lower and in adult formsgenerally fused to a wavy line. The teeth of thecarina are broadly triangular.

Both P ? pridaensis and especially P. ? post-pridaensis are not typical Paragondolella ac-cording to their basal cavity. However, a distinctsecond pit on the end of furrow behind the pit,typically for all Neogondolella species, is alsonot present. Because the basal cavity of P. ? pri-daensis is near related to the basal cavity of typ-ical Paragondolella, both species are placed ten-tatively into Paragondolella.

Paragondolella ? pridaensis posteroacutan. subsp.

(PI. 1, Figs. 7, 8, 10,20,21,24)

1981 Gondolella pridaensis sp. n., pars - NICO-RA, KOZUR & MIETTO, p. 762-763,only P. 89, Fig. 3.

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Derivatio nominis: According to the pointedposterior end.Locus typicus: Weiße Wand, W Loiblpaß,Southern Karawanken Mountains (SouthernAlps), Carinthia (Austria).Stratum typicum: Fissure filling of red pelagiclimestone (Weiße Wand Member) of the LoiblFormation in the uppermost part of the ContrinFormation.Material: 15, mostly broken specimens.Diagnosis: Unit small. Platform pointed both atthe anterior and posterior end, widest in or be-hind the mid-length. Prominent terminal cuspfused with the platform end. Carina high, denti-cles in the posterior part nearly of the same size,in the anterior part somewhat longer.Description: Small conodonts with unusualplatform shape, sharply pointed at both ends.The platform sometimes does not reach the pos-terior edge of the terminal, prominent, posterior-ly inclined cusp that is fused with the posteriorplatform margin. The platform is widest in orsomewhat behind the midlength, their lateralmargins are only slightly thickened and slightlyto moderately upturned. The smooth furrowalong the carina is rather wide. The carina ishigh, the denticles are not fused to a ridge. In theposterior part the denticles have the same lengthor their length increases slightly anteriorly, likein the anterior part of the carina. The keel is nar-row, with rather wide basal furrow. Its posteriorend is pointed. The basal cavity is only slightlyforeward-shifted with respect to the keel end. Atthe beginning of the basal cavity lies a rather lar-ge pit, followed by a short furrow that does notwiden to a secondary pit.Occurrence: Lower Ladinian of Nevada. TheEuropean occurrences are somewhat doubtful inthe age (see stratigraphie evaluation of the co-nodont faunas). However, P. ? pridaensis poste-roacuta is seemingly a little more advanced thanP. ? pridaensis pridaensis from the Paracera-tites cricki beds of Nevada. Because these bedsare contemporaneous with upper P. trinodosusZone of the European Tethys, P. ? pridaensisposteroacuta should belong to the latest Illyrianor earliest Fassanian. Because the Radiolarian

fauna is pre-Fassanian (older than Reitzi -Zone),a latest Illyrian age is probably, but somewhatolder ages cannot be excluded from the faunalevidence.Remarks: P. pridaensis pridaensis (NICORA,KOZUR & MEETTO, 1981) has always a roun-ded posterior end and the denticles of the carinaare strongly fused.

N. acuta (KOZUR, 1972) displays also apointed posterior platform end. This species isclearly distinguished by its low carina.

Genus Budurovignathus KOZUR, 1988

Synonyma: Carinella BUDUROV, , 1973; Se-phardiella MARCH et al., 1988.Type species: Polygnathus mungoensis DIE-BEL, 1956.Remarks: The genera Budurovignathus KO-ZUR, 1988 and Sephardiella MARCH et al.,1988 have been established in the same abstractvolume of the Frankfurt ECOS V to replace thehomonym genus Carinella BUDUROV, 1973.Because a homonym was replaced by a new na-me, none of the taxa has an taxonomic advan-tage and the procedure of the first revising au-thor is not necessary.. Therefore the page prior-ity is decisive. Moreover, the real publicationappeared later, in 1990. There, BudurovignathusKOZUR has been published some months be-fore Sephardiella MARCH et al. For these rea-sons, Budurovignathus has clearly the priority.

Budurovignathus gabriellae n. sp.(PI. 3, Figs. 1,4,7, 19,22,25)

Derivatio nominis: In honour of Prof. Dr. Ga-briella Bagnoli, Pisa.Holotypus: The specimen on PL 3, figs. 1, 4, 7,rep.-no. KKM 1993 III-lLocus typicus: Section at the Zelenitza forestroad, W Loiblpass, Southern Karawanken(Southern Alps), Carinthia (Austria).

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Stratum typicum: Sample ZE 6a, red pelagiclimestones of the Loibl Formation, Upper Fassa-nian.Material: 4 specimens.Diagnosis: Smooth Budurovignathus with sig-moidally bent broad platform, broadly roundedplatform end, widely separated long denticles onthe carina and a basal cavity situated at the be-ginning of the posterior third of the unit.Description: Platform relatively broad, flat,with thickened, only slightly upturned platformmargin. The smooth furrow along the carina isnarrow and shallow, the honeycomb microstruc-ture covers therefore the largest part of the plat-form surface. The platform is slightly sigmoidal-ly bent and the asymmetric posterior margin isbroadly rounded. The carina bears few (7-8,widely separated, long, uniformly posteriorly in-clined denticles, somewhat larger in the anteriorpart than in the posterior part. No distinct cusp.

