Geologia Croatica 60/1 1–113 18 Figs. 26 Pls. ZAGREB 2007

Stratigraphy and Palaeobiogeography of Mesozoic Benthic of the Karst Dinarides (SE Europe)

Ivo VELIĆ

Key words: Benthic foraminifera, Chronostratigraphy, Within the 257 determined taxa belonging to different foramini­ feral families which lived through the Mesozoic, numerous differ­ Biostratigraphy, Palaeobiogeography, Mesozoic, ent index fossils occur in assemblages indicating various ages: Early Karst Dinarides, Adriatic Carbonate Platform, Slove­ , Anisian, Carnian, Norian–Rhaetian, Late Sinemurian, Early nia, Croatia, Bosnia and Herzegovina, Montenegro. and Late Pliensbachian (Carixian and Domerian), Early and Late Aalenian, Early and Late Bajocian, Early and Late Bathonian, Callo­ vian, Early and Late Oxfordian, Kimmeridgian, Tithonian, Berriasian, Valanginian, Late Hauterivian, Late Barremian, Early and Late Aptian (Bedulian and Gargasian), Early and Late Albian, Early, Middle and Abstract Late Cenomanian, Turonian, Coniacian, Santonian, Early and Late The Adriatic Carbonate Platform (AdCP), was a separate shallow- Campanian and Early and Late Maastrichtian. marine depositional system characterized by a lack of terrigenous A total of 64 biostratigraphic units – biozones of different catego­ input and was connected to Gondwana towards the South via Gav­ ries, from subzone to superzone, were defined within the stratigraphic rovo–Tripolitza or Apulia. It existed for approximately 120 MY, from interval from the Carnian to the Late Maastrichtian. This enabled very the Early (Pliensbachian/) to the end of the Creta­ detailed biostratigraphic subdivision of the carbonate deposits within ceous, resulting in a 4000–6500 m thick succession of almost pure the Karst Dinarides. This is one of the most precise sequences, not carbonates. However, this is part of a thicker (>8000 m) sequence of only in this area, but also among former shallow marine deposits of predominantly carbonate rocks which forms the Karst Dinarides, and the entire Neotethyan realm in the present Mediterranean region. was deposited during more than 270 MY – at least from the Carbonif­ The palaeobiogeographic characteristics of biotopes and the erous (Moscovian) to the Late Eocene. composition of foraminiferal assemblages during the Mesozoic were Among many different groups of fossil organisms, benthic controlled by the position of the study area within the Neotethyan bio­ foraminifera are especially abundant and well preserved, so they, provinces. Until the Albian, this area represented part of the Southern along with calcareous algae (Dasycladales), are the most important Neotethyan bioprovince, while from the Cenomanian to its final disin­ fossils used for age determination and stratigraphic subdivision of tegration at the end of the it belonged to a separate, Central shallow-marine carbonate deposits. Mediterranean Neotethyan bioprovince.

Ovaj rad posvećujem članovima Velebitsko–kapelske ekipe This paper is dedicated to the members of the Velebit–Kapela (1962–1980) – kolegama Anti Ivanoviću, Leonu Nikleru, team (1962–1980) – my colleagues Ante Ivanović, Leon Branku Sokaču, Ivanu Galoviću i Stjepanu Markoviću s Nikler, Branko Sokač, Ivan Galović and Stjepan Marković. kojima sam zajedno sudjelovao u izradbi Osnovne geološke Together we produced a Basic Geological Map at 1:100,000 karte 1:100.000 u Velebitu, Lici, Velikoj i Maloj Kapeli scale, and performed detailed stratigraphical subdivision of i Ogulinskome kraju. Tada su se, prvenstveno prema the Jurassic and Cretaceous rocks of Velebit Mt., Lika, Velika mikrofosilima, a među njima i bentičkim foraminiferama, and Mala Kapela Mts. and the Ogulin area. Based on benthic provele prve detaljnije stratigrafske raščlambe jurskih i microfossils – foraminifera and dasyclad algae, this was the krednih naslaga u Krškim Dinaridima. To je bilo polazište first such subdivision in the Dinarides. That work represented za kasnija paleontološko–stratigrafska istraživanja koja su a foundation for subsequent palaeontological–stratigraphic rezultirala mnogobrojnim znanstvenim radovima, pa tako i investigations which resulted in numerous scientific papers, ovim radom. of which the present paper is included.

Croatian Geological Survey, Sachsova 2, HR-10000 Zagreb, Croatia; e-mail: [email protected]  Geologia Croatica 60/1

CONTENTS

1. Introduction...... 2 2. Foraminiferal taxa and assemblages in chronostratigraphic succession...... 3 2.1. Triassic...... 4 2.1.1. Early Triassic – Scythian (Induan and Olenekian)...... 5 2.1.2. Middle Triassic...... 5 2.1.3. Late Triassic...... 5 2.2. Jurassic...... 6 2.2.1. Early Jurassic...... 6 2.2.2. Middle Jurassic...... 9 2.2.3. Late Jurassic...... 11 2.3. Cretaceous...... 12 2.3.1. Early Cretaceous...... 12 2.3.2. Late Cretaceous...... 16 3. Biostratigraphy...... 20 3.1. Previous investigations...... 20 3.2. Biostratigraphic zonation and subdivision...... 21 3.2.1. Late Triassic...... 21 3.2.2. Early Jurassic...... 22 3.2.3. Middle Jurassic...... 24 3.2.4. Late Jurassic...... 27 3.2.5. Early Cretaceous...... 29 3.2.6. Late Cretaceous...... 33 4. Palaeobiogeography of Mesozoic benthic foraminifera of the Karst Dinarides...... 38 4.1. Triassic...... 38 4.2. Jurassic...... 38 4.3. Cretaceous...... 42 4.4. Correlation...... 46 5. Review of the results...... 48 6. References...... 51

Appendix: List of Taxa...... 59

1. INTRODUCTION stratigraphically well determined forms; especially the Orbitopsella spp. of the Lower Jurassic of Morocco Benthic foraminifera are among the most important fos­ (SEPTFONTAINE, 1984), hauranids and pfenderinids sil organisms for age determination and stratigraphic in the Middle Jurassic and kurnubiinids in the Upper Jurassic of the Western Mediterranean in Europe and subdivision of shallow-marine carbonate deposits of the Northern Africa (BASSOULLET, 1997), orbitolinids Mesozoic Karst Dinarides. Some taxa (or even entire in the Hauterivian–Aptian (‘Urgonian’) deposits of SE assemblages) enable chronostratigraphic determination, France (e.g. CLAVEL et al., 1995; ARNAUD et al., because their stratigraphic ranges were evaluated by 1998; SCHROEDER et al., 2002), or within Upper Bar­ correlation with ammonite zones, mostly in the Western remian–Middle Cenomanian stratigraphic range in the Mediterranean realm, e.g. Moroccan Atlas, Spanish Bet­ Prebetian Platform of SE Spain (CASTRO et al., 2001). ic Cordillera, and especially in France: Pyrenees, Aqui­ Other examples include the orbitolinids and alveolinids taine and Paris Basins, Provence, Western Alps, etc. The of Vraconian and Cenomanian deposits of the Eastern Jurassic lituolids can be highlighted, as an example of Pyrenees (BILOTTE, 1985), etc. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 

During the last couple of decades, new and sophis­ Bosnian to the N and NE, Krasta–Budva–Cukali to the ticated methods have been introduced into stratigraphic SE, Ionian–Adriatic–Belluno to the S and SW and the and palaeontologic investigations, e.g. dating on the Slovenian Basin to the NW. basis of isotopic or palaeomagnetic analyses, or deter­ During the late Maastrichtian and Palaeogene, the mination of DNA within fossil organisms, and many platform significantly disintegrated, and carbonate papers were published. Recently, when the term pal­ depositional environments were established only within aeontology has become increasingly substituted by the restricted areas, particularly several narrow carbonate term palaeobiology, the purpose of a paper such as this, ramps along flysch–molasse basins surrounding uplifted i.e. the stratigraphy of benthic foraminifera within the areas of the former AdCP (VLAHOVIĆ et al., 2002). shallow-marine Mesozoic carbonates of the Adriatic Car­ Despite significant tectonic disintegration, which bonate Platform, may be questioned. This is especially had already begun in the Mesozoic and lasted practi­ true since papers on similar topics in the neighbouring cally throughout the Cenozoic, the Adriatic Carbonate carbonate platforms were published more than 10 years Platform is one of the best preserved fossil carbonate ago (e.g. CHIOCCHINI et al., 1994: Apenninic Plat­ platforms in the World (except for its marginal parts). form). However, this is the first synthesis of the results of stratigraphic investigation of benthic foraminifera Unlike the aforementioned areas of the Western of the AdCP in the last fifty years, especially on Upper Mediterranean, which were characterized by interfinger­ Triassic to Uppermost Cretaceous deposits. ing of shallow-marine and open-marine/basinal facies, allowing chronostratigraphic determination of many The Adriatic Carbonate Platform (AdCP, Fig. 1) is a foraminiferal taxa, no such possibilities occurred on term for a palaeogeographic entity which existed from the AdCP. This platform was geotectonically relative­ the Early Jurassic to the end of the Cretaceous, and ly stable throughout its existence, without significant which forms an important part of the Karst Dinarides. palaeogeographic and palaeoenvironmental changes. However, shallow-marine environments mostly charac­ Therefore, intra-platform shallow-marine deposits were terised by carbonate deposition existed for much longer only sporadically accompanied by temporary drowned- in the study area. The oldest carbonate rocks cropping platform environments containing ammonites or other out in the Karst Dinarides are in age, pelagic organisms enabling stratigraphic correlation. while almost continuous carbonate deposition lasted for However, platform margin and slope facies, which inter­ some 220 MY, from the Middle to the Eocene fingered with basinal deposits, were overlain by young­ (VELIĆ et al., 2002). er, Tertiary and Quaternary clastic rocks, or by nap­ Before the individualization, i.e. during the Late pes composed of Palaeozoic and older Triassic rocks. Palaeozoic and Early Triassic, the area of the future Therefore, numerous taxa of benthic foraminifera, the Karst Dinarides represented part of the Gondwanian chronostratigraphic range of which were already deter­ epicontinental shelf (see discussion in VLAHOVIĆ et mined in other Peri-Mediterranean platforms, represent al., 2005). a very important, and sometimes even the only tool for From the Middle Triassic to the middle Early the stratigraphic subdivision of the Karst Dinarides. Jurassic, the area of the present Karst Dinarides was This paper considers benthic foraminiferal taxa and part of the huge, more or less isolated carbonate plat­ assemblages from the stratigraphic viewpoint, as well form (free from continental influences), which covered as examining their importance for the stratigraphic a large part of present Southern and Central Europe. determination of shallow-marine carbonate sequences, This was called the Southern Tethyan Megaplatform especially for geological mapping. In addition, the pal­ by VLAHOVIĆ et al. (2005), and it disintegrated into aeobiogeography of the most important (mostly index several smaller, relatively isolated carbonate platforms fossils), is presented. Major attention was given to the during Late Sinemurian to Late Toarcian times. These so-called larger foraminifera, while smaller forms were smaller platforms included, among others, the Adriatic, discussed only if they represent index fossils. Apulian and Apenninic platforms (VLAHOVIĆ et al., 2002, 2005; DRAGIČEVIĆ & VELIĆ, 2002). The Adriatic Carbonate Platform s.str. existed from 2. FORAMINIFERAL TAXA AND the late Early Jurassic (Pliensbachian/Toarcian) to near ASSEMBLAGES IN THE CHRONO­ the end of the Cretaceous, when it was finally destroyed. STRATIGRAPHIC SUCCESSION It represented an almost completely isolated shallow- marine depositional system, which was probably con­ Some periods in the geological history of the Adriatic nected towards the S and SE to the Kruja and Gavrovo– Carbonate Platform and its basement were character­ Tripolitza platforms (VLAHOVIĆ et al., 2005). During ized by the predominance of certain groups of benthic the Mesozoic period the platform continuously migrated foraminifera. For example, the Early and Middle Tri­ from the Southern Tethyan realm, where it was placed assic were characterized by amodiscids, Late Triassic during the Early Jurassic, towards the N and NE, to the by involutinids, and the Jurassic by lituolids. During Northern areas of this palaeo-ocean in the Late Maas­ the Cretaceous, diversification was significant. Several trichtian. It was surrounded by deep-marine basins: the groups of smaller foraminifera characterized the Neo­  Geologia Croatica 60/1

Fig. 1 Recent area covered by the Adriatic Carbonate Platform deposits. comian. From the Barremian to the Middle Cenoma­ Genera and species, either individually or within nian the most important foraminifera were orbitolinids foraminiferal assemblages, are mostly presented in their accompanied by ataxophragmiids, lituolids, cyclammi­ order of occurrence. nids as well as frequent miliolids, cuneolinids, etc. The evolution of soritids and alveolinids can be traced from 2.1. Triassic the Aptian, while orbitoids and siderolitids were impor­ As the shallow-marine deposits accumulated during the tant during the Campanian and Maastrichtian. Triassic belong to the basement of the Adriatic Carbon­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera...  ate Platform, only their general foraminiferal taxa and 2.1.2. Middle Triassic assemblages are presented. After the Late Permian glo­ General characteristics: Anisian and Ladinian forami­ bal biological crisis, when most of the Late Palaeozoic niferal assemblages show great differences, both foraminifera (including the most important group, the in their composition and stratigraphic value. While fusulinids) became extinct, there was a gradual restora­ Anisian deposits contain several index species, none tion of shallow-marine environments, which were eco­ have so far been discovered in the Ladinian sedi­ logically suitable for the existence of benthic foraminif­ ments, where smaller foraminifera predominate. era. During the Early Triassic, carbonate deposition was 2.1.2.1. Anisian characterized by an important terrigenous influence, Foraminiferal assemblage: Meandrospira dinarica, and mixed carbonate–siliciclastic environments were Pilammina densa, Pilamminella semiplana, Hoy­ not favourable as niches allowing the larger foraminif­ enella gr. sinensis, Duostomina alta, Neoendothyra era to thrive. Therefore, the Early Triassic represented reicheli, Turriglomina mesotriassica, Arenovidalina a period during which smaller foraminifera (mostly chialingchiangensis, Endotriadella wirtzi, Fron­ ammodiscids) predominated. In the Middle Triassic, dicularia woodwardi, Ammobaculites radstadtensis, shallow-marine environments characterized by ‘pure’ Aulotortus sp., Haplophragmium sp., Glomospira carbonate deposition were restored, but in some areas sp., Glomospirella sp. and other smaller foramini­ temporary lateral replacement by deeper-marine car­ fera. bonate–volcanoclastic deposits occurred. From a palae­ Discussion: Anisian index species, Meandrospira dina­ ontological point of view, an abundant flora of calcar­ rica, Pilammina densa and Pilamminella semiplana, eous algae, including in particular the Diploporaceaen are widely distributed, and have been described in family of the Dasycladales, characterized these environ­ the area of the Karst Dinarides (PANTIĆ, 1965; ments. Reefal–perireefal environments favourable for KOCHANSKY-DEVIDÉ & PANTIĆ, 1966). the development of corals, hydrozoans, molluscs, etc. They were also cited in later papers (PANTIĆ- were also very common. Benthic foraminifera were not PRODANOVIĆ, 1975; GRGASOVIĆ et al., 2000; very important, but the most frequent were still ammo­ GRGASOVIĆ, 2003). Other taxa have a wider discids, meandrospirids, endotriadids, duostominids stratigraphic range, and most of them also occur in etc. During the Late Triassic involutinids were the most Ladinian deposits. important foraminifera. 2.1.2.2. Ladinian 2.1.1. Early Triassic – Scythian Foraminiferal assemblage: Most taxa cited in the Ani­ (Induan and Olenekian) sian assemblage also occur in Ladinian deposits, General characteristics: Early Triassic foraminifera except for Anisian index forms. However, the most were not systematically investigated in the Karst frequent are specifically undeterminable forms of Dinarides, although they are very abundant at some Diplotremina and Duostomina. localities. Their occurrence is of great stratigraphic Discussion: Ladinian foraminifera have not been pre­ value, because they frequently represent the only viously systematically determined because of their palaeontological evidence for the determination of scarcity and the lack of index forms. Therefore, Lower Triassic deposits. Therefore, the most fre­ stratigraphic determination and distinction between quent genera and index species are mentioned here. Anisian and Ladinian deposits has been based Foraminiferal assemblage: Cyclogyra mahayeri, on calcareous algal assemblages. On the basis of Meandrospira pusila, Ammodiscus parapriscus, detailed analysis by GRGASOVIĆ (2003) it may be Arenovidalina chialingchiangensis, Glomospirella concluded that there are rare occurrences of Aulotor­ triphonensis and numerous other undeterminable tus species in the Ladinian (e.g. A. praegaschei and smaller foraminifera, mostly ammodiscids. A. sinuosus). In the present area of the Karst Dinar­ Discussion: This assemblage is characteristic of the ides these also occur in Anisian deposits, reaching Early Triassic of the Karst Dinarides in the Induan– maximum frequency during the Norian and Rha­ Olenekian stratigraphic range. In his comprehensive etian. and detailed treatise of Croatian Triassic stratigra­ 2.1.3. Late Triassic phy, which also included the NE parts of the AdCP basement in the vicinity of Karlovac GRGASOVIĆ General characteristics: Late Triassic foraminiferal (2003) separated two foraminiferal zones: the older assemblages are not especially rich in genera and Cyclogyra mahayeri zone and younger Meandros­ species. This is the result of palaeogeographic rela­ pira pusila zone, and he cited most of the species tionships, since the area of the present Karst Dinar­ listed above. Some of these forms were also listed in ides was emergent from the Late Ladinian to the some older papers discussing the platform area, e.g. Early Norian (VELIĆ et al., 2002; VLAHOVIĆ et KOCHANSKY-DEVIDÉ & PANTIĆ (1966) and al., 2005). Therefore, older Upper Triassic deposits PANTIĆ-PRODANOVIĆ (1975). (Carnian and Lower Norian), can only be observed  Geologia Croatica 60/1