The keel is narrow to moderately wide, slight-ly sigmoidally bent like the platform, but withpointed posterior end. Until the almost not ex-panded basal cavity the basal furrow is distinct.The basal cavity has two pits connected by a rel-atively long furrow. After the posterior pit thebasal furrow is missing or very indistinct.Occurrence: Upper Fassanian of the type local-ity.Remarks: Budurovignathus truempyi(HIRSCH) displays a pointed posterior platformend and the keel is more distinctly expandedaround the basal cavity.

BAGNOLI et al. (1984) figured a form withidentical platform shape from the Ladinian ofSardinia. This form is a little more advanced,because the basal cavity lies only a little behindthe mid-length of the platform. This form occurstogether with a new subspecies of B. truempyi,transitional between typical B. truempyi(HIRSCH) and B. hungaricus (KOZUR &VÉGH). From this it can be concluded that ournew species is a little older, perhaps contempo-raneous with B. truempyi truempyi or even still alittle older. The relatively slightly foreward-shif-ted basal cavity indicates that B. gabriellae is

the oldest known Budurovignathus species, stilltransitional to Neogondolella.

5. Stratigraphie evaluation of the conodontfaunas and some remarks to the Anisian-La-dinian boundary

The conodonts are the most important guideforms for the Triassic biostratigraphy. Most ofthe stage and substage boundaries are better de-fined by conodonts than by any other fossilgroup. Only at the Anisian-Ladinian boundarythe conodonts are said to be stratigraphically un-important or of minor importance. However,this depends on two facts: (1) the Anisian-La-dinian boundary is placed by different authorsand in different faunal realms in different levels.Early Ladinian guide forms of one author be-come Illyrian forms by other authors. By this, II-lyrian-Fassanian ranges are constructed for sev-eral species that would be in reality restricted toone of these substages, if a uniform Anisian-La-dinian boundary will be applied. (2) Both the Il-lyrian and Fassanian (except the upper curionii -Zone) are characterized by smooth gondolellids,a taxonomically difficult conodont group. Manyspecies are used in a wide sense comprising of-ten different species or subspecies that straddlethe Anisian-Ladinian boundary. Sometimes the-se species are subdivided into different morpho-types of different stratigraphie ranges(KOVÄCS et al, 1990). However, in a formtaxonomy all conodont species of a form genusare form taxons and therefore morphotypes.This term should be in the form taxonomy onlyused for intraspecific form groups with the samestratigraphie range. If the Middle Triassic Neo-gondolella- and Paragondolella species areused in a more restricted sense, then they aregood guideforms also for this time-interval.

The main problem of the Middle Triassic stra-tigraphy is the Anisian-Ladinian boundary orbetter to say, the problems that the ammonoidspecialists made with this boundary. The priorityof this boundary is clear and a priority boundary

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should not be changed, if this boundary is notproblematical by large time gap or overlap bet-ween the two zones, with the boundary of whicha stage or substage boundary has been defined.There will be always still a little "better" boun-dary in the future, if we use other fossil groupsor other stratigraphical methods or simply be-cause of the fact that a certain level is betterstudied than another one. The stability of thestratigraphie boundaries requires the recognitionof the priority rules also in the stratigraphy, be-cause otherwise certain schools will try to pressthe international geological community to ac-cept their boundaries.

BÖCKH (1873) established for the first timethe 'Niveau des Ceratites reitzï in the BalatonHighland, one of the first recognized ammonoidzones in the Triassic. MOJSISOVICS (1874) es-tablished the Norian stage as the second stage ofthe Triassic. Due to a later polemic, this stagewas later re-named as Ladinian stage and theterm Norian was used for the former JuvavianUpper Triassic stage. The Norian stage of MOJ-SISOVICS (1874) = Ladinian stage of later usewas the best defined one of the MediterraneanTriassic stages. Already MOJSISOVICS (1874)defined this stage with the 'Horizont des Tra-chyceras reitzï and with the Buchenstein Lime-stone of Gröden/Val Gardena for the lower partand with the 'Zone der Daonella lommeli unddes Trachyceras archelaus' and Wengen Bedsfor the upper part. MOJSISOVICS (1879)slightly redefined this zonation and placed intothe Lower Ladinian (in this time his Lower No-rian) the 'Zone des Trachyceras curionii und desTrachyceras reitzi and in the upper Ladinianagain the Archelaus ammonoid zone and theLommeli bivalve zone. As youngest pre-Ladin-ian ('Norian') zone he regarded the 'Trachyce-ras' trinodosus - Zone. The same zonation wasused by MOJSISOVICS (1882).

In the first comprehensive work about theMediterranean Triassic the terms Fassanian (forthe Lower 'Norian' = Lower Ladinian) and Lan-gobardian (for the Upper 'Norian' = Upper La-dinian) have been introduced. The Fassanianwas defined with the Protrachyceras curionii -

Zone (below) and the Dinarites avisianus - Zo-ne (above) and the Langobardian was defined bythe Protrachyceras archelaus - TJQWZ,. The topof the Anisian was again defined by the top ofthe Trinodosus - Zone. The Curionii - Zonewas at that time used for the whole time intervalfrom the base of the Reitzi - Zone until the topof the Recubariense - Zone as clearly indicatedin several papers. Reitzi - Zone s.l. and Curionii- Zone s.l. have been used in this time in the sa-me sense. Thus, MOJSISOVICS (1882) listedamong the fossils of the 'Zone des Trachycerasreitzi', for instance, Trachyceras reitzi, T. chie-sense, T. recubariense and T. curionii. At thatmoment, when these zones have been again sep-arated, the base of the Ladinian has been placedwithout any exception at the base of the Reit-ziites reitzi - Zone for about 70 years. Only theposition of the Aplococeras avisianum - Zonewas disputed. Many geologists placed the An-isian-Ladinian boundary between the A. avisia-num- and the R. reitzi - Zone. Only KOZUR(1972 and later papers) stated that the Avisianum- Zone has the same Lower Ladinian microfaunaas the Reitzi - Zone. KOZUR & MOSTLER (inpress since 1991) could now prove by detailedradiolarian correlations that the Avisianum- Zo-ne corresponds to the upper Reitzi - Zone andsomewhat younger beds. The hardly under-standable position of the Avisianum - Zone abo-ve the Curionii - Zone in MOJSISOVICS; WA-AGEN & DIENER (1895) was by this resulteasily explainable. The Avisianum - Zone ischaracteristic for rather shallow water sedimentsnear the platform slope, the Reitzi - Zone for pe-lagic beds. If pelagic beds of the Reitzi - Zones.str. (= lower Curionii - Zone sensu MOJSI-SOVICS, WAAGEN & DIENER, 1895) havebeen overlain by a carbonate platform then inthe transitional faciès Aplococeras avisianumcan be found above the Reitzi - Zone s.l. (= Cu-rionii - Zone s.l.). Because the carbonate plat-form is free of ammonites, ammonoid faunas ofthe Curionii - Zone s.l. are overlain by ammo-noid faunas of the Avisianum - Zone.

Independent from the position of the Avisia-num - Zone, the base of the Reitzi - Zone was

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invariably used as base of the Ladinian for abouta century, one of the clearest priorities in the Te-thyan Triassic stratigraphy. This priority is notinvalid by the fact that in the Southern Alps oth-er ammonoids have been misinterpreted as 'Ce-ratites' reitzi. The Reitzi - Zone was from thebeginning defined in the Balaton Highland, itstype locality is the Forrâshegy section at Fel-söörs, where all 4 subzones of the Reitzi - Zoneare present and also the underlying Illyrian bedsare rich in ammonoid faunas. This section is ex-cellently exposed and was studied in detail bymany Hungarian and foreign geologists. Thescope of a zone does not change, if somebodymakes a misidentification of the index speciesoutside the type locality of the zone.

PIA (1930) showed that in the Reitzi - ZoneDiplopora annulata appears for the first time, adasycladacean guideform for the whole Ladin-ian in the widespread Tethyan carbonate plat-forms.

TOZER (1967) placed the base of the Ladin-ian considerably higher, at the base of the Eo-protrachyceras subasperum - Zone which corre-sponds to the base of the E. curionii - Zone inthe European Tethys. Geologists, who worked inthe Arctics and the RIEBER school from Zürichfollowed him. This boundary is a good ammo-noid boundary, but it lies considerably above thepriority boundary and it is not correlable by anypronounced boundary in conodont and radiolar-ian faunas and neither in the dasycladacean as-sociation (inside the Diplopora annulata flora)nor in sporomorph zonations. Therefore thisboundary is hardly correlable in ammonoid-freeTriassic successions that compris by far morethan 90 % of the known Triassic in the world.Moreover, by this high boundary the Fassanianwould be reduced to a very short time intervalbetween the base of the Curionii - Zone and thebase of the Gredleri - Zone.

A third Anisian-Ladinian boundary was pro-posed by KRYSTYN (1983) in the Epidaurossection. According to his studies the first ap-pearance of Nevadites coincides with a sharpboundary in the conodont fauna, indicated bythe first appearance of P. trammeri. This boun-

dary was overtaken by the Milano school (GAE-TANI, NICORA) and for a time also byKOVACS who meanwhile again supports thepriority boundary at the base of the Reitzi - Zo-ne. In an ad hoc voting on the Anisian/Ladinianboundary field workshop (27. 6. - 4. 7. 93, appli-cation made by TOZER), 4 participants votedfor the base of the Curionii - Zone, 4 partici-pants for the base of the Nevadites fauna and 7for the base of the Reitzi - Zone as Anisian-La-dinian boundary.

There was general agreement that the base ofthe Curionii - Zone is not supported by any dis-tinct change in microfauna. However, accordingto our opinion, there are not yet enough conod-ont data from this boundary. According to thedata of the Italian colleagues, there are no chan-ges in the conodont faunas near the base of theCurionii -Zone. However, in Hungary Buduro-vignathus truempyi occurs in the upper part ofthe Curionii - Zone, but the ammonoid correla-tions in this section (Köveskal) are rather weak.

There was no agreement about the assumedsharp conodont boundary at the base of the Nev-adites fauna. Wheareas KRYSTYN and NICO-RA continue to believe in this boundary, KO-ZUR has shown that P. trammeri appears con-siderably earlier in agreement with the data pub-lished by KOZUR & MOSTLER (1982). A dis-tinct conodont boundary can be, in turn, ob-served at the base of the Reitzi - Zone, whereone of the two major radiolarian boundaries inthe Late Illyrian to Longobardian interval oc-curs.