along the marginal parts of the platform area, e.g. in Discussion: Triasina hantkeni is the index fossil of the the vicinity of Karlovac, from where GRGASOVIĆ Rhaetian. The maximum frequency of the cited spe­ (2003) listed rich foraminiferal assemblages. During cies of Aulotortus which appeared in the Late Nori­ the Norian and Rhaetian, limestones and early-dia­ an continued into the Early Rhaetian. genetic dolomites were deposited within peritidal environments. Limestones mostly suffered late-dia­ 2.2. Jurassic genetic dolomitization, so rocks of this age, usu­ During the Jurassic in the area of the present Karst ally known as the Main Dolomite (Hauptdolomit Dinarides, lituolids represented the major foraminif­ or Dolomia principale), only sporadically contain eral group. They occurred in the Sinemurian and had undolomitized lenses and thinner packages of lime­ already reached their maximum in the Pliensbachian. stones with Late Triassic fossil assemblages, includ­ They were very frequent in the Middle Jurassic, when ing frequent foraminifera. The most important group together with cyclamminids, valvulinids (especially are involutinids, including some index species of the pfenderinids), as well as other groups they represent the Late Triassic. major fossils used for stratigraphic division. The larg­ 2.1.3.1. Carnian est number of index fossil taxa occurred in Bathonian deposits. During the Late Jurassic, the number of taxa, Foraminiferal assemblage: Lamelliconus multispirus, as well as individual specimens decreased. L. procerus, Aulotortus friedli, A. praegaschei, A. sinuosus, A. tenuis and numerous smaller foraminif­ 2.2.1. Early Jurassic era of greater, Middle and Upper Triassic strati­ graphic range. General characteristics: In several very important papers, SEPTFONTAINE (1980, 1984, 1988; SEP­ Discussion: Due to emergence, Carnian deposits are TFONTAINE et al., 1991) presented the best evo­ either missing, or are represented by terrigenous lution, classification/taxonomy, biostratigraphy clastic sediments over much of the Karst Dinar­ and palaeobiogeography of the Jurassic lituolids. ides. Shallow-marine carbonates were only depos­ Subsequently BASSOULET (1997), in his revi­ ited in some places which were not emergent, e.g. sion of the stratigraphic ranges of larger Jurassic central Slovenia, or the vicinity of Karlovac in foraminifera, mostly lituolids, confirmed Septfon­ Croatia along the NE margin, as well as Komiža on taine’s thoughts on the Early Jurassic lituolids. This the island of Vis, the vicinity of Trebinje in Herze­ is especially important for the discussion of Early govina or in Montenegro along the SW and SE mar­ Jurassic foraminiferal assemblages, because almost gins. Within these rocks a Carnian algal–foraminif­ all genera and species that Septfontaine cited from eral assemblage has been discovered, including the Moroccan High Atlas (Fig. 2b), and Bassoulet in index forms foraminifera Lamelliconus multispirus Northern Africa and Western Europe (Fig. 2c) occur and L. procerus. In addition to these two forms, in the Adriatic Carbonate Platform (Fig. 2a). How­ GRGASOVIĆ (2003) also cited numerous other ever, their occurrence did not commence at the Tri­ species of smaller foraminifera. assic–Jurassic boundary but later in the Sinemurian. 2.1.3.2. Norian 2.2.1.1. Hettangian Foraminiferal assemblage: Aulotortus friedli, A. prae­ Foraminiferal assemblage: Involutina liassica, Duo­ gaschei, A. sinuosus, A. tenuis, A. tumidus, Auloco­ taxis metula, Amijiella amiji, Siphovalvulina varia­ nus permodiscoides, Trocholina crassa, T. acuta and bilis and Everticyclammina sp. – ?E. praevirguliana. Turrispirillina minima. Discussion: There are no very important foraminif­ Discussion: The aforementioned species of Aulotortus eral taxa in the Hettangian deposits. Therefore, the reached their maximum development in the Late stratigraphy of this stage on the AdCP is based on Norian and Rhaetian. They were documented in the Dasycladacean calcareous algae, mostly on species regionally very well known Periadriatic and Central of Palaeodasycladus (SOKAČ, 2001), among which European formations of the Main Dolomite (Haupt­ P. mediteraneus is the most frequent and ranges dolomit or Dolomia principale) and Dachstein lime­ from the Hettangian to the end of the Pliensbachi­ stones of Upper Triassic age. In the Karst Dinarides, an. Involutina liassica occurs throughout the Lower the Main Dolomite is of Late Norian and Rhaetian Jurassic deposits, but only in the rocks originating age. In older papers (PANTIĆ, 1966/67; PANTIĆ- at the platform margins and slopes. Duotaxis metula PRODANOVIĆ, 1975) Turrispirillina minima was is characterized by a stratigraphic range throughout described as a Norian species. the Early Jurassic, while Amijiella amiji has an even wider range, from the Early to Middle Jurassic. 2.1.3.3. Rhaetian 2.2.1.2. Sinemurian Foraminiferal assemblage: Triasina hantkeni, Aulo­ conus permodiscoides, Aulotortus friedli, A. sinu­ Foraminiferal assemblage: osus, A. tenuis, A. tumidus, A. pokornyi, Trocholina (a) Early Sinemurian: Mesoendothyra sp., Siphovalvu­ crassa and T. acuta. lina gibraltarensis, S. colomi, S. variabilis, Duotaxis Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 

Hett. Sinemurian Pliensbachian Toarcian

A

Pliensbachian Hett. Sinemurian Toarcian Carixian Domerian

B

Hettangian Sinemurian Pliensbachian Toarcian

Early Late Carixian Domerian C

Fig. 2 Stratigraphic ranges of the Early Jurassic benthic foraminifera on the AdCP (A) and comparison with stratigraphic ranges in High Atlas (B – after SEPTFON- TAINE, 1984) and Western Europe and Northern Africa (C – after BASSOULLET, 1997). Stratigraphic ranges are shown according to their duration in MY (time scale after GRADSTEIN et al., 2004).  Geologia Croatica 60/1

metula, Amijiella amiji, Everticyclammina praevir­ in the Early Carixian, O. praecursor in the Late guliana, Meandrovoluta asiagoensis, Pseudocycla­ Carixian, and the occurrence of the Early Domer­ mmina sp. and Involutina liassica. ian index fossil O. dubari is assumed. P. liassica is (b) Late Sinemurian: Mesoendothyra sp., A. amiji, D. an index form of the Late Domerian. Observations metula, S. variabilis, S. gibraltarensis, S. colomi, E. of biokovinids – Biokovina gradacensis and Bos­ praevirguliana, M. asiagoensis, Lituosepta recoa­ niella oenensis – are rare and mostly occur in the rensis, Orbitopsella primaeva, Haurania deserta, tectonized parts of the southern Karst Dinarides, so Lituolipora termieri, Planisepta compressa, Agerina their stratigraphy may be approximated to within the martana and I. liassica. Pliensbachian. P. compressa reaches its maximum frequency in the Domerian. L. termieri is an index Discussion: The Early Sinemurian is characterized form for the Late Sinemurian and Carixian. SEPT­ by the appearance of more abundant foraminiferal FONTAINE (1980, 1988) considered the genus Bos­ assemblage, but without very good index forms. The niella as a synonym of the genus Mesoendothyra. most important is Mesoendothyra sp., which accord­ ing to SEPTFONTAINE (1984, 1988) is the source L. recoarensis has been discovered within the ear­ form of the most important Early Jurassic lituolids in liest Pliensbachian (Early Carixian) in the Karst the Southern Tethyan Bioprovince. This foraminifera Dinarides, although occurrences are rare. This con­ has been found in many localities on the AdCP, and firms the partial overlap of its stratigraphic range is especially abundant in Velebit Mt. (TIŠLJAR et with that of O. primaeva in detailed profiles inves­ al., 1991), where it occurs in the Early Sinemurian. tigated in the Velebit Mt. area. In Western Tethys it According to SEPTFONTAINE (1984, 1988), two is exclusively a Sinemurian species, as illustrated lines of Early Jurassic lituolids evolved from this by SEPTFONTAINE (1984) and BASSOULLET form in the Late Sinemurian and Pliensbachian: (1) (1997). Towards the East, in Italy, on the Apennin­ Mesoendothyra sp. – Lituosepta? recoarensis – O.? ic Carbonate Platform CHIOCCHINI et al. (1994) primaeva – O. praecursor – O. dubari, and (2) Mes­ found this form within (or even after) the strati­ oendothyra sp. – Lituolipora termieri – Planisepta graphic range of O. praecursor, in the Monte Sor­ compressa, which has also been confirmed on the genza and Monte della Selva columns, indicating AdCP. KABAL & TASLI (2003) described Lituoli­ that in the Apennines L. recoarensis was definitely pora termieri as a synonym of both Paleomayncina of Carixian, and possibly even Early Domerian age. termieri (HOTTINGER, 1967; SEPTFONTAINE, 2.2.1.4. Toarcian 1988) and Lituolipora polymorpha (GUŠIĆ & Foraminiferal assemblage: E. praevirguliana, Mesoe­ VELIĆ, 1978). The only index forms for the Late ndothyra croatica, I. liassica, A. martana, S. varia­ Sinemurian are L. recoarensis (its entire stratigraph­ bilis, S. gibraltarensis, S. colomi and M. asiagoen­ ic range is within this part of the stage), accompa­ sis. nied by O. primaeva occurring in Late Sinemurian and Early Pliensbachian. During the Sinemurian H. Discussion: Due to palaeoenvironmental and sedimen­ deserta occurs which is similar to A. amiji, and is tological circumstances, larger foraminifera have never been observed in the youngermost Lower observed in the later stages (Pliensbachian, Aalenian Jurassic deposits of the Adriatic Carbonate Plat­ and Bajocian). form, nor on neighbouring platforms in the South­ 2.2.1.3. Pliensbachian ern Tethyan realm. As a consequence of the intense Foraminiferal assemblage: regional extensional tectonic movements described in the introduction, the Adriatic Carbonate Plat­ (a) Early Pliensbachian – Carixian: Mesoendothyra form separated from the neighbouring Apulian and sp., L. recoarensis, Orbitopsella praecursor, O. pri­ Apenninic ones during the Toarcian (VLAHOVIĆ maeva, H. deserta, A. amiji, P. compressa, L. ter­ et al., 2002, 2005; DRAGIČEVIĆ & VELIĆ, 2002). mieri, Biokovina gradacensis, Bosniella oenensis, A. Western areas of the Adriatic Carbonate Platform martana, D. metula, S. variabilis, S. gibraltarensis, were temporarily drowned, resulting in deposition S. colomi, E. praevirguliana, M. asiagoensis and I. of bioturbated mudstones and subordinate peletal– liassica. ooid wackestone/packstones, the so-called ‘spotty (b) Late Pliensbachian – Domerian: Mesoendothyra limestones’. This event correlates with a Toarcian sp., O. praecursor, O. ?dubari, H. deserta, A. amiji, Oceanic Anoxic Event (JENKYNS, 1988, 1991; L. termieri, P. compressa, B. gradacensis, B. oenen­ VLAHOVIĆ et al., 2005). Penecontemporaneous­ sis, D. metula, Pseudocyclammina liassica, S. varia­ ly, the NE margin emerged, while in the SE, high- bilis, S. gibraltarensis, S. colomi, E. praevirguliana, energy shallows were locally formed, characterized M. asiagoensis, A. martana, in places calcisphaer­ by deposition of beach and bar ooid grainstones. ulids and I. liassica. Generally, depositional environments throughout Discussion: Within the Carixian foraminiferal assem­ the platform area were unfavourable for the growth blage, the occurrence of Orbitopsella species are of benthic organisms of all kinds, not only for very important. O. primaeva reached its maximum foraminifera, which are very scarce. Interesting first Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 

Aalenian Bajocian Bathonian Callovian

Fig. 3 Stratigraphic ranges of the Mid- dle Jurassic ben- thic foraminifera on the AdCP and dasycladalean alga Selliporella donzellii for comparison with foraminiferal ranges.

occurrences of rare forms close to M. croatica were 2.2.2.1. Aalenian recorded in the ‘spotty limestones’, representing an Foraminiferal assemblage: age probably from the beginning of the Toarcian to (a) Early Aalenian: Mesoendothyra croatica, Gutni­ the beginning of the Aalenian. This species had the cella cayeuxi, Pseudocyclammina maynci, Amijiella major part of its range in the Middle Jurassic (to the amiji, Haurania deserta, Redmondoides lugeoni, Early Bathonian), and its maximum in the Bajocian. Siphovalvulina variabilis and Everticyclammina sp. The Toarcian represents the end of the stratigraphic range of I. liassica, which has never been found in (b) Late Aalenian: M. croatica, G. cayeuxi, P. maynci, the inner part of the platform. A. amiji, H. deserta, R. lugeoni, S. variabilis, Timi­ donella sarda, Spiraloconulus perconigi and Everti­ 2.2.2. Middle Jurassic cyclammina sp. General characteristics: Before late 1980’s, Middle Discussion: All cited taxa were determined in the S Jurassic platform carbonates were considered to be and SE parts of the AdCP, and locally in the NE, quite unfossiliferous, except for the part belonging to marginal parts of the platform. In these areas Mid­ the Bathonian. However, new investigations proved dle Jurassic carbonates are mostly represented by that they are relatively rich in a number of fossils. ooid, bioclastic and intraclastic limestones depos­ According to the number of determined fossil taxa, ited in the higher water energy environments of the they represent the most fossiliferous Jurassic depos­ platform shallows. They are relatively rich in their its (Fig. 3). Among the larger foraminifera, lituolids, foraminiferal assemblages. However, in the central, hauraniids, cyclamminids and pfenderinids were relatively deeper and more restricted areas, predom­ determined. Most of these taxa represent index fos­ inantly thick-bedded mudstones were deposited, sils, most frequently in the Upper Aalenian, Lower characterized by relatively rare findings of the cited Bajocian and Bathonian deposits. taxa, with the exception of M. croatica which is rel­ atively frequent. Among the stratigraphically more important species, G. cayeuxi has a stratigraphic 10 Geologia Croatica 60/1

range through most of the Aalenian, and is accom­ 2.2.2.3. Bathonian panied by S. perconigi and T. sarda in the Upper Foraminiferal assemblage: Aalenian deposits. These two latter species have (a) Early Bathonian: M. croatica, S. variabilis, R. luge­ not been previously determined within the Middle oni, P. striata, P. arabica, T. gr. alpina, P. elongata, Jurassic carbonates of the AdCP. P. crusei, Paleopfenderina salernitana, P. trochoi­ 2.2.2.2. Bajocian dea, Nautiloculina gr. oolithica and Everticyclam­ Foraminiferal assemblage: mina sp. (a) Early Bajocian: M. croatica, P. maynci, G. cayeuxi, (b) Late Bathonian: S. variabilis, R. lugeoni, P. stria­ T. sarda, A. amiji, H. deserta, R. lugeoni, S. variabi­ ta, P. arabica, T. gr. alpina–elongata, P. elongata, lis, Spiraloconulus giganteus, S. perconigi, Pseudo­ P. crusei, P. salernitana, P. trochoidea, Alzonella dictyopsella jurassica, Marzoella ficcarellii, Proto­ cuvillieri, Satorina apuliensis, Satorina mesojurassi­ peneroplis striata, Nautiloculina sp. – probably N. ca, Kilianina blancheti, Orbitammina elliptica, N. gr. oolithica, Trocholina sp. and Everticyclammina gr. oolithica and Everticyclammina sp. sp. Discussion: The Bathonian is the richest stage within (b) Late Bajocian: M. croatica, S. giganteus, A. amiji, Middle Jurassic deposits of the AdCP with regard H. deserta, R. lugeoni, S. variabilis, P. striata, Par­ to the abundance of larger foraminifera. Within this avalvulina complicata, Pseudoeggerella elongata, assemblage index fossils of the Early Bathonian are Pfenderella arabica, Praekurnubia crusei, N. gr. absent, but the age is reliably indicated between the ?oolithica, Trocholina gr. alpina, Trocholina sp. and beginning of the stratigraphic range of P. salerni­ Everticyclammina sp. tana and the first occurrence of the following index fossils of the Late Bathonian: A. cuvillieri, S. apuli­ Discussion: Forms typical of Aalenian deposits also ensis, S. mesojurassica, K. blancheti and O. ellip­ occur in the earliest Bajocian, including G. cayeuxi, tica. P. elongata, S. mesojurassica and A. cuvillieri S. perconigi and T. sarda. Above these, during the have only a single record in the AdCP area (VELIĆ end of the Early and the beginning of the Late Bajo­ et al., 1995), but have not been previously illustrat­ cian, P. jurassica and M. ficcarellii (index species) ed. O. elliptica has been observed here for the first occur with N. gr. ?oolithica. Although BASSOUL­ time. LET (1997) indicated a stratigraphic range for S. giganteus from the Early Bajocian to the Early P. salernitana is the most frequent and most widely Callovian, with the most definite occurrences in the distributed Bathonian species on the AdCP. S. apuli­ Bathonian, this form only occurs in the Bajocian ensis, and other Bathonian species are less common. on the AdCP. M. croatica reaches its maximum in 2.2.2.4. Callovian the Early Bajocian, while P. striata is at the begin­ Foraminiferal assemblage: P. salernitana, S. variabi­ ning of its stratigraphic range, which extends until lis, R. lugeoni, P. striata, P. arabica, T. gr. alpina– the end of the Jurassic in the AdCP. In the topmost elongata, P. crusei, N. gr. oolithica, Mohlerina basi­ levels of Bajocian carbonates, P. arabica and T. liensis, Kurnubia jurassica, probably Chablaisia gr. alpina were determined, and these also occur chablaisensis and Everticyclammina sp. in the younger stages. The index form of younger Upper Bajocian deposits is P. complicata. P. elon­ Discussion: There are no specific index forms in the gata occurs in the continuous succession from top­ Callovian sediments: this assemblage has a ‘transi­ most Upper Bajocian beds and are more common tional’ character with taxa that can be traced from in the Bathonian. Trocholina species of the T. alpi­ older Middle Jurassic to Late Jurassic stages: R. na–elongata group were reported within the genus lugeoni, P. crusei, P. arabica, P. striata and T. gr. Andersenolina (NEAGU, 1994). This view has been alpina–elongata. This assemblage also comprises followed by some, e.g. BUCUR et al. (2004), but the beginning of the M. basiliensis range (TASLI, not by all authors, e.g. CHIOCCHINI et al. (1994), 1993 and CHIOCCHINI et al., 1994, this form being RADOIČIĆ (2005), etc. Until recently S. giganteus, attributed to Archaeosepta). By the end of the Callo­ P. jurassica, M. ficcarellii, P. complicata and P. vian the first kurnubias occur, e.g. K. jurassica. elongata were not known within the Bathonian, or Ch. chablaisensis has only been observed at one generally in the Middle Jurassic in the area of the locality in Biokovo Mt., within tectonically very AdCP. They have very recently been reported by disturbed deposits also containing P. crusei. Other VELIĆ (2005). The first occurrences of P. crusei index forms are absent, but neighbouring older occur in transitional beds at the base of the Batho­ outcrops are of Bathonian age. Ch. chablaisensis nian. By the end of the Early Bajocian Selliporella is only previously known from Oxfordian deposts donzellii appears, and its stratigraphic range includes in the AdCP area (VELIĆ & TIŠLJAR, 1988; the earliest Late Bathonian. This dasyclad alga is TIŠLJAR & VELIĆ, 1993; VELIĆ et al., 1995). At very important for the stratigraphy of the Middle the aforementioned localities other Oxfordian forms Jurassic shallow-marine carbonate succession of the are also absent. Therefore, it could be concluded that Karst Dinarides. this occurrence probably indicates somewhat older, Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 11