How may these different opinions be ex-plained ? KRYSTYN (1983) found his conodontboundary (first appearance of P. trammeri) at thebase of the Nevadites fauna in the Epidaurossection. According to his opinion, this section iscondensed in the Anisian part, but uncondensedin the Ladinian part (including the Nevaditesfauna). This view could not be confirmed byKOZUR & MOCK (in prep.) who re-sampledthis section. Also the Ladinian part is stronglycondensed. The bed with the first Nevadites oc-curs above a several cm thick manganese oxidelayer indicating strong condension. Also the co-

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nodont fauna of the level with Nevadites isstrongly condensed. Thus the very strange co-nodont ranges shown by KRYSTYN (1983)from this section (e.g. beginning of P. trammeriat the base of the Nevadites fauna, beginning ofthe Longobardian B. hungaricus at the base ofthe Curionii - Zone) can be explained. In un-condensed sections P. trammeri begins by farearlier. This was unintentionally confirmed byKRYSTYN (1983) and KOVÂCS et al. (1990)that means by all conodont workers that believein a strong conodont boundary at the base of theNevadites - Zone.

KOZUR & MOSTLER (1982) recorded a co-nodont fauna with N. longa and P. trammeripraetrammeri in ammonoid dated beds belong-ing to the Kellnerites fauna (R. reitzi - Zone). Tothe paper of KOZUR & MOSTLER (1982)KRYSTYN (1983, p. 257) remarked the follow-ing: 'In obiger Arbeit wird aus der angeblichdurch Parakellnerites abgesicherten Probe FQeine Conodontenfauna genannt, die verglichenmit Epidaurus und anderen Profilen ganz sicherfür die Nevadites - Zone und keineswegs für dieParakellnerites - Zone charakteristisch ist. Indiesem Lichte sind die stratigraphischen Schluß-folgerungen der Autoren hinsichtlich der Einstu-fung ihrer Probe und der enthaltenen Plattform-Conodonten ('typisch unterladinisch') bemer-kenswert, weil sie sich insbesondere auf Gondo-lella trammeri und Gondolella pseudolongastützen. Die neubeschriebene Gondolella tram-meri praetrammeri hat hier zur Umbenennungvon Gondolella praetrammeri in Gondolella eo-trammeri n. sp. genötigt. Überflüssigerweise lei-der, weil G. trammeri praetrammeri KOZUR &MOSTLER eindeutig in die Synonymie von G.trammeri zu verweisen ist, während sie sich an-dererseits von G. eotrammeri n. sp. artlich sicherunterscheidet.' From this Statement the follow-ing facts can be taken: 1. KRYSTYN (1983) ag-rees that this conodont fauna is a typical LowerLadinian conodont fauna. He was so sure aboutthe Lower Ladinian character that he assignedthe ammonoid fauna without investigation intothe Nevadites fauna. 2. KRYSTYN agrees thatthe form described by KOZUR & MOSTLER

(1983) is a P. trammeri. According to a pers.comm., Dr. KRYSTYN has meanwhile restud-ied the ammonoid fauna and could confirm thedata of KOZUR & MOSTLER (1982) that thisfauna belongs to the Kellnerites fauna (R. reitzi -Zone). Therefore he has unintentionally con-firmed that the R. reitzi - Zone contains a typicalLower Ladinian fauna with P. trammeri. He nowagrees that this form is a little more primitiveand can be separated in subspecies level (see re-marks in the systematic part). KO VACS et al.(1990) also confirmed that P. trammeri prae-trammeri is a 'typical' P. trammeri and also theyregarded therefore the horizon from where thesample has been derived as Early Ladinian inage. Therefore all conodont workers that regardthe first appearance of P. trammeri as an impor-tant marker for defining the Anisian-Ladinianboundary have unintentionally confirmed thatthe R. reitzi - Zone has an undoubtedly LowerLadinian conodont fauna.

Medium advanced P. trammeri that are hardlyseparable from the primitive P. trammeri prae-trammeri occur in the Felsöörs section in the up-per Subzone of the R. reitzi - Zone (see alsoKOVÄCS in GAETANI, 1993). In the samesection P. trammeri praetrammeri is present inthe Liepoldti - Subzone, the second lowest Sub-zone of the R. reitzi - Zone. KO VACS wrote:'Appearance of G trammeri evolved from G.aff. eotrammeri near to the base (?) of the Costo-sus horizon. This is the most remarkable andeasiest recognizable one among the four eventsdiscussed here, being G. trammeri a very com-mon and characteristic form in the Tethyan eu-pelagic faciès' We agree that the first appear-ance of P. trammeri is an important conodontevent, but not the appearance of medium ad-vanced forms at the base of the upper subzone ofthe Reitzi - Zone, but the first appearance of thespecies itself (first appearance of P. trammeripraetrammeri). It is very difficult to separate themedium advanced P. trammeri of the upper Re-itzi - Zone and of the Nevadites fauna from themore primitive P. trammeri of the lower Reitzi -Zone as well demonstrated by KRYSTYN(1983) and KOVÀCS et al. (1990) by regarding

188

the primitive P. trammeri praetrammeri as typi-cal P. trammeri. P. trammeri praetrammeri wasestablished to separate it from the advanced P.trammeri of the uppermost Curionii - Zone andof the Langobardian. Even this separation is nottoo easy. The medium advanced forms from theupper Reitzi - Zone and Nevadites fauna aretransitional between these two subspecies! Ofcourse, the very difficult separation of theseforms from P. trammeri praetrammeri (KRYS-TYN, 1983 and KOVÀCS et al., 1990 failed inseparating P. trammeri praetrammeri of the Re-itzi - Zone from typical P. trammeri) cannot beused as an important conodont event, only thefirst appearance of P. trammeri themselves andthis event is in the lower Reitzi - Zone ! At thebase of the Reitzi - Zone N. mesotriassica ap-pears, a typical Ladinian form. N. cornuta ladin-ica n. subsp., N. balkanica s.str., N. longa, N.basisymmetrica and N. aldae first appear at ornear the base of the Reitzi - Zone. All theseforms are typical Lower Ladinian forms, oftendifficult to separate from each other, especiallyin juvenile and medium ontogenetic stages. Thecommon feature of all these forms is that theyhave rather long and slender juvenile forms ofconstricta type, in which already in medium on-togenetic or late juvenile stages the basal cavityis somewhat fore ward-shifted, a typical Ladin-ian (and younger feature), unknown in any Illyr-ian platform conodont.

Also the Paragondolella associations, charac-teristic for deep pelagic associations, changedistinctly at or near the base of the Reitzi - Zo-ne. At the base of the Reitzi - Zone appears P.alpina (and its junior synonym P. szaboi, thejuvenile forms of P. alpina). P. alpina belongs tothe P. trammeri group that dominates among thegondolellids of the deep pelagic Ladinian co-nodont faunas. P. trammeri itself first appears inthe lower Reitzi - Zone with the primitive P.trammeri praetrammeri that is almost insepara-ble from medium advanced forms that begin inthe upper Reitzi - Zone. Also among the P. ex-celsa group distinct changes can be observed,but the investigation of the range of the differenttaxa of the P. excelsa group is not yet finished.

Typical P. excelsa with very high anterior plat-form and broad platform brim behind the lastdenticles appear seemingly near the base of theReitzi - Zone.

The next distinct conodont change is onlywithin the Curionii - or Recubariense - Zone,where in a still unknown exact level the genusBudurovignathus appears. In the interval bet-ween these two conodont events only slightmodifications within the Lower Ladinian conod-ont fauna can be observed.

The distinct conodont boundary at the base ofthe Reitzi - Zone is not faciès controlled. Transi-tional forms to Illyrian forerunners can be foundin the underlying Asseretoceras- and Lardaroc-eras faunas.

The distinct change in the conodont faunas atand near the base of the Reitzi - Zone is accom-panied by a drastic change in the dominantradiolarian groups. The first typical Oertlispon-gidae with curved main spine (genus Oertlis-pongus) appeared at the base of the Reitzi - Zone(KOZUR & MOSTLER, in press, DOSZTÂLY,in press). In the same level Triassocampe dewe-veri and T. scalaris and some near related formsthat dominate the Ladinian Nasselarian faunasappear. The common and characteristic Ladin-ian Eptingium manfredi and the genus Yeharaiaappear also in this level. Further common andimportant Ladinian guideforms appear in the sa-me level. Therefore the radiolarian faunas of theupper Paraceratites trinodosus - Zone and thebasal Reitzi - Zone are definitely different intheir most important components. Within theFassanian, a gradual development of the Ladin-ian faunas can be observed without changing thegeneral Lower Ladinian character of the faunas.Only at the base of the Budurovignathus mun-goensis - Zone the next drastic change in themain components of the radiolarian faunas canbe observed. Also the radiolarian faunas supportthe placement of the Anisian-Ladinian boundaryat the base of the Reitzi - Zone.

Near the base of the Reitzi - Zone the Ladin-ian phase of the sporomorph development sensuBRUGMAN starts. According to GOCZÀN (inGAETANI (1993) the decisive important forms

189

for the Ladinian phase, Cannanoropollis scheu-ringi, C. brugmani, Kuglerina meieri in the Fel-söörs section first appear in beds 98, immediate-ly below the base of the Reitzi - Zone within theLardaroceras ammonoid fauna. In the same lev-el also a distinct change in the foraminifer faunacan be observed (ORAVECZ-SCHEFFER inGAETANI, 1993). As known since long time,also the Ladinian dasycladacean flora with Dip-lopora annulata appears near the base of the Re-itzi -Zone.

The priority Anisian-Ladinian boundary istherefore supported by all important microfossilgroups that allow correlations within the pelagicammonoid-bearing faciès, within the deep pe-lagic ammonoid-free faciès (radiolarites), withinthe ammonoid-free shallow-water faciès and inthe continental Triassic (sporomorphs). None ofthe other proposed ammonoid boundaries havethis advantage and moreover, they are not inagreement with the priority. For this reason, weplace the base of the Ladinian in agreement withthe more than 100 year old priority and use atthe base of the Reitziites reitzi - Zone.