Oxfordian Kimmeridgian Tithonian

Fig. 4 Stratigraphic ranges of the Late Jurassic benthic foraminifera on the AdCP.

probably Callovian or even Bathonian deposits (see (b) Late Oxfordian: Labyrinthina mirabilis, Kurnubia Fig. 3). wellingsi, K. jurassica, K. palastiniensis, K. uzbekis­ tanica, Ch. chablaisensis, N. oolithica, N. circu­ 2.2.3. Late Jurassic laris, T. alpina, T. elongata, T. gigantea, P. lituus, General characteristics: Among the foraminifera in S. variabilis, R. lugeoni, P. arabica, P. crusei, M. the Upper Jurassic carbonates of the AdCP, those basiliensis, P. striata and E. virguliana. belonging to the genus Kurnubia are the most Discussion: In the assemblage cited above, the only important. In spite of the occurrence of either other form which represents an index fossil for the Oxfor­ index species or those species that are very abundant dian is K. uzbekistanica (occuring in quiet subtidal in certain levels, the stratigraphic range of different and/or lagoonal facies). T. gigantea, determined in species from Late Callovian to the Middle Tithonian Middle Oxfordian deposits is significant, but is very designates this genus as typically indicating a Late rare (mostly in higher energy and tempestite facies). Jurassic age (Fig. 4). Several species of the genus Kurnubia forms predominate, and K. palastiniensis Kurnubia have been described in the Perimediter­ is the most frequent species, while maximum fre­ ranean area, while on the AdCP the following spe­ quency was also achieved by T. alpina, T. elongata cies have been documented: K. palastiniensis, K. and C. chablaisensis. The latter form disappeared wellingsi (e.g. RADOIČIĆ, 1966; GUŠIĆ, 1969a, in the late Oxfordian, and it frequently occurs in b; VELIĆ, 1977, etc.) and K. jurassica (GUŠIĆ, the tempestitic grainstones. The beginning of the 1969a, b). stratigraphic range of L. mirabilis and K. wellingsi is considered to represent the approximate boundary 2.2.3.1. Oxfordian between the Early and Late Oxfordian. N. circula­ Foraminiferal assemblage: ris appeared by the end of the Early Oxfordian, and (a) Early Oxfordian: Pseudocyclammina lituus, reached its maximum in the Late Oxfordian. Siphovalvulina variabilis, Redmondoides lugeoni, 2.2.3.2. Kimmeridgian Pfenderella arabica, Praekurnubia crusei, Mohle­ rina basiliensis, Protopeneroplis striata, Kurnubia Foraminiferal assemblage: palastiniensis, K. jurassica, Karaisella uzbekista­ (a) Early Kimmeridgian: S. variabilis, E. virguliana, nica, Trocholina alpina, T. elongata, T. gigantea, R. lugeoni, P. arabica, P. crusei, P. striata, K. jura­ Chablaisia chablaisensis, Nautiloculina oolithica, ssica, K. palastiniensis, K. wellingsi, L. mirabilis, T. N. circularis and Everticyclammina virguliana. alpina, T. elongata, M. basiliensis, P. lituus, Coni­ 12 Geologia Croatica 60/1

cocurnubia orbitoliniformis, Alveosepta jaccardi, if orbitolinids are the most important, cuneolinids and Kilianina? rahonensis and Parurgonina caelinensis. miliolids were the most abundant Cretaceous foraminif­ (b) Late Kimmeridgian: S. variabilis, E. virguliana, R. era. Besides these groups, Early Cretaceous foraminife­ lugeoni, P. striata, T. alpina, T. elongata, M. basil­ ral assemblages include numerous smaller forms – invo­ iensis, K. palastiniensis, K. wellingsi, P. lituus, C. lutinids, ataxophragmiids, novalesinids, and cyclammi­ orbitoliniformis, A. jaccardi, K.? rahonensis and P. nids. From the Aptian to the Late Cretaceous, alveoli­ caelinensis. nids and nezzazatids became more and more frequent, while during the Late Cretaceous soritids, fischerinids, Discussion: Index forms for the Kimmeridgian deposits pavonitids, dicyclinids, rhapydioninids and rotaliids of the AdCP are C. orbitoliniformis and A. jaccar­ increased noticeably. Orbitoidids and siderolitids were di, and partially K.? rahonensis, which has a strati­ important on the platform slopes and in the deeper intra­ graphic range through the major part of the Kim­ platform troughs formed in the Late Cretaceous. meridgian, and P. caelinensis in the terminal Early and Late Kimmeridgian. However, the latter two 2.3.1. Early Cretaceous species can also be observed in Lower Tithonian General characteristics: The end of the Jurassic and deposits. L. mirabilis, P. crusei, P. arabica and K. the beginning of the Cretaceous are character­ jurassica disappeared in the Early Kimmeridgian. ized by the emergence of a large part of the central For stratigraphic correlation, the important dasyclad AdCP (VELIĆ et al., 2002). Deposits from areas algae are Pseudoclypeina cirici (index fossil of the with continuous deposition were late-diagenetically Early Kimmeridgian), as well as the first occurrence dolomitized, while in the shallowest areas early-dia­ of the Late Jurassic species Clypeina jurassica. genetic dolomites were formed. Therefore, within Dasyclad algae are distributed throughout the AdCP these unfavourable conditions foraminiferal assem­ area from the Middle Kimmeridgian to the transi­ blages were reduced, especially those of the larger tional level between the Tithonian and Berriasian. foraminifera. Until the first occurrence of orbi­ 2.2.3.3. Tithonian tolinids in the Late Barremian, smaller foraminifera Foraminiferal assemblage: were predominant. The richest foraminiferal assem­ (a) Early Tithonian: S. variabilis, E. virguliana, R. blages were determined from Early Aptian and lugeoni, P. striata, T. alpina, T. elongata, M. basil­ Albian deposits (Fig. 5). During the Late Aptian, iensis, K. palastiniensis, P. lituus, K.? rahonensis there was a decrease in the number of foraminife­ and P. caelinensis. ral taxa, as a result of the regional relative sea-level fall and frequent short periods of emergence (e.g. (b) Late Tithonian: E. virguliana, R. lugeoni, P. stria­ VLAHOVIĆ et al., 2005), as well as longer local ta, T. alpina, T. elongata, M. basiliensis, K. palas­ emergent phases, e.g. in Western Istria (VELIĆ et tiniensis, P. lituus, Protopeneroplis ultragranulata, al., 1989). Anchispirocyclina lusitanica and/or A. neumannae. The most important studies of palaeontology, bios­ Discussion: In the youngest Upper Jurassic carbonates tratigraphy and modern systematics of orbitolinids of the AdCP there are almost no typical index forms are those of Schroeder (particularly SCHROEDER, of foraminifera. Very rarely, specimens of Anchispi­ 1975, 1979). These two papers on the general evo­ rocyclina occur, which are specifically almost lutionary trends, definitions and origin of orbitoli­ impossible to determine because of significant mic­ nids from the Early Aptian to Middle Cenomanian ritization, probably belong to index Tithonian forms resulted from his earlier studies (e.g. SCHROED­ including A. lusitanica and/or A. neumannae. All the ER, 1962, 1964a, b, c). The evolutionary line, which cited species except the trocholinas, R. lugeoni, P. commenced in the Early Aptian, with Praeorbitoli­ lituus and P. ultragranulata disappeared in Titho­ na cormyi – P. wienandsii – Mesorbitolina lotzei nian: first P. caelinensis then K.? rahonensis in the has also been documented on the Adriatic Carbon­ Early Tithonian, K. palastiniensis at the beginning ate Platform (VELIĆ & SOKAČ, 1978a, b; VELIĆ, of the Late Tithonian, and A. lusitanica/neuman­ 1988). nae at the end of Tithonian. According to BUCUR (1993), P. ultragranulata has a wider stratigraphic 2.3.1.1. Berriasian range from the Middle Tithonian to the Early Bar­ Foraminiferal assemblage: remian, but on the AdCP it has only been found in (a) Early Berriasian: Mohlerina basiliensis, Trocholi­ the Late Tithonian and Early Berriasian. na alpina, T. elongata, Pseudocyclammina lituus, Protopeneroplis ultragranulata, Mayncina bulgari­ 2.3. Cretaceous ca and Charentia cuvillieri. The orbitolinids are the most important Cretaceous larg­ (b) Late Berriasian: M. basiliensis, T. alpina, T. elon­ er foraminifera. On the AdCP they appear in the Late gata, P. lituus, M. bulgarica, C. cuvillieri, Montsa­ Barremian, and inhabited the platform, with temporary levia salevensis, Meandrospira favrei, Haplophrag­ breaks, until the end of the Cretaceous. Almost all orbi­ moides joukowskyi, Vercorsella tenuis, V. campo­ tolinid species have stratigraphic importance. However, saurii and Praechrysalidina sp. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 13

Fig. 5 Stratigraphic ranges of the Early Creta- ceous benthic foraminifera on the AdCP. 14 Geologia Croatica 60/1

Discussion: In the transitional level between the Titho­ Campanellula capuensis, Pseudonummoloculina sp. nian and Berriasian, and for the Early Berriasian, and Praechrysalidina sp. the index form is P. ultragranulata. The Berriasian Discussion: N. distorta occurs in the Early Hauterivian. represents the end of the stratigraphic range of M. The Late Hauterivian is characterized by the disap­ basiliensis. The beginning of this stage is character­ pearance of T. gr. alpina–elongata. The entire stage ized by the first occurrences of M. bulgarica (with is characterised by several species which continued a stratigraphic range throughout the Early Creta­ from the Valanginian, e.g. T. sagittaria, V. scarsell­ ceous) and infrequent observations of C. cuvillieri ai, M. bulgarica, C. cuvillieri and S. minuta. Species (the stratigraphic range of which continued into the appearing for the first time in the latest Hauterivian, Late Cretaceous). First occurrences of small Prae­ and occurring throughout the younger stages of the chrysalidina sp. (which may be ancestors of Prae­ Early Cretaceous are D. hahounerensis, N. producta chrysalidina infracretacea that ranges from the Late and V. laurentii. This group also contains species Barremian until the Late Albian), were documented of Pseudonummoloculina, similar to P. aurigerica. from Upper Berriasian to Lower Barremian depos­ Nezzazata germanica represents an index species, its. Near the end of the Berriasian, important species but is very rare, while in the latest Hauterivian and of smaller foraminifera occur, including M. saleven­ transitional beds to the Barremian C. capuensis is sis, M. favrei, H. joukowskyi, V. tenuis and V. cam­ very common. posaurii, which reached their maximum in the Val­ anginian. 2.3.1.4. Barremian Foraminiferal assemblage: 2.3.1.2. Valanginian (a) Early Barremian: N. distorta, N. cornucopia, N. Foraminiferal assemblage: Trocholina sagittaria, T. producta, Vercorsella wintereri, V. scarsellai, V. lau­ delphinensis, T. alpina–elongata, P. lituus, M. bul­ rentii, M. bulgarica, T. sagittaria, D. hahounerensis, garica, C. cuvillieri, M. salevensis, M. favrei, H. joukowskyi, Vercorsella scarsellai, V. camposaurii, S. minuta, Spiroloculina cretacea, Rumanoloculina V. tenuis, Sabaudia minuta, as well as Novalesia sp. robusta, Belorussiella textilarioides, Glomospira and Praechrysalidina sp. watersi, Pseudonummoloculina sp., Praechrysalidi­ na sp. and involutinids – ‘Involutina’ sp. Discussion: Index fossils for the Valanginian of the AdCP are M. salevensis, H. joukowskyi, M. favrei, (b) Late Barremian: all taxa from the Early Barremian V. tenuis, and V. camposaurii, which is particular­ are present but are also accompanied by Rectodic­ ly common. All these forms appear near the end of tyoconus giganteus, Palorbitolina lenticularis, Tro­ the Berriasian, and disappear in the Early Hauteriv­ cholina molesta, T. odukpaniensis, Vercorsella are­ ian. There are additional forms that first occur in nata, Derventina filipescui, Neotrocholina fribur­ the Valanginian, and others that are also present in gensis, N. aptiensis, Nautiloculina broennimanni, younger intervals of the Early Cretaceous, includ­ N. cretacea, Charentia cuvillieri, Praechrysalidina ing T. sagittaria, V. scarsellai, and very rarely S. infracretacea, Pseudonummoloculina aurigerica minuta. Throughout the Valanginian, trocholinids and ‘Involutina’ sp. of the T. alpina–elongata group, P. lituus and Prae­ Discussion: During the earliest Barremian, there were chrysalidina sp. are still present. T. delphinensis no significant changes in the foraminiferal assem­ was determined exclusively in Valanginian depos­ blage compared to the Late Hauterivian. This its. Although Valanginian deposits contain the larg­ marks the beginning of the stratigraphic range of est number of index fossils of all the Neocomian some species which are typical of the Early Creta­ units, more detailed stratigraphic subdivision of Val­ ceous, as well as S. cretacea, R. robusta, B. texti­ anginian deposits is not presently possible since all larioides and V. wintereri, which were documented these index forms, together with numerous calcare­ in the Barremian of the Western part of the AdCP ous algae (Dasycladales), have stratigraphic ranges by CVETKO TEŠOVIĆ (2000). C. cuvillieri also spanning the whole of the Valanginian. occurs in these deposits, although it has not been observed in the Lower Barremian deposits. 2.3.1.3. Hauterivian Near the end of the Barremian, orbitolinids occur as Foraminiferal assemblage: the first genuine larger foraminifera on the AdCP. (a) Early Hauterivian: T. gr. alpina–elongata, T. sagit­ The oldest forms belong to R. giganteus observed taria, M. bulgarica, C. cuvillieri, S. minuta, M. sale­ along the northern platform margin between Banja vensis, H. joukowskyi, M. favrei, V. tenuis, V. campo­ Luka and Kupres, and P. lenticularis which is saurii, V. scarsellai, Novalesia distorta, Nezzazata widely distributed across the platform. VELIĆ & germanica and Praechrysalidina sp. SOKAČ (1978a) proposed that the stratigraphic (b) Late Hauterivian: T. gr. alpina–elongata, T. sagit­ range of P. lenticularis began, before the occurrence taria, M. bulgarica, C. cuvillieri, S. minuta, N. ger­ of the Early Aptian praeorbitolinids, indicating the manica, Vercorsella laurentii, V. scarsellai, Debari­ Late Barremian. This approach was later confirmed na hahounerensis, Novalesia producta, N. distorta, by GUŠIĆ (1981), who compared the dimensions of Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 15

the proloculuses and embrional apparatuses of this Early Aptian and ends in the Middle Cenomanian. species, and documented the gradual increase in This was confirmed on the AdCP by VELIĆ & these dimensions from older, Barremian, to younger, SOKAČ (1978a) and VELIĆ (1988), where the Aptian palorbitolinids. Furthermore, in the absence Aptian forms occur in the stratigraphic succession of of other criteria within the palorbitolinid limestones P. cormyi – P. wienandsi – M. lotzei (Early Aptian) of the AdCP, the size of the proloculus can even be – M. parva – M. texana (Late Aptian). used to determine the boundary between the Bar­ The Early Aptian foraminiferal assemblage also remian and the Aptian (GUŠIĆ, 1981). contains other index fossils, including V. murgensis, In addition, several other foraminifera of a wider S. briacensis, C. decipiens, D. filipescui, N. fribur­ stratigraphic range have been determined – Trocholi­ gensis, N. aptiensis, P. globosa and P. conica. Oth­ na sagittaria, T. molesta, T. odukpaniensis, Ver­ ers, characterized by a wider stratigraphic range corsella arenata, Pseudonummoloculina aurigerica throughout the Early Cretaceous include P. auriger­ and P. infracretacea, as well as all the taxa found ica, N. broennimanni and N. cretacea, as well as an in the Lower Barremian deposits. At the very end of alveolinid form very close to A. reicheli, which has the Barremian Derventina filipescui, Neotrocholina a stratigraphic range through the Late Aptian and friburgensis, N. aptiensis, Nautiloculina broenni­ Early Albian. Furthermore, numerous specimens of manni and N. cretacea occur before the first appear­ involutinids also occur, but due to intense recrystal­ ance of the praeorbitolinids indicating the beginning lization, can only be determined as ‘Involutina’ sp. of the Aptian. The evolutionary trend of genuine orbitolinids is 2.3.1.5. Aptian continued in the Late Aptian by the new species M. parva and M. texana. Among other orbitolinids in Foraminiferal assemblage: the youngermost Aptian deposits O. aquitanica was (a) Early Aptian: P. lenticularis, R. giganteus, Paleo­ also determined. dictyoconus actinostoma, Praeorbitolina cormyi, P. The most abundant foraminifera are of somewhat wienandsi, Mesorbitolina lotzei, Voloshinoides mur­ wider stratigraphic ranges, e.g. D. hahounerensis, gensis, Sabaudia briacensis, S. minuta, Choffatella M. bulgarica, P. infracretacea, S. minuta, S. capi­ decipiens, D. filipescui, N. friburgensis, N. aptiensis, tata, S. auruncensis, V. laurentii, V. scarsellai and T. molesta, T. odukpaniensis, Pfenderina globosa, V. arenata, including the first occurrences of H. Pseudolituonella conica, D. hahounerensis, M. bul­ globosus (or Trochaminoides coronus according garica, P. infracretacea, V. laurentii, V. scarsellai, to CHIOCCHINI et al., 1994) and N. simplex. A. V. arenata, N. producta, N. distorta, N. cornuco­ reicheli, which can also be observed locally in mes­ pia, N. broennimanni, N. cretacea, C. cuvillieri, P. orbitolinid tempestites, reaches its maximum in the aurigerica, Glomospira urgoniana, G. watersi, S. Aptian. Smaller forms, together with some still not cretacea, R. robusta, Archaealveolina sp. and ‘Invo­ completely determined forms, cited by CVETKO lutina’ sp. TEŠOVIĆ (2000) include G. urgoniana, G. watersi, (b) Late Aptian: Mesorbitolina parva, M. texana, N. isabellae, S. cretacea and R. robusta. In the Late N. aptiensis, D. hahounerensis, M. bulgarica, P. Aptian, species of Novalesia disappeared except for infracretacea, Sabaudia auruncensis, S. capitata, N. angulosa. S. minuta, V. laurentii, V. scarsellai, V. arenata, Novalesia angulosa, N. producta, N. distorta, N. 2.3.1.6. Albian cornucopia, Haplophragmoides globosus, Archal­ Foraminiferal assemblage: veolina reicheli, P. aurigerica, Nezzazata isabellae, (a) Early Albian: Mesorbitolina pervia, M. subconca­ N. simplex, G. urgoniana, G. watersi, S. cretacea, R. va, M. parva, M. texana, Paracoskinolina sunni­ robusta and Orbitolinopsis aquitanica. ladensis, Cuneolina pavonia, C. parva, C. sliteri, Discussion: Occurrence of the first orbitolinids near Sabaudia dinapolii, S. minuta, S. auruncensis, S. the end of the Barremian marks the beginning of the capitata, Nezzazatinella picardi, Pseudonummolo­ dominance of these larger foraminifera on the AdCP culina heimi, P. aurigerica, S. cretacea, R. robusta, which lasted until the Middle Cenomanian. One of G. urgoniana, N. isabellae, N. simplex, D. hahoune­ the most frequent and widely distributed species is rensis, H. globosus, M. bulgarica, P. infracretacea, P. lenticularis, which reaches its maximum during A. reicheli, V. laurentii, V. scarsellai, V. arenata, N. the Early Aptian. In some localities it is so abundant angulosa and Orbitolinopsis sp. cf. O. aquitanica. that it has lithogenetic value as some tempestite lay­ (b) Late Albian: ‘Valdanchella’ dercourti, Neoiraquia ers are composed almost exclusively of tests of this insolita, N. convexa, Simplorbitolina broennimanni, foraminifera (see cover page of this issue). In the Paracoskinolina fleuryi, P. sunnilandensis, C. pavo­ undisturbed succession of Lower Aptian deposits, nia, C. parva, Scandonea pumila, S. phoenissa, N. the first part of the evolutionary trend of orbitolinids simplex, N. isabellae, N. gyra, ?Trochospira avni­ can be traced, which, according to SCHROEDER melechi, ?Biconcava bentori, N. picardi, P. heimi, (1962, 1964a, b, c, 1975, 1979), commences in the P. aurigerica, S. cretacea, S. minuta, S. auruncen­ 16 Geologia Croatica 60/1