Accepting this boundary, all the investigatedconodont faunas of the Southern KarawankenMountains with exception of the samples WWSand ZG 1, ZG 2 and ZG 3 (see below) belong tothe Lower Ladinian. This is indicated by thepresence of Paragondolella trammeri, P. alpina,advanced P. excelsa, Neogondolella cornuta la-dinica, N. mesotriassica, N. longa, N. balkanica.In agreement with the above statements, withinthe Lower Ladinian only in few samples a spec-ification is possible by conodonts. Sample ZE6a contains already very primitive Budurovigna-thus and should be therefore Late Fassanian inage.

The samples ZG 1 to ZG 3 yielded amongother forms Budurovignathus mungoensis. Theybelong therefore to the B. mungoensis A.Z. ofthe Langobardian.

Sample WWS from a fissure filling of red pe-lagic limestones (Weiße Wand Member of theLoibl Formation) in the uppermost part o theunderlying platform carbonates (Contrin Forma-tion) contained Nicoraella kockeli (TATGE).

The disappearance of N. kockeli only in theNorthern Tethys is a good time-marker for thetop of the Pelsonian. In the Southern Tethys thisspecies ranges up into younger beds and still inthe Cordevolian the very similar N. postkockelioccurs (KOZUR, 1993). Also the other conod-onts are not very diagnostic. An early juvenileParagondolella (PI. 1, Fig. 6) cannot be deter-mined, because in these early juvenile stages thegondolellid conodonts are not well distinguish-able. It could be an early juvenile P. trammeri,but also an early juvenile P. eotrammeri, the ageof which is not clear because of the strong con-dension in the Epidauros section. P. ? pridaensisposteroacuta is probably an advanced P. pri-daensis, because this subspecies is also presentin the Ladinian of Nevada. If this is true, the fau-na of sample WWS must be younger than theParaceratites cricki beds and (because of theradiolarian fauna) older than the base of the Re-itzi - Zone. This would indicate a latest Illyrianage. However, because the forerunner of P. ?pridaensis is unknown, it cannot be totally ex-cluded that P. ? pridaensis posteroacuta is aprimitive P. pridaensis. Typical Paragondolellan. sp. ex gr. excelsa that are dominant in sampleWWS are known from the Lower Ladinian ofJapan, but the whole range of the species is un-known. Thus, for the moment a latest Illyrianage is most probably for this sample that has anrather unusual species composition, perhaps pre-dominantly forms that lived in a fissure.

6. Conclusion

The conodont fauna derived from pelagiclimestones of the Loibl Formation, containingParagondolella trammeri, P. alpina, advancedP. excelsa, Neogondolella cornuta ladinica, N.mesotriassica, N. longa, N. balkanica and al-ready primitive Budurovignathus, points to Ear-ly to Late Fassanian age.

Early Fassanain age is also indicated by theradiolarian fauna from one locality (WeißeWand Member of the Loibl Formation at the

190

Weiße Wand; KRAINER & MOSTLER 1992,KRAINER & MOSTLER in prep.) and by am-monites from red limestones exposed along theZelenica forest road (BAUER 1980).

Red fissure fillings in the uppermost ContrinFormation at the Weiße Wand containing Nico-rella kockeli may be of latest Illyrian age.

Late Illyrian age is also reported from red pe-lagic limestones from the peak of the Zeller Pra-potnik mountain by BAUER (1984) due to theoccurrence of Kellnerites sp.

The conodont fauna derived from the Bu-chenstein Formation is of Fassanian (sectionZimpasser Gupf west) and Late Langobardianage (section Zimpasser Gupf north), the latterindicated by Budurovignathus mungoensis.

This is in good conformity with the radiolar-ian fauna obtained from samples of the Buchen-stein Formation from other localities in the Ka-rawanken Mountains which also points to an ageranging from the Late Fassanian to the LateLangobardian.

This indicates that the volcanic rocks and clas-tic sediments between the Loibl Formation andthe Buchenstein Formation were formed during avery short time span within the Fassanian.

Acknowledgements

We are very grateful to the Jubiläumsfonds ofthe Oesterreichische Nationalbank (Project no.3935) for the financial support of this study.

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Anschrift der Verfasser:Dr. se. Heinz Kozur, Rézsii u. 83, H-1029 Budapest, Hungary;Univ.-Doz. Dr. Karl Krainer, Univ.-Prof. Dr. Helfried Mostler,Institut für Geologie und Paläontologie der Universität Inns-bruck, Innrain 52, A-6020 Innsbruck, Austria.

submitted: January 21, 1994 accepted: February 5, 1994

193

Explanation of Plates

Plate 1All figured specimens, except Fig. 22, have been derived from sample WWS, a fissure filling of redpelagic limestones (Weiße Wand Member of the Loibl Formation) in the underlying platform carbo-nates (Contrin Formation) of the section Weiße Wand, W Loiblpass. Uppermost Illyrian (?), accord-ing to the radiolarian fauna somewhat older than the base of the R. reitzi - Zone.