sis, V. laurentii, V. scarsellai, N. angulosa, G. urgo­ tional level to the Cenomanian, N. simplex occurs niana, P. infracretacea, Protochrysalidina elongata, together with the first appearance of other nezzaza­ Ovalveolina maccagnoae and ?Sellialveolina viallii. tids including N. gyra, Trochospira avnimelechi and Discussion: Albian deposits on the AdCP contain rich Biconcava bentori, which reached their maximum in foraminiferal assemblages dominated by different the Cenomanian. miliolids. Among other larger foraminifera, the most 2.3.2. Late Cretaceous important are the orbitolinids because of both their stratigraphic value and abundance in certain levels. General characteristics: Foraminiferal assemblag­ The most important event for determination of the es in the AdCP Upper Cretaceous deposits are not boundary between Upper Aptian and Lower Albian regularly distributed (Fig. 6). If the frequency and deposits is the first occurrence of species of cune­ abundance of all taxa within specific stages were olina, including C. pavonia, C. parva and C. sliteri analysed statistically, Cenomanian foraminiferal which are certain indicators of the beginning of assemblages would probably be the richest in the the Albian. Albian deposits also contain numerous entire succession of shallow-marine carbonates. In foraminifera known from Barremian (or even ear­ contrast, the Turonian is characterized by an evi­ lier) and Aptian sediments: P. aurigerica, D. hahou­ dent decrease in the diversity and abundance of nerensis, M. bulgarica, P. infracretacea, A. reicheli foraminiferal assemblages, as a consequence of the as well as all previously mentioned species of prominent global sea-level rise in the youngermost Sabaudia and Vercorsella. During the Early Albian, Cenomanian and oldermost Turonian, first recog­ the widely distributed species P. heimi and N. picar­ nised in the AdCP by GUŠIĆ & JELASKA (1990, 1993). This sea-level rise resulted in the temporary di, occur for the first time. These, together with the drowning of a major part of the AdCP (see JELAS­ cuneolinids, continue to the Late Cretaceous in the KA et al., 1994; VLAHOVIĆ et al., 1994, 2003, area of the AdCP. Of these, P. aurigerica is the most 2005; TIŠLJAR et al., 2002; VELIĆ et al., 2002). common. According to CVETKO TEŠOVIĆ (2000), Another earlier regionally important sea-level rise Lower Albian carbonates also contain smaller was determined within the youngermost Albian and foraminifera, including G. urgoniana, N. isabellae, oldest Cenomanian deposits on the island of Cres S. cretacea and R. robusta. Within the orbitolinid (HUSINEC et al., 2000; KORBAR et al., 2001) and assemblage, the mesorbitolinids M. parva and espe­ some neighbouring areas. After restoration of the cially M. texana are important. These, together with shallow marine environments and ecosystems in the P. lenticularis, constitute the most abundant Early Late Turonian, a gradual recolonization of benthic Cretaceous larger foraminifera on the AdCP. M. foraminifera took place, but this was not character­ texana is also frequently lithogenic, especially along ized by such a richness of taxa and specimens as the NE platform margin from Banja Luka towards during the Cenomanian. However, the next resur­ Jajce and Kupres. The largest, most advanced mes­ gence in the foraminiferal assemblages took place in orbitolinids are M. pervia and M. subconcava, and the Late Santonian and, especially, the Campanian. these are also index fossils for the Albian. In the Maastrichtian assemblages were characterized by a area of the AdCP, the latter species represents the relative decrease in the number of taxa, which is not final member of the mesorbitolinid part of the evo­ surprising considering that by this time the platform lutionary trend of the ‘genuine’ orbitolinids (after had already largely disintegrated and most of it was SCHROEDER, 1975). These deposits rarely con­ emergent, resulting in a significant decrease of the tain specifically undeterminable (mostly micritized) shallow-marine areas. specimens of Orbitolinopsis sp., as well as P. sun­ nilandensis. 2.3.2.1. Cenomanian Upper Albian deposits are characterized by two Foraminiferal assemblage: Foraminifera determined zones with orbitolinids: the first one in the older within the Cenomanian deposits of the AdCP can part, and the second one in uppermost part of the generally be divided into two groups. Vraconian, below the Cenomanian. In the first orbit­ The first group comprises forms which originated in olinid-rich zone, ‘V.’ dercourti, N. insolita, S. broen­ the Early Cretaceous or the Cenomanian, and some nimanni and P. fleuryi were determined, while the of them continue into younger periods. Forms from second one contains only ‘V.’ dercourti and N. con­ the Early Cretaceous include Cuneolina pavonia, vexa. During the Late Albian S. pumila and S. phoe­ C. parva, Pseudonummoloculina heimi, Spirolo­ nissa occur together with other foraminifera, and in culina cretacea, Nezzazatinella picardi, Nezzazata the youngermost part there is also P. elongata and simplex, N. gyra, Trochospira avnimelechi, Bicon­ O. maccagnoae. From the Early Albian some forms cava bentori, Charentia cuvillieri and Scandonea with wider stratigraphic ranges continued, of which pumila. Since Cenomanian they were accompanied the most important include the pseudonummolo­ by Biplanata peneropliformis, Nezzazata conica and culinids, cuneolinids and sabaudias cited above. In Rotalia mesogeensis, as well as Cuneolina conica, the terminal part of the Late Albian, and the transi­ Murgeina apula and Spiroloculina robusta, which Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 17

Cenomanian Turonian Campanian Coniacian Santonian Maastrichtian

Pseudorhipidionina murgiana

Fig. 6 Strati- graphic ranges of the Late Cre- taceous benthic foraminif- era on the AdCP. 18 Geologia Croatica 60/1

can be found also in younger Upper Cretaceous as well as the exact stratigraphic range of this form, deposits. remains open. The second group comprises foraminifera that are 2.3.2.2. Turonian either index fossils for some parts of the Cenoma­ Foraminiferal assemblage: nian, or were found only in some levels of the Ceno­ manian of the AdCP although they are of a wider (a) Early Turonian: P. heimi, C. pavonia, C. parva, S. stratigraphic range in the Cretaceous. In this group, cretacea, N. simplex, N. gyra, N. conica, T. avnime­ the following forms are characteristic: lechi, B. bentori, N. picardi and M. apula. (a) Early Cenomanian: Ovalveolina maccagnoae, O. (b) Late Turonian: P. heimi, Cuneolina compressa, C. crassa, ?Sellialveolina viallii, Praealveolina iberi­ pavonia, C. parva, S. cretacea, N. simplex, N. gyra, ca, P. simplex, Cisalveolina lehneri, Orbitolina gr. N. conica, T. avnimelechi, B. bentori, N. picardi, M. sefini–concava, Conicorbitolina conica, C. cuvil­ apula, Pseudocyclammina sphaeroidea, Monchar­ lieri, C. gr. corbarica–conica, Pseudocyclammina montia apenninica and M. compressa. rugosa, Trocholina arabica and Pastrikella biplana. Discussion: The Early Turonian is characterized by (b) Middle Cenomanian: Conicorbitolina conica, C. platform drowning, resulting in impoverishment of gr. corbarica–conica, C. cuvillieri, O. gr. sefini–con­ the shallow-marine benthic foraminifera assemblag­ cava, P. iberica, C. lehneri, O. crassa, Chrysalidina es, which are mostly composed of taxa with broad gradata, Nummoloculina? regularis, Pseudolituo­ stratigraphic ranges. Restoration of assemblages in nella reicheli, Pseudorhapydionina dubia, Merlin­ subtidal, lagoonal and peritidal environments took gina cretacea, Peneroplis turonicus, Pastrikella bal­ place at the end of the Early and during the Late canica and P. biplana. Turonian. The most frequent forms were those with their maximum abundance in the Cenomanian, (c) Late Cenomanian: C. gradata, N.? regularis, P. including C. pavonia, C. parva, C. conica, P. heimi, reicheli, P. balcanica, P. dubia, Cisalveolina fraasi, S. cretacea, N. simplex, N. gyra, N. conica, T. avn­ Vidalina radoicicae, V. hispanica, Murgeina ceno­ imelechi, B. bentori, N. picardi and M. apula. New mana, Pseudorhapydionina laurinensis, Pseudorhi­ species like P. sphaeroidea, M. compressa and C. pidionina casertana, P. murgiana, M. cretacea, compressa occurred in the Late Turonian. Peneroplis parvus and P. turonicus. 2.3.2.3. Coniacian Discussion: The maximum abundances of P. heimi, C. pavonia, C. parva and nezzazatids, some of which Foraminiferal assemblage: C. pavonia, C. parva, C. previously occur in the Late Albian (VELIĆ & conica, C. compressa, Dicyclina schlumbergeri, P. SOKAČ, 1979), and some in the Early Cenomanian, heimi, N. picardi, N. simplex, N. gyra, N. conica, B. are particularly important in the first group. Among bentori, T. avnimelechi, M. apula, M. compressa, M. the second group, the alveolinids O. maccagnoae, apenninica, P. sphaeroidea, Scandonea samnitica, O. crassa, S. viallii, C. lehneri, P. simplex and P. Accordiella conica and Idalina antiqua. iberica are stratigraphically very important for the Discussion: The Coniacian foraminiferal assemblage Early Cenomanian. Index fossils for the Middle also does not contain index forms, since most of the Cenomanian alone are absent, except for P. biplana determined taxa first appeared in Turonian or older which is known as the Early to Middle Cenomanian deposits. The Coniacian is characterized by the form (CHERCHI & SCHROEDER, 1980, 1985), first occurrence of D. schlumbergeri, S. samnitica, and is frequent in Middle Cenomanian beds with P. A. conica and I. antiqua, the stratigraphic range of balcanica on the AdCP. Reliable definition of this which extends into younger Cretaceous stages. part of the Cenomanian is based on the overlapping 2.3.2.4. Santonian stratigraphic ranges of the cited orbitolinids, which ended their stratigraphic range near the end of the Foraminiferal assemblage: Middle Cenomanian, together with foraminifera of (a) Early Santonian: C. pavonia, C. parva, C. conica, characteristic Middle to Late Cenomanian strati­ P. heimi, S. samnitica, P. sphaeroidea, M. com­ graphic range, including C. gradata, N.? regularis, pressa, M. apenninica, N. simplex, N. gyra, B. ben­ P. reicheli, P. balcanica and P. dubia. Index fossils tori, T. avnimelechi, N. picardi, D. schlumbergeri, of the Late Cenomanian are C. fraasi, V. radoicicae, M. apula, A. conica, I. antiqua, Keramosphaerina V. hispanica, P. laurinensis, P. casertana, P. mur­ ?sarda, K. tergestina, Pseudorhapydionina mediter­ giana and P. parvus. ranea and Murgella lata. POLŠAK et al. (1982) cited S. minuta as being of (b) Late Santonian: all species from the Early San­ Early Cenomanian age, although they commented tonian and new forms: Reticulinella fleuryi, Dicty­ that their Early Cenomanian foraminiferal assem­ opsella kiliani, Dictyopselloides cuvillieri, Spirolina blage does not differ from that of the Late Albian. cretacea, Minouxia lobata, Broekinella neumannae, Therefore, the question of the position of the exact Archiacina munieri, Pseudolituonella marie, Pseu­ boundary between the Albian and the Cenomanian, docyclammina massiliensis, Rotorbinella scarsellai, Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 19

Stensiöina surrentina, Raadshovenia salentina and which is in close agreement with the opinion of Orbitoides gr. hottingeri. GUŠIĆ & JELASKA (1990). Discussion: In the Early Santonian, some species first 2.3.2.5. Campanian appear that continue into the Early Campanian Foraminiferal assemblage: including, for example, K. tergestina and M. lata, or into the Maastrichtian for example P. mediter­ (a) Early Campanian: C. pavonia, C. parva, C. coni­ ranea. The first occurrence of the index species M. ca, P. heimi, D. schlumbergeri, N. simplex, N. gyra, lata in the Santonian is very important (after GUŠIĆ T. avnimelechi, B. bentori, N. picardi, P. sphaeroi­ & JELASKA, 1990), together with P. mediterranea, dea, P. massiliensis, M. apula, M. apenninica, M. K. tergestina, R. fleuryi, S. cretacea, D. kiliani, D. compressa, A. conica, P. mediterranea, S. samniti­ cuvillieri, R. scarsellai, S. surrentina and O. gr. hot­ ca, K. tergestina, M. lata, S. cretacea, I. antiqua, M. tingeri, which are also index fossils for the Early lobata, Calveziconus lecalvezae, B. neumannae, A. Campanian. The beginning of the stratigraphic range munieri, P. marie, Reticulinella reicheli, R. fleuryi, of K. tergestina in the Early Santonian is character­ D. kiliani, D. cuvillieri, R. scarsellai, S. surrentina, ized by small specimens, among which forms indi­ Orbitoides gr. hottingeri, O. douvillei, O. tissoti and cating K. sarda were observed. The largest speci­ Praesiderolites douvillei. mens of K. tergestina occur exclusively in upper­ (b) Late Campanian: C. pavonia, C. parva, N. sim­ most Santonian and lowermost Campanian deposits. plex, N. gyra, T. avnimelechi, N. picardi, M. apula, B. neumannae and A. munieri occur for the first time M. apenninica, M. compressa, D. schlumbergeri, A. in the Late Santonian–Early Campanian foraminif­ conica, R. scarsellai, P. mediterranea, S. samnitica, eral assemblage of the AdCP. POLŠAK et al. (1982) P. sphaeroidea, P. massiliensis, R. reicheli, R. fleur­ and VENTURINI (2005) also cited R. salentina yi, D. kiliani, D. cuvillieri, I. antiqua, C. lecalvezae, from the Santonian–Campanian assemblages. Nummofallotia cretacea, Praesiderolites douvillei, Due to the pronounced similarity of Late Santonian Orbitoides douvillei, O. tissoti, O. media and Pseu­ and Early Campanian assemblages it is not easy to dosiderolites vidali. reliably establish the boundary between these two Discussion: There is little difference between the stages. Another, relatively new problem, is connect­ foraminiferal assemblages of the Late Santonian and ed to the general division of younger Upper Creta­ the Early Campanian, when new forms Orbitoides ceous deposits of the AdCP. Strontium isotope anal­ douvillei, O. tissoti and Praesiderolites douvillei ysis performed by STEUBER et al. (2005) resulted appeared. Most of the taxa continued to the Late in revision of the stratigraphic ranges of the majority Campanian, where they are accompanied by new of larger benthic foraminifera index forms, and con­ forms – R. reicheli, N. cretacea and Pseudosidero­ sequently the stratigraphy of the Coniacian–Maas­ lites vidali and Orbitoides media, which is also an trichtian deposits of the island of Brač as determined index fossil for the Maastrichtian. by GUŠIĆ & JELASKA (1990). According to this 2.3.2.6. Maastrichtian investigation M. lata occurred in the Late Coniacian, P. mediterranea and A. conica had a stratigraphic Foraminiferal assemblage: D. schlumbergeri, Cuneo­ range from the Late Coniacian to the Early Campa­ lina ketini, C. pavonia, N. picardi, M. apula, R. nian, and K. tergestina and Calveziconus lecalvezae scarsellai, Murciella cuvillieri, M. renzi, Rhapydio­ occur in the Early Campanian (their first occurrence nina liburnica, Laffitteina mengaudi, Fleuryana would mark the beginning of the Campanian). How­ adriatica, O. media, Siderolites calcitrapoides, ever, VENTURINI (2005) published results of the Omphalocyclus macroporus, Lepidorbitoides minor strontium isotope analysis from Santonian–Campa­ and Hellenocyclina beotica. nian deposits of the Slovenian–Trieste Karst region, Discussion: The foraminiferal assemblage of the Maas­ according to which K. tergestina and M. lata occur trichtian comprises several species which continued in Upper Santonian, and C. lecalvezae in Lower from earlier stages, e.g. C. pavonia, D. schlumber­ Campanian limestones. On the basis of Sr isotopes geri, N. picardi and O. media. Exclusively Maas­ MARITAN et al. (2003) proposed Santonian age for trichtian index fossils are M. cuvillieri, M. renzi, K. tergestina and Santonian–Lower Campanian age R. liburnica, C. ketini, F. adriatica, L. mengaudi, for M. lata in the Trieste Karst area. Furthermore, S. calcitrapoides, O. macroporus, L. minor and H. FRIJIA & PARENTE (2007) suggest on the basis beotica. of strontium and carbon isotope stratigraphy in the Upper Campanian and Maastrichtian deposits are southern Apennines, that the first occurence of M. not very common on the AdCP. They occur only in lata occurs in the Early Coniacian and K. tergestina its SW marginal areas – in Divača area (SW Slov­ in the Early Campanian. enia), near Trogir and Dubrovnik, and on the islands Here, on the AdCP, the stratigraphic ranges of M. of Brač and Hvar (S, SE Croatia) (MARKOVIĆ, lata and K. tergestina are proposed as extending 1975; GUŠIĆ & JELASKA, 1990; JURKOVŠEK from the Early Santonian to the Early Campanian, et al., 1996; MARINČIĆ, 1997). Such a restricted 20 Geologia Croatica 60/1