Figs. 1-5, 11-19: Paragondolella n. sp. ex gr. excelsa MOSHER, Figs. 1-3: lateral, oblique lower andupper views of one specimen, Figs. 11, 14, 17: lateral, upper and lower views of a specimenwith partly damaged platform, Figs. 12, 15, 18: lateral, upper and lower views of a juvenilespecimen, Figs. 13, 16, 19: lateral, upper and lower views of a specimen with partly damagedplatform and carina; rep.- no. KKM 1993 III-2, Figs. 1, 3-5: x 130, Fig. 2: x 120, Figs. 11-19:xlOO.

Figs. 6, 9: Early juvenile Paragondolella probably of the P. trammeri group, Fig. 6, lateral view, Fig.9: somewhat oblique lower view; x 160, rep.-no. KKM 1993 III-3.

Figs. 7, 8, 10, 20, 21, 24: Paragondolella ? pridaensis posteroacuta n. subsp., Figs. 7, 8, 10: lateral,upper and lower views of holotype, x 110, rep.-no. KKM 1993 III-4; Figs. 20, 21, 24: lateral,lower and oblique upper view of a broken specimen (anterior part missing), x 130, rep.-no.KKM 1993 m-5.

Fig. 23: Nicoraella kockeli (TATGE), lateral view, x 160, rep.-no. KKM 1993 III-6.Fig. 22: Neogondolella balkanica BUDUROV & STEFANOV, lateral view of a broken specimen

(anterior part missing). Cusp terminal, but not fused with the platform margin, sample ZE 6 a,red pelagic limestones (Loibl Formation) at the Zeller Prapotnik (E of Loiblpass), upper Fas-sanian, x 80, rep.-no. KKM 1993 III-7.

Plate 2All figured specimens are from sample PR 1, outcrops at the Zeller Prapotnik E of Loiblpass (LoiblFormation). The fauna displays a distinct Fassanian character, but their position inside the Fassaniancannot be determined (surely below the upper Fassanian Budurovignathus truempyi - Zone). With ex-ception of figures 1, 2,4, 5 and 7, 8 all specimens are broken (mostly crashed during the sticking), butthe posterior half is well suitable for determination of Ladinian Neogondolella, at least for recogni-tion, whether Illyrian or Fassanian forms are present.

Figs. 1, 4, 7: Lateral, upper and lower views of a juvenile form (constricta stage) of a slender, longNeogondolella, x 120, rep.-no. KKM 1993 III-8. An exact determination of the species is im-possible in such juvenile forms. However, such long, slender, little arched juvenile forms donot occur before the base of the Reitzi - Zone. They are world-wide distributed and in deeppelagic rocks (radiolarites) beside juvenile Paragondolella often the only conodonts, becausein such faciès adult gondolellid conodonts are rare. In restricted pelagic environments near theécologie tolerance boundary for platform conodonts juvenile Neogondolella are the only plat-form conodonts. If the base of the Ladinian is placed at the base of the R. reitzi - Zone, thenthe Ansisian-Ladinian boundary can be well determined by the first appearance of such long,slender juvenile Neogondolella in conodont faunas from all these environments, that are most-ly free of ammonoids. If we use the base of the Nevadites fauna or the base of the E. curionii -Zone as base of Ladinian, then all these faunas can be only assigned as Illyrian to Fassanian

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faunas, because these long, slender juvenile Neogondolella dominate all neogondolellid fau-nas from the base of the Reitzi - Zone up to the top of the Fassanian.

Figs. 2, 5, 8: Paragondolella cf. alpina (KOZUR & MOSTLER), juvenile specimen with unusuallylittle arching of the unit, x 140, rep.-no. KKM 1993 III-9.

Figs. 3, 6, 9: Neogondolella balkanica BUDUROV & STEFANOV, typical specimen with platformbrim and prominent, broad, conical cusp, x 100, rep.-no. KKM 1993 III-10.

Figs. 10, 11, 13, 14, 16, 17: Juvenile stages (constricta stage) of Neogondolella sp., Fig. 10, 13, 16: x120, rep.-no. KKM 1993 ffl-11, Fig. 11, 14, 17: x 150, rep.-no. KKM 1993 111-12. Brokenjuvenile stages of Illyrian and Fassanian Neogondolella are in general indeterminable in spe-cies level.

Figs. 12, 15, 20: Neogondolella mesotriassica (KOZUR & MOSTLER), transitional form to N cor-nuta ladinica n. subsp., x 100, rep.-no. KKM 1993 HI-13.

Figs. 18, 21, 22: Neogondolella longa BUDUROV & STEFANOV, lateral, upper and lower view ofa late juvenile modified constricta stage ('pseudolonga' stage), x 130, rep.-no KKM 1993 III-14.

Figs. 19, 23, 24: Paragondolella trammeri (KOZUR), upper, lateral and oblique lower view of ajuvenile specimen, x 150, rep.-no. KKM 1993 III-15.

Plate 3The specimens on Figs. 1-18, 19, 22, 25 have been derived from sample ZE 6 a (outcrops along theZelenitza forest road, W of Loiblpass). Because of the occurrence of primitive Budurovignathus, thissample can be placed into the upper Fassanian. The specimens on Figs 20, 23, 26-29 have been de-rived from sample PR 1 (see PI. 2). The specimen on figs. 21, 24 has been derived from sample WW3 (Weiße Wand section W Loiblpass). The fauna of this level is extremely poor in conodonts, but be-longs probably to the Lower Ladinian.