distribution is the consequence of a decreased depo­ stratigraphic sequences and regional correlation and sitional area due to the intense disintegration of the synthesis. Recent data can only be found in relation to platform that began in the Cenomanian, gradual Upper Triassic deposits, and therefore biostratigraphic emergence of its central parts and the culmination zonation can be attempted from this level onwards. The of uplift in the Maastrichtian. Depositional environ­ previously acknowledged subdivision of the Upper Tri­ ments favourable for the development of shallow- assic deposits of Karlovac area (Central Croatia) is that marine benthic communities were also frequently published by GRGASOVIĆ (1997). However, results of influenced by open marine conditions. At the NE older investigations from different parts of the platform margin of the AdCP (DRAGIČEVIĆ & VELIĆ, are also used for biostratigraphic zonation presented in 2002), and in the intraplatform troughs formed with­ this paper. in the central part of the platform (W Bosnia and Jurassic deposits have interested biostratigraphers Herzegovina, S Croatia – VELIĆ et al., 2002), clas­ for a long time, as witnessed by a long list of papers tic–carbonate deposits accumulated with predomi­ published since the 1960’s. Authors usually followed nantly planktonic foraminiferal assemblages, but a scheme proposed for the Apennines by SARTONI & also rich assemblages of benthic foraminifera, espe­ CRESCENTI (1962). For the AdCP, the first such sub­ cially orbitoids and siderolitids. The SW platform division was published by FARINACCI & RADOIČIĆ margin is presently located beneath the Adriatic Sea, (1964), followed by RADOIČIĆ (1966), NIKLER & mostly covered by Tertiary and Quaternary deposits. SOKAČ (1968), GUŠIĆ (1969b), GUŠIĆ et al. (1971), However, slope deposits of Maastrichtian age were VELIĆ & SOKAČ (1974 – only for Late Jurassic), determined in off-shore exploratory wells, e.g. W VELIĆ (1977), VELIĆ & SOKAČ (1978a) and and SW of Istria, containing orbitoid and siderolitid SOKAČ & VELIĆ (1979a). SEPTFONTAINE et al. assemblages (VESELI, 1999). (1991) presented a general biostratigraphic zonation of Jurassic imperforate foraminifera on Tethyan carbonate 3. BIOSTRATIGRAPHY platforms of the Mediterranean, which included the area of the AdCP. VELIĆ et al. (1995) biostratigraphically subdivided younger Middle and Upper Jurassic deposits Mesozoic carbonate deposits of the Adriatic Carbon­ of Western Istria. ate Platform are, although partially, relatively well biostratigraphically investigated. Most of the studies Papers dealing with Cretaceous microfossil assem­ were restricted to relatively small areas. However, the blages are very numerous. One of the first papers illus­ work of RADOIČIĆ (1966) is more of an attempt at the trating Cretaceous assemblages is that by RADOIČIĆ regional biostratigraphic analysis of a major part of the (1960). Besides stratigraphic relationships more and platform. An integrated biostratigraphic subdivision of more attention was gradually given to biostratigraphic the entire platform is presented here from the Upper Tri­ subdivision. Some of the aforementioned papers deal­ assic to the end of the Cretaceous, based also on results ing with Jurassic biostratigraphy also included, either of all available former investigations. partly or completely, Cretaceous deposits, e.g. FARI­ Lower and Middle Triassic deposits have not been NACCI & RADOIČIĆ (1964), VELIĆ (1977), VELIĆ systematically stratigraphically investigated, so there & SOKAČ (1978a, b) and SOKAČ & VELIĆ (1979a). are no reliable recent data that are needed for their Lower Cretaceous microfossil assemblages and/or bio­ stratigraphic zonation. zonations were also investigated, including those by VELIĆ (1973), SOKAČ & VELIĆ (1978, 1979a, b), It should be noted that benthic foraminifera on the SOKAČ et al. (1978), VELIĆ & SOKAČ (1978a, b, AdCP were most frequently biostratigraphically ana­ 1979, 1980, 1983), VELIĆ et al. (1979) and FUČEK lyzed together with calcareous algae. Therefore, this et al. (1995). The most detailed study of foraminiferal paper represents the first zonation based exclusively biostratigraphy of Lower Cretaceous deposits is that by on benthic foraminifera that can be used platformwide. VELIĆ (1988). CVETKO TEŠOVIĆ (2000) analyzed It will be a reliable starting point not only for more foraminiferal assemblages of the Barremian, Aptian detailed biozonation, but also for stratigraphic analysis and Albian, while the orbitolinid biostratigraphy of the and determination. so-called Mid-Cretaceous (stratigraphic range Early Aptian–Middle Cenomanian), was presented by HUSI­ 3.1. Previous investigations NEC et al. (2000). ARNAUD-VANNEAU et al. (1991) Triassic deposits of the Karst Dinarides have been presented stratigraphic ranges of Early Cretaceous ben­ relatively poorly investigated, and there are no data thic foraminifera of the AdCP and neighbouring plat­ on detailed recent studies. Palaeontological papers on forms, particularly the Apenninic, Apulian and Hellenic. Early and Middle Triassic foraminiferal assemblages VELIĆ et al. (1995) presented microfossil assemblages are scarce, and several decades old. Furthermore, they and determined biozones within a major part of the usually refer to specific localities dealing with narrow Lower Cretaceous deposits of Western Istria. stratigraphic ranges, specific taxa, etc., generally lack­ Upper Cretaceous deposits of the AdCP are not so ing reliable data based on the studies of continuous well biostratigraphically subdivided as Jurassic and Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 21

Lower Cretaceous deposits. Foraminiferal assemblag­ tial for biostratigraphic subdivision, because it is based es, mostly from the Adriatic coastal area and islands, mostly on the order of occurrence, i.e. stratigraphic suc­ were studied by MAMUŽIĆ & GRIMANI (1970), cession of individual taxa and assemblages. MAMUŽIĆ et al. (1976, 1980, 1981a, b, 1982, 1983, 1984), POLŠAK et al. (1982), GUŠIĆ et al. (1988), 3.2.1. Late Triassic FUČEK et al. (1990) and others. VELIĆ & VLAHOVIĆ GRGASOVIĆ (1997) established three biozones within (1994) published detailed biostratigraphic zonation the Upper Triassic deposits of Žumberak (NW Croatia): of the Cenomanian deposits. Regionally important (1) Clypeina besici taxon-range zone in the Carnian, results on the biostratigraphy of benthic foraminifera (2) Aulotortus sinuosus interval zone of approximately on the AdCP are presented in monographs published by Norian age, and (3) Triasina hantkeni taxon-range zone GUŠIĆ & JELASKA (1990) and JURKOVŠEK et al. in the Rhaetian. This subdivision is acceptable and usa­ (1996), as well as in papers by CVETKO TEŠOVIĆ et ble throughout the AdCP area. However, zonation based al. (2001) and KORBAR et al. (2001), using biozona­ exclusively on benthic foraminifera will produce zones tion after FLEURY (1980). which are not completely correlatable with those based on dasyclad algae. According to the stratigraphic ranges 3.2. Biostratigraphic zonation and subdivision of Late Triassic foraminifera the following superim­ posed Upper Triassic foraminiferal biozones could be From the introduction above it is clear that the distri­ established: (1) Lamelliconus procerus taxon-range bution of biostratigraphic investigations of benthic zone, (2) Lamelliconus procerus–Triasina hantkeni foraminifera over the entire AdCP area is uneven. There interval zone, and (3) Triasina hantkeni taxon-range is more or less continuous coverage over the Croatian part of the platform, but papers dealing with neighbour­ zone. ing areas are infrequent. The least investigated areas are 3.2.1.1. Lamelliconus procerus taxon-range zone Bosnia and Herzegovina and Montenegro, so mostly older papers were relied on here. Boundaries: stratigraphic range of L. procerus. Since Lower and Middle Triassic rocks were rela­ Index and important taxa: L. procerus, L. multispirus, tively poorly investigated, biostratigraphic zonation Aulotortus friedli, A. sinuosus, A. praegaschei and cannot be based exclusively on benthic foraminifera. A. tenuis. Detailed investigation of Early Triassic and Anisian Stratigraphic range: Carnian. foraminiferal assemblages reveals the presence of index Discussion: Within the cited assemblage only L. procer­ taxa of narrow stratigraphic range. However, Ladin­ us and L. multispirus are index fossils, while spe­ ian carbonate deposits do not contain index forms of cies of Aulotortus have wider stratigraphic ranges foraminifera, and biostratigraphic zonations of Middle through the Late Triassic. Triassic deposits are based mainly on dasyclad algae. The first such zonation was published by GRGASOVIĆ 3.2.1.2. Lamelliconus procerus–Triasina hantkeni (2003). Therefore, in this paper biozonation of the shal­ interval zone low-marine carbonate deposits of the AdCP will be lim­ Boundaries: from the last occurrence of L. procerus to ited to the stratigraphic range Late Triassic–end Creta­ the first occurrence of T. hantkeni. ceous. Important taxa: Aulotortus friedli, A. sinuosus, A. prae­ The biozones and accompanying tables only list gaschei, A. tenuis, A. tumidus, Trocholina crassa index fossils and the more important taxa for the and T. acuta. foraminiferal assemblages in order to avoid repetition Stratigraphic range: Late Carnian–Early Rhaetian. of all determined taxa previously cited in the descrip­ tion of assemblages. There are several cases when one Discussion: Within well studied Upper Triassic depos­ biozone was determined as e.g. lineage zone or interval its of Karlovac area, GRGASOVIĆ (1997) sepa­ zone or partial-range zone. These zones were defined rated the A. sinuosus interval zone of Norian strati­ as being between the first occurrences of two index or graphic range (from the last occurrence of Carnian more important foraminiferal taxa. Priority was given to algae Clypeina besici to the first occurrence of the lineage zones, followed by partial-range zones named Rhaetian foraminifera T. hantkeni). If only ben­ after species with narrow chrono- and biostratigraphical thic foraminifera are considered the zone would be range, and interval zones if the older taxa is of a wider somewhat wider, but would not contain any index stratigraphic range. For each biozone, the chronostrati­ forms, since the stratigraphic ranges of cited species graphic range is presented, to enable correlation of the of Aulotortus commenced in the Anisian. Separation studied deposits. However, there are cases when new of this, as well as the previous zone in the area of investigations significantly changed the chronostrati­ the AdCP can only be made in areas where Upper graphic ranges of some important foraminiferal taxa, Triassic deposits are not dolomitized, as discussed for example the strontium isotope analysis of the Upper in section 2.1.3. Cretaceous deposits of the island of Brač by STEUBER et al. (2005). Such changes are important, but not essen­ 22 Geologia Croatica 60/1

Hett. Sinemurian Pliensbachian Toarcian

Fig. 7 Biostratigraphic subdivision of the Lower Jurassic deposits of the AdCP. Legend: 1) Triasina hantkeni–Mesoendothyra sp. interval zone; 2) Mesoendothyra sp. lineage zone; 3) Lituosepta recoarensis lineage zone; 4) Orbitopsella primaeva lineage zone: 4.1) Orbit- opsella primaeva–Lituosepta recoarensis concurrent-range subzone, 4.2) Lituosepta recoarensis–Orbitopsella praecursor interval sub- zone; 5) Orbitopsella praecursor taxon-range zone: 5.1) Orbitopsella praecursor–O. primaeva concurrent-range subzone, 5.2) Orbit- opsella praecursor abundance subzone; 6) Orbitopsella praecursor–Pseudocyclammina liassica interval zone; 7) Pseudocyclammina liassica taxon-range zone; 8) Pseudocyclammina liassica–Gutnicella cayeuxi interval zone.

3.2.1.3. Triasina hantkeni taxon-range zone, the AdCP, where more than 90% of genera and species GRGASOVIĆ (1997) are common and occur in the same stratigraphic ranges. Boundaries: stratigraphic range of T. hantkeni. Therefore, the relationships between different taxa are Index and important taxa: T. hantkeni, Auloconus also the same as in the Lower Jurassic of the High Atlas permodiscoides, Aulotortus friedli, A. sinuosus, A. (compare Fig. 2 with Figs. 3 & 7). tenuis, A. tumidus, A. pokornyi, Trocholina crassa 3.2.2.1. Triasina hantkeni–Mesoendothyra sp. and T. acuta. interval zone Stratigraphic range: Rhaetian. Boundaries: from the last occurrence of T. hantkeni to Discussion: Foraminiferal assemblages of this zone the first occurrence of Mesoendothyra sp. cannot be traced through the entire sequence of Index and important taxa: not applicable, only small Rhaetian deposits, due to extensive late-diagenetic valvulinids and lituolids without significant strati­ dolomites, while non-dolomitized limestones can be graphic value, e.g Duotaxis metula, Amijiella amiji, found in alternation with dolomites in certain levels Siphovalvulina variabilis and Everticyclammina sp. or as lenses and interbeds within dolomites. Index – ?E. praevirguliana. fossils for this zone are T. hantkeni and A. permo­ Stratigraphic range: Late Rhaetian–Early Sinemurian. discoides. Discussion: During the Late Rhaetian and Hettang­ 3.2.2. Early Jurassic ian, environmental conditions were very unfavour­ able for foraminifera, due to extensive coverage of Due to the evident decrease of fossils during the earli­ the platform by tidal flats or very shallow subtidal est Jurassic, biostratigraphic studies and subdivisions seas. Rhaetian carbonate deposits are represented by based exclusively on benthic foraminifera have not pre­ the alternation of early- and late-diagenetic stroma­ viously been attempted. Authors dealing with Jurassic tolitic dolomites (Hauptdolomite – Main Dolomite biostratigraphy only defined the subzone Orbitopsella Formation). The Hettangian deposits consist of the praecursor in the middle part of the Early Jurassic with­ alternation of limestones and late-diagenetic dolo­ in the Palaeodasycladus mediterraneus zone (NIKLER mites with rare intercalations of early-diagenetic & SOKAČ, 1968; GUŠIĆ, 1969b; GUŠIĆ et al., 1971; dolomites. In Hettangian environments dasyclad VELIĆ, 1977; VELIĆ & SOKAČ, 1978a). algae sporadically flourished. Although all the stud­ Biozonation within the Lower Jurassic deposits is ied sections contain benthic foraminifera there are presented here (Fig. 7). It is based upon results of bio­ no index species, which only occur later in the Early stratigraphic investigations of Early Jurassic lituolids Sinemurian. Therefore, this part of the sequence is made by SEPTFONTAINE (1984) in shallow-marine, marked as an interval zone. mostly carbonate sequences of Western Mediterranean, especially in the Lower Jurassic of the High Atlas in 3.2.2.2. Mesoendothyra sp. lineage zone or Mesoendo­ Morocco, from where HOTTINGER (1967), described thyra sp. interval zone, SEPTFONTAINE (1984) most of the Early Jurassic lituolid species. This subdi­ or Mesoendothyra sp. partial-range zone vision not only correlates well with the biostratigraphy Boundaries: from the first occurrence of Mesoendothy­ of lituolids in Velebit Mt., but also in the wider area of ra sp. sensu SEPTFONTAINE (1988) to the first Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 23

occurrence of Lituosepta recoarensis. (1984). In Velebit Mt. and Lika the index fossil O. Important taxa: Mesoendothyra sp. and Lituolipora primaeva also reaches maximum abundance with­ termieri. in this zone, so it may also be defined as an abun­ dance zone. However, if one analyzes the ranges of Stratigraphic range: transitional beds between Lower and Upper Sinemurian. the three most important index fossils found within Lower Jurassic deposits – L. recoarensis, O. primae­ Discussion: Within the previously established Lower va and O. praecursor – this zone can be further sub­ Jurassic, Hettangian, Sinemurian and Pliensbachian divided into two subzones: Orbitopsella primaeva united zone, a dasyclad algal – Palaeodasycladus – Lituosepta recoarensis concurrent-range subzone mediterraneus zone was separated on the AdCP (e.g. and Lituosepta recoarensis – Orbitopsella praecursor FARINACCI & RADOIČIĆ, 1964; NIKLER & interval subzone. SOKAČ, 1968; GUŠIĆ et al., 1971; VELIĆ, 1977). Co-occurring foraminifera have no stratigraphic val­ 3.2.2.4.1. Orbitopsella primaeva–Lituosepta recoarensis ue. Foraminiferal biozones were established, as sub­ concurrent-range subzone zones of the aforementioned algal zone only in the Boundaries: from the first occurrence of O. primaeva Upper Sinemurian and Pliensbachian deposits, as a to the last occurrence of L. recoarensis. result of the significant development of orbitopselli­ nids. In the biostratigraphic investigations of solely Index and important taxa: Mesoendothyra sp., L. ter­ foraminiferal assemblages, this is the first time that mieri, L. recoarensis, P. compressa and O. primae­ a Mesoendothyra sp. lineage zone has been deter­ va. mined in the Lower Jurassic deposits of the AdCP. Stratigraphic range: Late Sinemurian–earliest Carix­ SEPTFONTAINE (1984) marked it as an inter­ ian. val-zone. However, according to the International Discussion: This biozone comprises part of the Lower Stratigraphic Guide (SALVADOR, 1994) it corre­ Jurassic deposits with Orbitopsella spp. in which the sponds to a lineage zone, since this form represents index species L. recoarensis and O. primaeva occur an ancestor of L. recoarensis within the evolutionary together. line of Early Jurassic lituolids. 3.2.2.4.2. Lituosepta recoarensis–Orbitopsella 3.2.2.3. Lituosepta recoarensis lineage zone, praecursor interval subzone SEPTFONTAINE (1984) or Lituosepta recoarensis partial-range zone Boundaries: from the last occurrence of L. recoarensis to the first occurrence of O. praecursor. Boundaries: from the first occurrence of L. recoarensis to the first occurrence of Orbitopsella primaeva. Index and important taxa: Mesoendothyra sp., L. ter­ mieri, P. compressa and O. primaeva. Index and important taxa: Mesoendothyra sp., L. ter­ mieri, L. recoarensis and Planisepta compressa. Stratigraphic range: Early Carixian. Stratigraphic range: older Late Sinemurian (Early Discussion: This subzone comprises a part of the Low­ Lotharingian). er Jurassic orbitopsella-bearing deposits containing only O. primaeva. Other forms continue from zones Discussion: This zone is determined as a lineage zone, older than this to the younger biozones. and not interval or range or taxon-range zone, because L. recoarensis within the evolutionary line 3.2.2.5. Orbitopsella praecursor taxon-range zone of Early Jurassic lituolids has a place between the (SARTONI & CRESCENTI, 1962) ancestor Mesoendothyra sp. and O. primaeva. The Boundaries: stratigraphic range of O. praecursor. stratigraphic range of L. recoarensis on the AdCP continues from this to the next zone. Index and important taxa: Mesoendothyra sp., L. ter­ mieri, P. compressa, O. primaeva, O. praecursor 3.2.2.4. Orbitopsella primaeva lineage zone, and O. ?dubari. SEPTFONTAINE (1984) or Orbitopsella primaeva Stratigraphic range: Early Pliensbachian (Late Carix­ partial-range zone ian). Boundaries: from the first occurrence of O. primaeva Discussion: The Orbitopsella praecursor biozone was to the first occurrence of Orbitopsella praecursor. first established by SARTONI & CRESCENTI Index and important taxa: Mesoendothyra sp., L. ter­ (1962) in the Apennines, as a subzone of the Pal­ mieri, L. recoarensis, P. compressa and O. primae­ aeodasycladus mediterraneus cenozone (assemblage va. zone), with a stratigraphic position in the middle Stratigraphic range: Late Sinemurian–Early Carixian. part of the Early Jurassic. This was also accepted in Discussion: Within the evolutionary line of the Early all investigations within the Karst Dinarides. Within Jurassic lituolids, O. primaeva is a ‘transitional’ this zone, all three cited orbitopsellas occur, but the form between L. recoarensis and O. praecur­ occurrence of O. dubari is dubious, although this sor. Therefore, this zone is referred to as a lineage is not critical either for chronostratigraphical or for zone in the manner proposed by SEPTFONTAINE biostratigraphical subdivision. Theoretically, this 24 Geologia Croatica 60/1

zone might be subdivided as in the Orbitopsella pri­ 3.2.2.7. Pseudocyclammina liassica taxon-range zone, maeva lineage zone into two subzones: Orbitopsella SEPTFONTAINE (1984) primaeva–Orbitopsella praecursor concurrent-range Boundaries: stratigraphic range of P. liassica. subzone and Orbitopsella praecursor abundance Index and important taxa: L. termieri, P. compressa subzone. and P. liassica. 3.2.2.5.1. Orbitopsella praecursor–Orbitopsella Stratigraphic range: Late Pliensbachian (Late Domer­ primaeva concurrent-range subzone ian). Boundaries: from the first occurrence of O. praecursor Discussion: Investigation of detailed undisturbed sec­ to the last occurrence of O. primaeva. tions of Lower Jurassic deposits highlighted layers Index and important taxa: Mesoendothyra sp., L. ter­ where P. liassica was determined, which had not mieri, P. compressa, O. primaeva, O. praecursor, previously been recorded in the area of the Karst Biokovina gradacensis and Bosniella oenensis. Dinarides. Investigations in the Western Mediter­ ranean realm (HOTTINGER, 1967; SEPTFON­ Stratigraphic range: Late Carixian. TAINE, 1984) showed the exact position of this spe­ Discussion: This zone covers part of the Lower Jurassic cies in the Late Domerian, and SEPTFONTAINE deposits in which the stratigraphic ranges of two (1984) founded a biozone of the same name. Com­ orbitopsella index forms overlap: the first part of the parison of the stratigraphic ranges of index species O. praecursor stratigraphic range and the last part of of Early Jurassic foraminifera of these areas with the the O. primaeva range. These forms are followed by Karst Dinarides (Fig. 2) shows perfect correlation, a standard assemblage of Early Jurassic lituolids, the and further supports the establishment of this zone. stratigraphic ranges of which continued from earlier biozones. B. gradacensis occurs in this zone for the 3.2.2.8. Pseudocyclammina liassica–Gutnicella cayeuxi first time. interval zone Boundaries: from the last occurrence of P. liassica to 3.2.2.5.2. Orbitopsella praecursor abundance subzone the first occurrence of Gutnicella cayeuxi. Boundaries: maximum abundance of the species O. Important taxa: Involutina liassica, Agerina martana praecursor. and Mesoendothyra croatica. Index and important taxa: Mesoendothyra sp., L. ter­ Stratigraphic range: Toarcian–Early Aalenian. mieri, P. compressa, Orbitopsella ?dubari, O. prae­ Discussion: Cited taxa have a wider stratigraphic range cursor, B. gradacensis and B. oenensis. within Lower and Middle Jurassic deposits, so this Stratigraphic range: Late Carixian–earliest Domerian. zone contains no index taxa or even a large number Discussion: The index form O. praecursor is the most of foraminiferal taxa as in previous biozones. This is frequent and this zone reflects its maximum abun­ a consequence of altered depositional environments, dance in the younger part of its stratigraphic range. from mostly subtidal and lagoonal to either deeper Along with O. praecursor and other typical Early or shallower water regions. In the NW part of the Jurassic lituolids the occurrence of O. dubari is sup­ platform, regional deepening was recorded, with posed. local terrigenous influences, while in the SE part of the platform shallowing occurred, marked by depo­ 3.2.2.6. Orbitopsella praecursor–Pseudocyclammina sition of mostly oolitic limestones. Such environ­ liassica interval zone ments were probably not very favourable for benthic Boundaries: from the last occurrence of O. praecursor organisms, so only small, undeterminable foraminif­ to the first occurrence of P. liassica. era and A. martana, which has a wider stratigraphic Index and important taxa: Mesoendothyra sp., L. ter­ range within Lower Jurassic, have been found in mieri and P. compressa. this zone. I. liassica has been determined only in deposits indicating an open marine influence, such Stratigraphic range: Late Pliensbachian (Domerian); as platform margins and slopes. M. croatica rarely probably also the beginning of Toarcian. appears in Toarcian beds, but is frequent in Middle Discussion: Although this zone contains frequent speci­ Jurassic deposits. mens of L. termieri and P. compressa (general index fossils of the middle part of the Early Jurassic 3.2.3. Middle Jurassic – Sinemurian and Pliensbachian), they cannot be Similar to the Lower Jurassic, the Middle Jurassic used for naming this zone, since they also occur in deposits of the Karst Dinarides were not biostrati­ younger and older biozones. Therefore, due to the graphically subdivided in detail. Previously, one zone lack of index taxa this zone is defined as an interval with three subzones had been proposed: (1) Trochami­ zone. nidae, Verneuilinidae and Textulariidae, (2) Sellipore­ lla donzellii, and (3) Pfenderinidae (FARINACCI & RADOIČIĆ, 1964), or one zone – Pfenderina salerni­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 25

8.1

Fig. 8 Biostratigraphic subdivision of the Middle Jurassic deposits of the AdCP. Legend: 1) Pseudocyclammina liassica–Gutnicella cayeuxi interval zone; 2) Gutnicella cayeuxi partial range zone; 3) Timidonella sarda taxon-range zone; 4) Timidonella sarda–Pseudodictyopsella jurassica interval zone; 5) Pseudodictyopsella jurassica and Marzoella ficcarellii assemblage zone; 6) Pseudodictyopsella jurassica–Par- avalvulina complicata interval zone; 7) Paravalvulina complicata taxon–range zone; 8) Paleopfenderina salernitana taxon-range zone; 8.1) Satorina apuliensis taxon range subzone; 9) Paleopfenderina salernitana–Kurnubia jurassica interval zone; 10) Kurnubia jurassica lineage zone. tana in younger part of the Middle Jurassic (NIKLER 3.2.3.2. Gutnicella cayeuxi partial range zone or & SOKAČ, 1968; GUŠIĆ, 1969b; GUŠIĆ et al., 1971), Gutnicella cayeuxi–Timidonella sarda interval zone or two zones: (1) Mesoendothyra croatica, and (2) Selli­ Boundaries: from the first occurrence of G. cayeuxi to porella donzellii (VELIĆ, 1977; VELIĆ & SOKAČ, the first occurrence of T. sarda. 1978a). Index and important taxa: G. cayeuxi, Mesoendothyra Biozonation of the Middle Jurassic deposits pre­ croatica and Pseudocyclammina maynci. sented here (Fig. 8) is based upon recent investigations, Stratigraphic range: Middle to Late Aalenian. during which numerous index species have been deter­ mined. Among them are taxa which were previously Discussion: Deposits of this zone and the next one relatively unknown in the area of the AdCP or have could also be defined as an assemblage zone with­ been discovered for the first time. in the stratigraphic range of both G. cayeuxi and T. sarda, as proposed by SEPTFONTAINE et al. 3.2.2.8. and 3.2.3.1. Pseudocyclammina liassica– (1991). Given the fact that the first occurrences Gutnicella cayeuxi interval zone or Mesoendothyra of the cited taxa represent important biohorizons, croatica partial range zone deposits between them were defined as an interval Boundaries: from the first occurrence of Mesoendothy­ zone. Besides G. cayeuxi, P. maynci is another index ra croatica to the first occurrence of G. cayeuxi. form for these deposits. Important taxa: M. croatica. 3.2.3.3. Timidonella sarda taxon-range zone, Stratigraphic range: Toarcian and Early Aalenian. SEPTFONTAINE (1980) Discussion: This interval-zone comprises Jurassic Boundaries: stratigraphic range of T. sarda. deposits between the first occurrence of M. croatica Index and important taxa: T. sarda, M. croatica, G. to the first occurrence of G. cayeuxi, but it can also cayeuxi, P. maynci, Spiraloconulus perconigi and S. be defined as the M. croatica partial range zone, giganteus. since it comprises the lowermost part of the strati­ graphic range of this taxon. M. croatica is the most Stratigraphic range: Late Aalenian–Early Bajocian. important of the index species in this zone. Within Discussion: T. sarda was discovered on the AdCP some this weakly fossiliferous interval, a chronostrati­ 20 years ago, but this fact was never published. The graphic boundary between the Lower and Middle beginning of its stratigraphic range coincides with Jurassic occurs, since the first occurrence of M. the last occurrence of G. cayeuxi. In the younger croatica probably took place in the early Toarcian part of its stratigraphic range S. perconigi and S. deposits. giganteus also occur. Given these relationships, this 26 Geologia Croatica 60/1

zone was defined as being of latest Aalenian and Discussion: Deposits of this biozone contain a rich earliest Bajocian age, according to the results of the assemblage of small foraminifera, most of which investigations on the AdCP, and comparison with belong to taxa of wider stratigraphic ranges. The well established chronostratigraphic positions of maximum frequency of M. croatica was determined, these taxa in the Western Mediterranean realm (e.g. so this zone could also be defined as an abundance BASSOULLET et al., 1976; SEPTFONTAINE, zone. This zone comprises a major part of the strati­ 1980; SEPTFONTAINE et al., 1991; BASSOUL­ graphic range of the important dasyclad species Sell­ LET, 1997). iporella donzelli, which is less frequent in deposits of the succeeding zone. 3.2.3.4. Timidonella sarda–Pseudodictyopsella jurassica interval zone 3.2.3.7. Paravalvulina complicata taxon-range zone Boundaries: from the last occurrence of T. sarda to the Boundaries: stratigraphic range of the species P. com­ first occurrence of P. jurassica. plicata. Important taxa: M. croatica, P. maynci and S. gigan­ Index and important taxa: P. complicata, Pseudoeg­ teus. gerella elongata and M. croatica. Stratigraphic range: younger part of the Early Bajo­ Stratigraphic range: latest Bajocian. cian. Discussion: The stratigraphic positions of P. compli­ Discussion: There are no index taxa, and only two other cata and P. elongata, which were recently unknown important taxa – P. maynci and S. giganteus contin­ in Jurassic deposits of the AdCP (VELIĆ, 2005), uing from the earlier biozone, as well as M. croati­ are the same as those defined by SEPTFONTAINE ca, which has a wider Middle Jurassic range. Gener­ (1988) for the Western Mediterranean. P. complica­ ally, from the biostratigraphic point of the view it is ta occurs somewhat earlier (probably at the very end important to note that this zone marks the beginning of Bajocian) than P. elongata, which occurs at the of the stratigraphic range of the important Bajo­ Bajocian–Bathonian transition and through Batho­ cian–early Bathonian dasyclad species Selliporella nian. Therefore this zone occurs in part of the Mid­ donzellii. dle Jurassic deposits directly underlying the horizon characterized by the first occurrence of Paleopfend­ 3.2.3.5. Pseudodictyopsella jurassica and Marzoella erina salernitana, the Bathonian index form which ficcarellii assemblage zone is widespread over the Mediterranean. Boundaries: stratigraphic ranges of P. jurassica and M. ficcarellii. 3.2.3.8. Paleopfenderina salernitana taxon-range zone SARTONI & CRESCENTI (1962) Index and important taxa: P. jurassica, M. ficcarellii, M. croatica and S. giganteus. Boundaries: stratigraphic range of P. salernitana. Stratigraphic range: middle part of the Bajocian. Index and important taxa: P. salernitana, P. trochoi­ dea, M. croatica, P. elongata, Alzonella cuvillieri, Discussion: This foraminiferal assemblage is character­ Kilianina blancheti, Satorina apuliensis, S. meso­ istically of high abundance but low species diversity, jurassica and Orbitammina elliptica. where the only index forms are P. jurassica and M. Stratigraphic range: Bathonian–?earliest Callovian. ficcarellii found in the central part of the Bajocian succession (Fig. 3), probably in transitional depos­ Discussion: P. salernitana is connected with the begin­ its between the early and late Bajocian of Biokovo ning of the Bathonian on the AdCP, so the lower Mt., together with S. giganteus. Despite this nar­ boundary of this biozone coincides with the Bajo­ row stratigraphic range, and their occurrence in only cian–Bathonian chronostratigraphic boundary. In one locality, a separate biostratigraphic unit was the younger part of this zone the S. apuliensis sub­ defined. However, the occurrence of these species zone was defined within the stratigraphic range of in one level, could also allow these biohorizons to the Late Bathonian. be termed the P. jurassica and M. ficcarellii FADs 3.2.3.8.1. Satorina apuliensis taxon range subzone (First Appearance Datums). Since investigation of the Middle Jurassic deposits of the AdCP represents Boundaries: stratigraphic range of S. apuliensis. a work in progress, new data which would confirm Index and important taxa: Satorina apuliensis, S. the establishment of this zone are expected. mesojurassica, P. salernitana, P. trochoidea, A. cuvillieri, K. blancheti, O. elliptica and P. elongata. 3.2.3.6. Pseudodictyopsella jurassica–Paravalvulina Stratigraphic range: Late Bathonian. complicata interval zone Discussion: This subzone comprises the younger part of Boundaries: from the last occurrence of P. jurassica to the P. salernitana taxon-range zone. The stratigraph­ the first occurrence of P. complicata. ic range of S. apuliensis on the AdCP is longer than Important taxa: M. croatica and S. giganteus. that of the other cited late Bathonian species, so it is Stratigraphic range: approximately Middle Bajocian. probable that it occurred until the end of the Batho­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 27

nian, or even to the earliest Callovian. However, the 3.2.4. Late Jurassic stratigraphic range of P. salernitana exceeds even Hitherto biostratigraphic subdivisions of the Upper the range of S. apuliensis, so it is almost certain that Jurassic deposits of the AdCP were mostly based on it also occurs in the early Callovian. calcareous algae (NIKLER & SOKAČ, 1968; GUŠIĆ, 3.2.3.9. Paleopfenderina salernitana–Kurnubia 1969b; GUŠIĆ et al., 1971; VELIĆ, 1977; VELIĆ & jurassica interval zone SOKAČ, 1974, 1978a). Although benthic foraminifera were cited in these papers they were not discussed in Boundaries: from the last occurrence of P. salernitana detail. Only GUŠIĆ (1969b) and GUŠIĆ et al. (1971), to the first occurrence of Kurnubia jurassica. following subdivision by SARTONI & CRESCENTI Important taxa: P. arabica, P. crusei and Trocholina (1962), separated the Kurnubia palastiniensis assem­ gr. alpina–elongata. blage zone in the older part of the Late Jurassic. Stratigraphic range: Callovian. K. palastiniensis has a stratigraphic range in the Discussion: Within the interval from the last occur­ AdCP from the earliest Oxfordian to the Late Tithonian, rence of P. salernitana to the first occurrence of K. so it has an important role in biostratigraphy of the Late jurassica the foraminiferal assemblage contains no Jurassic. Therefore the major part of the Upper Jurassic index taxa. However, the stratigraphic range of this deposits can be defined as a K. palastiniensis superzone zone is unambiguous, since the last occurrence of P. (Fig. 9). salernitana at the AdCP was in the early Callovian, and the first occurrence of K. jurassica took place in 3.2.4.1– 3.2.4.6. Kurnubia palastiniensis superzone the late Callovian. Therefore, the range of this zone Boundaries: Stratigraphic range of K. palastiniensis. is Callovian. Index and important taxa: Includes all taxa cited in the next four biozones. 3.2.3.10. Kurnubia jurassica lineage zone or Kurnubia jurassica partial-range zone or Kurnubia jurassica– Stratigraphic range: Oxfordian–Late Tithonian. Kurnubia palastiniensis interval zone Discussion: This superzone comprises the following Boundaries: from the first occurrence of K. jurassica biozones: to the first occurrence of K. palastiniensis. – Kurnubia palastiniensis lineage zone, Important taxa: P. arabica, P. crusei, Trocholina gr. – Kurnubia wellingsi lineage zone, alpina–elongata and K. jurassica. – Chablaisia chablaisensis–Alveosepta jaccardi inter­ Stratigraphic range: latest Callovian–earliest Oxford­ val zone, ian. – Alveosepta jaccardi partial-range zone, Discussion: This interval zone comprises the young­ – Parurgonina caelinensis–Conicokurnubia orbitoli­ est part of the Middle Jurassic and the oldest part niformis concurrent range-zone, and of the Upper Jurassic deposits with clearly defined – C. orbitoliniformis–K. palastiniensis interval zone boundaries between two biohorizons, the first occur­ (highest-occurrence zone). rences of two cited species of Kurnubia. Therefore, this zone includes the chronostratigraphic boundary 3.2.4.1. Kurnubia palastiniensis lineage zone or Kur­ between the Middle and Late Jurassic. nubia palastiniensis partial–range zone or Kurnubia palastiniensis–Kurnubia wellingsi interval zone Alternatively, this zone can also be defined as a lin­ eage zone because of the taxonomic position of K. Boundaries: From the first occurrence of K. palastini­ jurassica as the first form in the evolutionary trend ensis to the first occurrence of K. wellingsi. of kurnubians. GUŠIĆ (1969a, b) discussed and Important taxa: K. palastiniensis, K. jurassica, Cha­ illustrated K. jurassica in Upper Jurassic deposits of blaisia chablaisensis and Karaisella uzbekistanica. Central Croatia from inner platform environments of Stratigraphic range: Early Oxfordian. the NW parts of the former AdCP. Since then, this Discussion: Separation of this part of the Upper Jurassic species has not been cited again in publications deal­ deposits as a lineage zone is based on the position ing with the AdCP. However, recent investigations of K. palastiniensis within the evolutionary trend of a series of detailed columns dispersed around the of kurnubians P. crusei – K. jurassica, K. palasti­ AdCP, suggest that this form occurs before K. pal­ niensis – K. wellingsi – C. orbitoliniformis. Due to astiniensis, indicating the youngermost Callovian. the lack of foraminiferal species with a stratigraphic Among kurnubians, K. jurassica represents a tran­ range confined to this interval, this unit could also sitional form between P. crusei and K. palastinien­ be defined as an interval zone, or as the K. palas­ sis. Therefore the definition of this zone as a lineage tiniensis partial-range zone, since it comprises the zone is also approved within the evolutionary line first part of the stratigraphic range of this species. of kurnubians: Praekurnubia crusei – K. jurassica – Within tempestites of the younger part of this zone Kurnubia palastiniensis – K. wellingsi – Conicokur­ and the older part of the next zone, Ch. chablaisen­ nubia orbitoliniformis. sis, Trocholina alpina and T. elongata (chablaisia– 28 Geologia Croatica 60/1