Figs. 1, 4, 7, 19, 22, 25: Budurovignathus gabriellae n. sp., x 120, Figs. 1,4, 7: upper, lateral andlower view of holotype, rep.-no. KKM 1993 III-1, Figs. 19, 22, 25: upper, lateral and lowerviews of a paratype, rep.-no. KKM 1993 EH-16.

Figs. 2, 5, 8, 11, 14, 17: Neogondolella cornuta ladinica n. subsp., Figs. 2, 5, 8: upper, lateral andlower views of holotype, x 100, rep.-no. KKM 1993 111-17, Figs. 11, 14, 17: upper, lateral andlower views of a broken specimen, consisting of the posterior half of the unit, x 86, rep.-no.KKM 1993 ffl-18.

Figs. 3, 6, 9, 27-29: Neogondolella balkanica BUDUROV & STEFANOV, Figs. 3, 6, 9: upper, later-"al and lower views of a broken specimen consisting of the posterior half of the unit, x 110,rep.-no. KKM 1993 III-19, Figs. 27-29: lateral, upper and oblique lower views of an unusualshort specimen with very long main cusp, representing perhaps a new subspecies, x 80, rep.-no. KKM 1993 m-20.

Figs. 10, 13, 16: Paragondolella trammeri (KOZUR), upper, lateral and oblique lower views of ajuvenile specimen, x 150, rep.-no. KKM 1993 111-21.

Figs. 12, 15, 18: Gladigondolella cf. malayensis NOG AMI, morphologically transitional to G. tethy-dis (Huckriede), upper and lateral views, x 80, rep.-no. KKM 1993 111-22.

Fig. 20: Neogondolella sp., early juvenile stage {constricta stage), upper view, x 140, rep.-no. KKM1993 m-23.

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Figs. 21, 24: Eccentric Neogondolella sp. aff. N. transita (KOZUR & MOSTLER), upper and lowerview of a broken specimen, x 100, rep.-no. KKM 1993 111-24.

Figs. 23, 26: Neogondolella sp., slender long juvenile form of constricta stage, transitional to mom-bergensis stage, rep.-no. KKM 1993 111-25, fig. 23: upper view, x 110, Fig. 26: lateral view, x115.

Plate 4

Figs. 1, 2: Budurovignathus mungoensis (DIEBEL), upper view, sample ZG 2, Langobardian, B.mungoensis A.-Z., section NW Zimpaserkogel (SW Eisenkappel), x 100, rep.-no.15-3/28/5/1993.

Figs. 3-5: Neogondolella longa BUDUROV & STEFANOV, adult specimen, sample Z 14, section WZimpaserkogel (SW Eisenkappel), Fassanian, rep.-no. 6-13/27/5/93, Fig. 3: lower view, x 100,Fig. 4, upper view, x 100, Fig. 5: lateral view, x 80.

Fig. 6: Paragondolella excelsa excelsa MOSHER, lateral view, x 100, sample Z 5, section W Zimpa-serkogel (SW Eisenkappel), Fassanian, rep.-no. 1-5/27/5/93.

Figs. 7-9: Mesogondolella mesotriassica (KOZUR & MOSTLER), oblique lateral, upper and lowerviews of an broken adult specimen (posterior part preserved), posterior platform ridge partlywith small teeth, x 100, sample PR 1 (see PI. 2), rep.-no 7-13/27/5/93.

Figs. 10, 11: Paragondolella inclinata (KOVACS), upper and lateral views of a juvenile specimen,sample ZG 2 (see Figs. 3-5), x 100, rep.-no. 16-3/27/5/93.

Figs. 12, 13, 15: Neogondolella constricta (MOSHER & CLARK), upper, lower and lateral views ofa medium ontogenetic stage, carina of constricta type, x 100, sample SD 1/1, western flank ofGosing Mt., near Siedig Eastern Alps (Austria), Illyrian N. constricta Zone, rep.-no7-3/27/5/93.

Figs. 14, 17, 20: Neogondolella constricta (MOSHER & CLARK), upper, lower and lateral view,main cusp indistinct, basal cavity almost terminal with respect to the keel end, x 100, sampleSD 1/1 (see above), rep.-no. 4-3/27/5/93.

Figs. 16, 18, 19, 21: Neogondolella cornuta BUDUROV & STEFANOV, lateral view of different on-togenetic stages, only the medium ontogenetic stages and adults can be subdivided in subspe-cies level, x 80, sample P 7, Karwendel, condensed upper Illyrian and lower Fassanian, Fig.16, juvenile stage {constricta stage), inseparable from the most other Illyrian and FassanianNeogondolella species, rep.-no. 21-6/27/5/93, Fig. 18: late juvenile stage of constricta type,transitional to mombergensis type (penultimate tooth is the cusp, but not much larger than thedenticles in front of it), rep.-no. 20-6/27/5/93, Fig. 19: medium ontogenetic stage, momber-gensis type without cusp, rep.-no. 18-6/27/5/93, Fig. 21: adult specimen of a primitive N. cor-nuta ladinica with distinct terminal cusp, fused with the posterior platform margin, its anteriormargin is posteriorly inclined, its posterior margin is erect, rep.-no. 12-6/27/5/93.

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Plate 1

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Plate 2

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Plate 3