Oxfordian Kimmeridgian Tithonian

1−6

Fig. 9 Biostratigraphic subdivision of the Upper Jurassic deposits of the AdCP. Legend: 1–6) Kurnubia palastiniensis superzone: 1) Kurnubia palastiniensis lineage zone; 2) Kurnubia wellingsi lineage zone; 3) Chablaisia chablaisensis-Alveosepta jaccardi interval zone; 4) Alveo- septa jaccardi partial-range zone; 5) Parurgonina caelinensis–Conicokurnubia orbitoliniformis concurrent range zone; 6) Conicokurnubia orbitoliniformis–Kurnubia palastiniensis interval zone (highest-occurrence zone); 7) Kurnubia palastiniensis–Protopeneroplis ultragranu- lata interval zone.

trocholina tempestites) are abundant in the western Important taxa: L. mirabilis, K. palastiniensis, K. well­ part of the AdCP – Velebit Mt., Gorski Kotar and ingsi and K. jurassica. Istria. So in this area, this part of the unit could be Stratigraphic range: younger part of the Late Oxford­ defined as an abundance zone. ian. 3.2.4.2. Kurnubia wellingsi lineage zone or Labyrin­ Discussion: There are no foraminiferal index-taxa in thina mirabilis–Karaisella uzbekistanica concurrent- the uppermost Oxfordian deposits. However, they range zone or Kurnubia wellingsi partial-range zone could be separated as an interval zone between two Boundaries: From the first occurrence of K. wellingsi biostratigraphic boundaries: the last occurrence of Ch. chablaisensis and K. uzbekistanica and the first to the first occurrence of Conicokurnubia orbitolini­ occurrence of A. jaccardi, C. orbitoliniformis and formis for the lineage and partial-range zones. From Kilianina? rahonensis. the first occurrence of L. mirabilis to the last occur­ rence of K. uzbekistanica for the concurrent-range 3.2.4.4. Alveosepta jaccardi partial–range zone zone. or Alveosepta jaccardi–Parurgonina caelinensis Stratigraphic range: older part of Late Oxfordian. interval zone Important taxa: K. wellingsi, K. palastiniensis, K. Boundaries: from the first occurrence of A. jaccardi to jurassica, Ch. chablaisensis, K. uzbekistanica and the first occurrence of P. caelinensis. L. mirabilis. Important taxa: L. mirabilis, K. palastiniensis, K. well­ ingsi, K. jurassica, A. jaccardi, C. orbitoliniformis Discussion: In the aforementioned evolution of Jurassic and K.? rahonensis. kurnubians, K. wellingsi represents a transitional form between K. palastiniensis and C. orbitolini­ Stratigraphic range: Early Kimmeridgian. formis, for the Late Oxfordian and the beginning of Discussion: This foraminiferal assemblage is currently the Kimmeridgian. Therefore, this zone is defined as the richest known in the Kimmeridgian of the AdCP. a lineage-zone. Alternatively, it may be defined as a Most of the taxa have a wide stratigraphic range, concurrent-range zone from the first occurrence of both those occurring for the first time in younger L. mirabilis to the last occurrence of K. uzbekista­ zones of the Middle Jurassic, and those first occur­ nica, the stratigraphic ranges of which overlap, or as ring in the oldest Late Jurassic zones. A. jaccardi, the K. wellingsi and L. mirabilis partial-range zone. C. orbitoliniformis and K.? rahonensis occur here K. wellingsi is common, which supports and enables for the first time, but they are not index forms, since the definition of this zone. Its first occurrence coin­ their stratigraphic ranges continue into younger cides with the first occurrence of L. mirabilis, and zones. Since former biostratigraphic zonations of they both continue into the next zone. the AdCP deposits were mostly based on rich calcar­ eous algal assemblages it should be noted that this 3.2.4.3. Chablaisia chablaisensis–Alveosepta jaccardi zone covers the stratigraphic range of Pseudocly­ interval zone peina cirici. Boundaries: from the last occurrence of Ch. chablai­ sensis to the first occurrence of A. jaccardi. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 29

3.2.4.5. Parurgonina caelinensis–Conicokurnubia posed, since SEPTFONTAINE et al. (1991) defined orbitoliniformis concurrent range zone or Parurgonina Anchispirocyclina lusitanica taxon-range zone in the caelinensis partial–range zone Late Tithonian (Portlandian). Boundaries: from the first occurrence of P. caelinensis to the last occurrence of C. orbitoliniformis. 3.2.5. Early Cretaceous Important taxa: K. palastiniensis, K. wellingsi, C. orbi­ Relatively rich foraminiferal assemblages in the Lower toliniformis, A. jaccardi, L. mirabilis, K.? rahonen­ Cretaceous platform carbonate deposits of the Adriatic sis and Parurgonina caelinensis. Carbonate Platform have previously enabled the most Stratigraphic range: Middle and Late Kimmeridgian. detailed biostratigraphic zonation (Fig. 10). Zonations Discussion: There are no index forms in this foraminif­ proposed in former studies, e.g. those by VELIĆ & eral assemblage. The upper boundary is marked SOKAČ (1978b) and VELIĆ (1988), will represent the by the last occurrence of A. jaccardi and C. orbi­ basis for biozonation proposed here. toliniformis, and this approximately corresponds to 3.2.5.1. Protopeneroplis ultragranulata taxon-range the Kimmeridgian–Tithonian chronostratigraphic zone (VELIĆ, 1988) boundary, and the beginning of the stratigraphic Boundaries: stratigraphic range of P. ultragranulata. range of dasyclad alga Clypeina jurassica. Index and important taxa: P. ultragranulata is the 3.2.4.6. Conicokurnubia orbitoliniformis–Kurnubia only index fossil, and other forms have a wider palastiniensis interval zone (highest-occurrence zone) stratigraphic range, either from the Jurassic into the Boundaries: from the last occurrence of C. orbitolini­ Early Cretaceous or through the Cretaceous, e.g. formis to the last occurrence of K. palastiniensis. Mohlerina basiliensis, Trocholina alpina, T. elon­ gata, Mayncina bulgarica and Charentia cuvillieri. Important taxa: K. palastiniensis, P. caelinensis and K.? rahonensis. Stratigraphic range: Late Tithonian–Early Berriasian. Stratigraphic range: Early Tithonian. Discussion: Central parts of the AdCP in present Croatia and W Bosnia were emergent during the Late Titho­ Discussion: In addition to K. palastiniensis, P. cae­ nian and Early Berriasian, while in other parts per­ linensis and K.? rahonensis, which have their last itidal environments predominated, which were very occurrences near the end of this zone, this assem­ unfavourable for benthic foraminifera. This is the blage contains other forms of much wider strati­ main reason why the foraminiferal assemblage is graphic range (Fig. 4). In the continuous succession very poor. of the Upper Jurassic, carbonate deposits of this zone can exceed 400 m thickness, and contain the 3.2.5.2. Protopeneroplis ultragranulata–Vercorsella middle part of the stratigraphic range of the dasyclad camposaurii interval zone index fossil Clypeina jurassica. Boundaries: from the last occurrence of P. ultragranu­ 3.2.4.7. Kurnubia palastiniensis–Protopeneroplis lata to the first occurrence of V. camposaurii. ultragranulata interval zone Important taxa: there are no important taxa, since only Boundaries: from the last occurrence of K. palastinien­ forms of wider stratigraphic range were determined, sis to the first occurrence of P. ultragranulata. such as Trocholina alpina, T. elongata, Mayncina bulgarica and Charentia cuvillieri. Important taxa: Anchispirocyclina lusitanica, probably also A. neumannae. Stratigraphic range: Middle and Late Berriasian. Stratigraphic range: Late Tithonian. Discussion: As for the previous zone, carbonate depos­ its of this zone were deposited in peritidal environ­ Discussion: In the youngermost part of the Upper ments, and temporary subtidal environments. There­ Jurassic deposits of the AdCP a significant decrease fore the foraminiferal assemblage is also poor and in frequency and diversity of the foraminiferal without index fossils. assemblage is observed. This is largely because the peritidal environments were more favourable for 3.2.5.3. Vercorsella camposaurii taxon-range zone abundant calcareous algae. This level is famous (SARTONI & CRESCENTI, 1962) throughout the platform for rich assemblages and Boundaries: stratigraphic range of V. camposaurii. the maximum development of dasyclad species Index and important taxa: Vercorsella tenuis, V. cam­ including Clypeina jurassica and Campbelliella stri­ posaurii, Montsalevia salevensis, Meandrospira ata. A. lusitanica and/or A. ?neumannae are index favrei, Haplophragmoides joukowskyi, Trocholina foraminifera but only for the youngermost part of delphinensis and T. sagittaria. this zone. They have only been discovered at two localities: Biokovo Mt., and around Dubrovnik. Sep­ Stratigraphic range: latest Berriasian, Valanginian and aration of a specific zone for the entire AdCP is not earliest Hauterivian. possible because of such very scarce and restricted Discussion: SARTONI & CRESCENTI (1962) defined observations, although theoretically it might be pro­ the Cuneolina camposaurii assemblage zone with 30 Geologia Croatica 60/1

7−10

Fig. 10 Biostratigraphic subdivi- sion of the Lower Creta- ceous deposits of the AdCP. Legend: 1) Protopeneroplis ultragranulata taxon-range zone; 2) Protopeneroplis ultragranulata–Vercorsella camposaurii interval zone; 3) Vercorsella camposaurii taxon-range zone; 4) Ver- corsella camposaurii–Cam- panellula capuensis interval zone; 5) Campanellula capuensis taxon-range zone; 6) Campanellula capuensis– Palorbitolina lenticularis inter- val zone; 7–10) Palorbitolina lenticularis superzone: 7) Rectodictyoconus gigan- teus and Palorbitolina lenticu- laris assemblage zone, 8) Praeorbitolina cormyi line- age zone, 9) Praeorbitolina wienandsi lineage zone, 10) Mesorbitolina lotzei line- age zone; 11) Mesorbitolina parva lineage zone; 12) Mesorbitolina texana line- age zone; 13) Mesorbitolina subconcava taxon-range zone; 14) ‘Valdanchella’ der- courti partial-range zone; 15) Neoiraquia convexa taxon-range zone.

the stratigraphic range of Valanginian–Aptian. (ARNAUD-VANNEAU et al., 1991). Almost all The stratigraphic range of V. camposaurii is also forms defined as C. camposaurii by VELIĆ (1973, stated as being Valanginian–Aptian, but they noted 1977) correspond to other species of Vercorsella. that elsewhere this species is of Valanginian age Other additional index forms in this assemblage (Provence, according to PFENDER, 1938) or Bar­ include V. tenuis, M. salevensis, M. favrei, H. jou­ remian age (Montenegro, according to RADOIČIĆ, kowskyi, T. delphinensis and T. sagittaria. Other 1960). Detailed investigations on the AdCP have taxa have different, but wider stratigraphic ranges proven that forms which correspond to the original through the Early Cretaceous (Fig. 5). description of V. camposaurii (SARTONI & CRES­ CENTI, 1962) are of exclusively Neocomian age, 3.2.5.4. Vercorsella camposaurii–Campanellula with a stratigraphic range of latest Berriasian–earli­ capuensis interval zone est Hauterivian. So this form can be considered as Boundaries: from the last occurrence of V. camposaurii an index form of the Valanginian. This species has or other index taxa of the previous zone to the first latterly been included into the genus Vercorsella occurrence of C. capuensis. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 31

Important taxa: No important taxa, only forms of wid­ Limestones’ (VELIĆ, 1988). Within this superzone, er stratigraphic range, e.g. T. alpina, T. elongata and which could also be defined as the P. lenticularis T. sagittaria. taxon-range zone (HUSINEC et al., 2000), four Stratigraphic range: Within the Hauterivian. zones have been determined (Fig. 12): Discussion: This is an interval within the Hauterivian – Rectodictyoconus giganteus and Palorbitolina len­ with a poor foraminiferal assemblage and no index ticularis assemblage zone, forms. All taxa have wider stratigraphic ranges – Praeorbitolina cormyi lineage zone, through the Early Cretaceous, and the cited trocholi­ – Praeorbitolina wienandsi lineage zone, and nas do not occur after the Hauterivian. – Mesorbitolina lotzei lineage zone. 3.2.5.5. Campanellula capuensis taxon-range zone, Within deposits of the P. lenticularis superzone on VELIĆ (1988) the AdCP, VELIĆ & SOKAČ (1978a) and VELIĆ Boundaries: stratigraphic range of C. capuensis. (1988) discovered all of the first (Aptian) part of Index and important taxa: C. capuensis, T. sagittaria. the evolutionary trend of orbitolinids according to SCHROEDER (1975, 1979). Therefore, these zones Stratigraphic range: latest Hauterivian and transition were defined as lineage zones, and were named after into the Barremian. the first three species within the evolutionary line. Discussion: C. capuensis is very frequent and important in this assemblage. Within the relatively poor assem­ 3.2.5.7. Rectodictyoconus giganteus and Palorbitolina blages in Lower Cretaceous deposits from the early lenticularis assemblage zone (VELIĆ, 1988) or Hauterivian to the late Barremian, its stratigraphic Palorbitolina lenticularis partial–range zone position is very important, as it represents an essen­ Boundaries: from the first occurrence of R. giganteus tial biofacies. Other forms have wider, but variable and P. lenticularis to the first occurrence of Praeor­ stratigraphic ranges through the Early Cretaceous. bitolina cormyi. This zone is also characterized by end of the strati­ Index and important taxa: R. giganteus, P. lenticula­ graphic ranges of T. alpina and T. elongata. ris, Derventina filipescui, Trocholina molesta, T. odukpaniensis, Neotrocholina aptiensis, N. fribur­ 3.2.5.6. Campanellula capuensis–Palorbitolina lenticularis interval zone gensis, Nautiloculina broennimanni and N. cretacea. Boundaries: Between two important biohorizons last Stratigraphic range: Late Barremian and beginning of occurrence of C. capuensis and first occurrence of the Early Aptian (Bedoulian). P. lenticularis. Discussion: R. giganteus is the index form for this zone. However, this orbitolinid was only discovered Important taxa: No important taxa, only forms of in the NE marginal area of the AdCP in the vicinity wider stratigraphic range, e.g. Vercorsella lauren­ of Banja Luka and Jajce. Elsewhere, over the major tii, Novalesia distorta, N. cornucopia, N. producta, part of the platform, deposits of this zone are rec­ Sabaudia minuta, Debarina hahounerensis and T. ognized as belonging to the lowermost part of the sagittaria. ‘Lower Orbitolina Limestones’, where the predomi­ Stratigraphic range: Early and Middle Barremian. nant foraminifera is P. lenticularis, and praeorbito­ Discussion: In this relatively rich foraminiferal assem­ linids are absent (VELIĆ & SOKAČ, 1978a). All blage there are no index forms. This zone marks other forms in the foraminiferal assemblage have a the beginning of the stratigraphic ranges of species wider stratigraphic range, except partially N. aptien­ that are relatively abundant in younger Lower Cre­ sis and N. friburgensis, which can also be found in taceous deposits of the AdCP – S. minuta and D. the next three zones of the Early Aptian. hahounerensis, as well as species of Vercorsella and Novalesia. 3.2.5.8. Praeorbitolina cormyi lineage zone (VELIĆ, 1988) or Praeorbitolina cormyi partial-range zone 3.2.5.7–3.2.5.10. Palorbitolina lenticularis superzone Boundaries: from the first occurrence of P. cormyi to (taxon-range zone, HUSINEC et al., 2000) the first occurrence of Praeorbitolina wienandsi. Boundaries: the stratigraphic range of P. lenticularis. Index and important taxa: P. lenticularis, R. gigan­ Index and important taxa: Comprises all taxa includ­ teus, P. cormyi, Paleodictyoconus actinostoma, ed in the next four biozones. It is important to note Sabaudia briacensis, Pfenderina globosa, D. filipes­ that the most important species in all of these zones cui, N. aptiensis, N. friburgensis, T. molesta, T. oduk­ is P. lenticularis. paniensis, N. broennimanni, N. cretacea, Choffatella Stratigraphic range: Late Barremian and Early Aptian. decipiens, Voloshinoides murgensis and Pseudo­ lituonella conica. Discussion: Carbonate deposits within the stratigraphic range of P. lenticularis on the AdCP represent the Stratigraphic range: Early Aptian (Early Bedoulian). regionally most important biofacies level of the Discussion: P. cormyi is the only index form of this Early Cretaceous, known as the ‘Lower Orbitolina zone. P. lenticularis, predominates the assemblage 32 Geologia Croatica 60/1

together with forms of variable stratigraphic range 3.2.5.11. Mesorbitolina parva lineage zone or through the Early Cretaceous, including index spe­ Mesorbitolina parva partial-range zone cies for the Late Barremian and Early Aptian – N. Boundaries: from the first occurrence of M. parva to aptiensis and N. friburgensis, as well as species the first occurrence of Mesorbitolina texana. determined only from the Early Aptian – S. briacen­ Index and important taxa: M. parva, N. aptiensis, sis, C. decipiens, V. murgensis and P. conica. At the Archalveolina reicheli, S. auruncensis, S. capitata NE margin of the AdCP (Mt. Vlašić in Central Bos­ and Haplophragmoides globosus. nia), the orbitolinid P. actinostoma was discovered, which is unknown in other areas of the platform, Stratigraphic range: transitional level Early–Late Apti­ although there are data on the discovery of the genus an and Late Aptian (Gargasian). Paleodictyoconus (e.g. VELIĆ & SOKAČ, 1978a). Discussion: The first occurrence of M. parva was noted immediately after the last occurrence of M. lotzei in 3.2.5.9. Praeorbitolina wienandsi lineage zone or the vicinity of Ogulin (Central Croatia). This assem­ Praeorbitolina wienandsi partial-range zone blage contains numerous specimens of N. aptiensis, Boundaries: from the first occurrence of P. wienandsi which on the AdCP is regarded as an index form of to the first occurrence of Mesorbitolina lotzei. the Early Aptian (e.g. VELIĆ & SOKAČ, 1978a; Index and important taxa: P. wienandsi and most of VELIĆ, 1988; HUSINEC et al., 2000). It is there­ taxa cited in the previous zone. fore probable that the first appearance of M. parva Stratigraphic range: middle part of the Early Aptian occurred in the final part of the Early Aptian and/ (Middle Bedoulian). or transition into the Late Aptian. H. globosus was Discussion: Foraminiferal frequencies in this assem­ recorded in this zone, but is illustrated as Trocham­ blage are almost identical to the assemblage of the minoides coronus by CHIOCCHINI et al. (1994). P. cormyi zone. It is however enriched by the index Other taxa of wider stratigraphic range through the form P. wienandsi, small trocholinas, and in places early Cretaceous were cited in the rich Late Aptian specifically undeterminable involutinids. P. lenticu­ fossil assemblage in the previous section. Among laris remains a predominant form. them, Praechrysalidina infracretacea reached its maximum in this zone. 3.2.5.10. Mesorbitolina lotzei lineage zone or Mesorbitolina lotzei taxon-range zone (VELIĆ, 1988) Deposits of this zone belong to the lowermost part Boundaries: from the first occurrence of M. lotzei to of the biofacies known as the ‘Upper Orbitolina the first occurrence of Mesorbitolina parva or with­ Limestones’ (VELIĆ & SOKAČ, 1978a; VELIĆ, in the stratigraphic range of M. lotzei. 1988), that are regionally distributed over the AdCP. It encompasses the stratigraphic range of Aptian Index and important taxa: As for the P. cormyi and P. and Albian mesorbitolinids, i.e. Late Aptian and wienandsi zones, enriched in M. lotzei and Sabaudia Early Albian, and, from the biostratigraphic point of capitata, S. auruncensis and Archaealveolina sp. view, deposits of this and the two following zones. Stratigraphic range: Late Early Aptian (Late Bedou­ In comparison with the evolution of important cal­ lian). careous algae, this and the following zone represent Discussion: The foraminiferal assemblage of this zone the maximum of the dasyclad alga Salpingoporella remains mostly unchanged from the composition dinarica RADOIČIĆ (VELIĆ & SOKAČ, 1978a). of the previous two zones, characterized by a pre­ dominance of P. lenticularis, and enrichment in M. 3.2.5.12. Mesorbitolina texana lineage zone (VELIĆ, lotzei. Poorly preserved, very micritized sections 1988) or Mesorbitolina texana partial-range zone determined as Archaealveolina sp. could represent Boundaries: from the first occurrence of M. texana to A. reicheli, described from Late Aptian deposits the first occurrence of Mesorbitolina subconcava. (DE CASTRO, 1966, 1980, 1988). If this specific Index and important taxa: M. texana, M. parva, A. determination is proven to be correct then the occur­ reicheli, S. auruncensis, S. capitata, H. globosus and rence of A. reicheli in the Early Aptian of the AdCP Orbitolinopsis aquitanica. would be the first occurrence of this form (VELIĆ & SOKAČ, 1978a). The end of this zone is char­ Stratigraphic range: Late Aptian. acterized by disappearance of index forms for the Discussion: M. texana is the most widespread mesorbi­ Late Barremian and Early Aptian: P. lenticularis, tolinid form on the AdCP, with a stratigraphic range P. cormyi, P. wienandsi, M. lotzei, P. globosa, D. from the middle of the Late Aptian to the Middle filipescui, N. friburgensis, as well as other trocholi­ Albian. It can be found in limestones originating nids and N. broennimanni, N. cretacea, C. decipiens, in different environments, and is very abundant in S. briacensis, V. murgensis and P. conica. Other lagoonal–subtidal environments and tempestites. forms have continuous stratigraphic ranges into the However, there are areas with very rare occur­ Late Aptian, and this zone represents the begin­ rences (e.g. SE Croatia and islands) or where it is ning of the stratigraphic ranges of S. capitata and S. absent (e.g. W Istria; Fig. 18), due to regional shal­ auruncensis. lowing and emergence (VELIĆ et al., 1989, 2002). Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 33

M. parva spreads through the entire zone, and this nimanni and P. fleuryi. Taxa of wider stratigraphic zone represents the maximum of H. globosus and A. range through several Early Cretaceous zones are reicheli. O. aquitanica occurs infrequently, and is V. scarsellai, V. laurentii, S. minuta, S. auruncensis, restricted to the latest Aptian and earliest Albian. M. bulgarica, and P. infracretacea, which disap­ peared by the end of the Albian, while P. auriger­ 3.2.5.13. Mesorbitolina subconcava taxon-range zone ica, P. heimi, S. cretacea, S. pumila, S. phoenissa, (VELIĆ, 1988) N. picardi and N. simplex continued into the Late Boundaries: stratigraphic range of M. subconcava. Cretaceous (GUŠIĆ & JELASKA, 1990; VELIĆ & Index and important taxa: M. subconcava, M. pervia, VLAHOVIĆ, 1994). Dasyclad alga Salpingoporella M. texana, M. parva, Paracoskinolina sunnilanden­ turgida is, among other microfossils, very impor­ sis, O. aquitanica, A. reicheli, Sabaudia dinapolii, tant for this zone, and it frequently accompanies the S. auruncensis, S. capitata, Cuneolina sliteri and H. cited orbitolinids (VELIĆ & SOKAČ, 1978a, 1979, globosus. 1983; VELIĆ et al., 1995; HUSINEC et al., 2000; Stratigraphic range: Transitional level between the CVETKO TEŠOVIĆ, 2000). Aptian and Albian and/or the Early Albian. 3.2.5.15. Neoiraquia convexa taxon-range zone Discussion: This zone forms the major part of the or Neoiraquia convexa–Conicorbitolina conica ‘Upper Orbitolina Limestones’, a typical facies interval zone of the AdCP (VELIĆ & SOKAČ, 1978a; VELIĆ, Boundaries: stratigraphic range of N. convexa. 1988). The foraminiferal assemblage is domi­ Index and important taxa: N. convexa, ‘V.’ dercourti, nated by mesorbitolinids, among which M. parva Ovalveolina maccagnoae, ?Sellialveolina viallii, occurs, while other species of this genus – M. tex­ Scandonea pumila, S. phoenissa and Protochry­ ana, M. subconcava and M. pervia – reached their salidina elongata. maximum, but disappeared at the end of this zone. Stratigraphic range: latest Albian (Vraconian)–earliest Within this zone the maximum of many other taxa Cenomanian. which range through several Early Cretaceous zones occurred, e.g. A. reicheli, Vercorsella scarsellai, V. Discussion: Index fossils are N. convexa, ‘V.’ dercourti laurentii, V. arenata, S. minuta, S. auruncensis, S. and P. elongata which did not survive into the capitata, S. dinapolii and M. bulgarica. A. reicheli, Cenomanian, as well as O. maccagnoae and S. vial­ V. arenata, S. auruncensis, S. capitata, S. dinapolii, lii, the stratigraphic range of which is Late Albian Rumanoloculina robusta and Glomospira urgoniana and Early Cenomanian. Therefore, this zone would disappear within this zone. In contrast, there is the comprise the boundary between the Albian and the first occurrence of important Cretaceous foraminif­ Cenomanian. Sabaudia minuta ends in this zone, era, which reached their maximum in the Late while all other taxa continue into the Late Creta­ Cretaceous – Cuneolina pavonia, C. parva, Nezza­ ceous. POLŠAK et al. (1982) also mentioned S. zatinella picardi and Pseudonummoloculina heimi. minuta in Lower Cenomanian deposits. The begin­ The lower boundary is, besides the first occurrence ning of the stratigraphic range of the nezzazatids N. of M. subconcava, marked by the occurrence of the gyra and N. conica occurs within this zone, and their cited cuneolinids, including C. sliteri, and approxi­ maximum was during the Cenomanian. mates the Aptian–Albian chronostratigraphic boun­ 3.2.6. Late Cretaceous dary. In spite of the numerous published papers with cited 3.2.5.14.“Valdanchella” dercourti partial-range zone taxa and assemblages of benthic foraminifera from or “Valdanchella” dercourti–Neoiraquia convexa Upper Cretaceous deposits of the AdCP, there are rela­ interval zone (VELIĆ & SOKAČ, 1978b) tively little data on the biostratigraphical zonation. As Boundaries: from the first occurrence of ‘V.’ dercourti previously mentioned, GUŠIĆ & JELASKA (1990) to the first occurrence of N. convexa. correlated lithostratigraphic units of the island of Brač Index and important taxa: ‘V.’ dercourti, Simplorbi­ with the biozonation after FLEURY (1980), and VELIĆ tolina broennimanni, Neoiraquia insolita, Para­ & VLAHOVIĆ (1994) determined five biozones in the coskinolina fleuryi, P. sunnilandensis and S. aurun­ Cenomanian deposits of N Istria. Zones defined here censis. (Fig. 11) represent the first attempt to biostratigraphi­ Stratigraphic range: Older part of the Late Albian. cally subdivide the Upper Cretaceous deposits of the AdCP on the basis of foraminiferal assemblages. Discussion: This foraminiferal assemblage has a mixed character, since it is composed of taxa of wider 3.2.6.1. Conicorbitolina conica/Conicorbitolina stratigraphic range through the Early Cretaceous, cuvillieri partial–range zone or Conicorbitolina conica/ together with index fossils and taxa which first Conicorbitolina cuvillieri–Chrysalidina gradata appeared in the Albian and continued into the Late interval zone Cretaceous. Index fossils for this zone are orbi­ Boundaries: from the first occurrence of C. conica and/ tolinid species ‘V.’ dercourti, N. insolita, S. broen­ or C. cuvillieri to the first occurrence of C. gradata. 34 Geologia Croatica 60/1

Cenomanian Turonian Campanian Maastrichtian Coniacian Santonian

2−4 9.1

Pseudorhipidionina murgiana

Fig. 11 Biostratigraphic subdivision of the Upper Cretaceous deposits of the AdCP. Legend: 1) Conicorbitolina conica/Conicorbitolina cuvillieri partial-range zone; 2–4) Chrysalidina gradata superzone: 2) Chrysalidina gradata partial-range zone; 3) Pastrikella balcanica–Conicorbi- tolina conica concurrent-range zone; 4) Vidalina radoicicae–C. gradata concurrent-range zone; 5) Chrysalidina gradata–Pseudocyclam- mina sphaeroidea interval zone; 6) Pseudocyclammina sphaeroidea–Scandonea samnitica interval zone; 7) Scandonea samnitica–Dicy- clina schlumbergeri interval zone; 8) Dicyclina schlumbergeri–Murgella lata interval zone; 9) Murgella lata partial-range zone; 9.1) Kera- mosphaerina tergestina abundance subzone; 10) Calveziconus lecalvezae taxon-range zone; 11) Calveziconus lecalvezae–Murciella cuvillieri interval zone; 12) Murciella cuvillieri and Rhapydionina liburnica assemblage zone; 12.1) Fleuryana adriatica taxon-range sub- zone. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 35

Index and important taxa: C. conica, C. cuvillieri, C. correlation of the stratigraphic ranges, and compari­ gr. corbarica–conica, Orbitolina gr. concava, O. gr. son with other Cenomanian foraminifera, it is pos­ sefini–concava, ?Sellialveolina viallii, Ovalveolina sible to conclude a very precise chronostratigraphic maccagnoae, O. crassa, Praealveolina simplex, P. definition for this zone in the early Middle Cenoma­ iberica, Cisalveolina lehneri, Pseudocyclammina nian. rugosa, Pastrikella biplana and Trocholina arabica. Within older Middle Cenomanian deposits of N Stratigraphic range: Early Cenomanian. Istria, VELIĆ & VLAHOVIĆ (1994) proposed an Discussion: The orbitolinids C. conica, C. cuvillieri, earlier Chrysalidina gradata partial-range zone and C. gr. corbarica–conica, O. gr. concava and O. gr. younger Chrysalidina gradata and Pastrikella bal­ sefini–concava were determined in the assemblage, canica assemblage zone. They divided the earlier together with index forms of Cenomanian alveo­ zone into two subzones: an older, C. gradata with linids, O. crassa, P. simplex and P. iberica. The orbitolinids, and younger, C. gradata without orbi­ stratigraphic ranges of these taxa were according to tolinids. During later investigations in other parts of SCHROEDER & NEUMANN (1985), undoubtedly the AdCP, partly published in a paper by HUSINEC defined as Early Cenomanian. et al. (2000), as well as during detailed investiga­ tions for this study, Cenomanian orbitolinids were 3.2.6.2–3.2.6.4. Chrysalidina gradata superzone discovered in deposits of ages up to the end of the Boundaries: stratigraphic range of Chrysalidina gra­ Middle Cenomanian in different parts of the plat­ data. form: it seems that their absence in some places is Index and important taxa: Cited in descriptions of the locally restricted. Therefore, it seems that definition following three zones. of three zones within the Middle and Late Cenoma­ nian, as proposed here, would be best. Stratigraphic range: Middle and Late Cenomanian. Discussion: C. gradata is probably the most widespread 3.2.6.3. Pastrikella balcanica–Conicorbitolina conica foraminiferal species within Middle and Upper concurrent-range zone (older part of the zone Cenomanian deposits of the AdCP. The chronostrati­ Broeckina balcanica FLEURY, 1980). graphic value of this species has been confirmed in Boundaries: From the first occurrence of P. balcanica numerous publications in the last two decades (e.g. to the last occurrence of C. conica. GUŠIĆ et al., 1988; FUČEK et al., 1990; GUŠIĆ Index and important taxa: Most of the species as in & JELASKA, 1990; VELIĆ & VLAHOVIĆ, 1994; the previous zone (Fig. 13), accompanied by P. bal­ HUSINEC et al., 2000; KORBAR et al., 2001). canica. Therefore, within these deposits, i.e. within its Stratigraphic range: Late Middle Cenomanian. stratigraphic range, one superzone, and at least three clearly differentiated zones can be defined: Discussion: P. balcanica is, together with C. gradata, the most common, and widely distributed index – Chrysalidina gradata partial-range zone, fossil within the Middle and Upper Cenomanian – Pastrikella balcanica–Conicorbitolina conica con­ deposits of the AdCP. Its first occurrence represents current-range zone, and an important biohorizon at the beginning of the – Vidalina radoicicae–C. gradata concurrent-range younger part of Middle Cenomanian deposits. This zone. provides a basis for the definition of this biozone, which is, according to its foraminiferal assemblage 3.2.6.2. Chrysalidina gradata partial-range zone or and stratigraphic position in the late Middle Cenom­ Chrysalidina gradata–Pastrikella balcanica interval zone anian, equivalent to the C. gradata and P. balcanica assemblage zone of VELIĆ & VLAHOVIĆ (1994). Boundaries: From the first occurrence of C. gradata to HUSINEC et al. (2000) have defined a maximum of the first occurrence of P. balcanica. the development (and abundance) of Conicorbitoli­ Index and important taxa: C. gradata, C. cuvillieri, na conica in the Middle Cenomanian limestones of C. conica, C. gr. corbarica–conica, O. gr. concava, the island of Cres, which are stratigraphically equiv­ O. gr. sefini–concava, P. iberica, O. crassa, C. leh­ alent to the top part of this zone. neri, Nummoloculina? regularis, Pseudolituonella reicheli, Pseudorhapydionina dubia, Merlingina 3.2.6.4. Vidalina radoicicae–Chrysalidina gradata cretacea, P. biplana and Peneroplis turonicus. concurrent-range zone (younger part of the P. balcanica zone after FLEURY, 1980; C. gradata and V. Stratigraphic range: Early Middle Cenomanian. radoicicae assemblage zone or P. balcanica and Discussion: Most of the Cenomanian orbitolinids still N.? regularis abundance zone after VELIĆ & occurred in the first part of the stratigraphic range VLAHOVIĆ, 1994). of C. gradata together with other index fossils of Boundaries: From the first occurrence of V. radoicicae the Middle–Late Cenomanian, including N.? regu­ to the last occurrence of C. gradata or range of the laris, P. turonicus and P. reicheli, continuing up to maximum abundance of P. balcanica and N.? regu­ the first occurrence of P. balcanica. By analysis and laris. 36 Geologia Croatica 60/1

Index and important taxa: C. gradata, V. radoici­ 3.2.6.7. Scandonea samnitica–Dicyclina cae, Nummoloculina? regularis, Pseudolituonella schlumbergeri interval zone or Scandonea reicheli, Pseudorhapydionina dubia, P. laurinen­ samnitica partial-range zone sis, Merlingina cretacea, Cisalveolina fraasi, Pseu­ Boundaries: From the first occurrence of S. samnitica dorhipidionina casertana, P. murgiana, Peneroplis to the first occurrence of D. schlumbergeri. turonicus and Peneroplis parvus. Important taxa: S. samnitica, P sphaeroidea, M. com­ Stratigraphic range: Late Cenomanian. pressa and M. apenninica. Discussion: The C. gradata, P. balcanica and V. radoi­ Stratigraphic range: Early Coniacian. cicae assemblage zone was defined within the Discussion: This zone has no index fossils: it contains Upper Cenomanian carbonates of N Istria by VELIĆ forms determined in the previous zone as well as & VLAHOVIĆ (1994). They noted that this zone some without biostratigraphic significance. could also have been defined as the P. balcanica and N. regularis abundance or acme-zone, because these 3.2.6.8. Dicyclina schlumbergeri–Murgella lata interval zone or Dicyclina schlumbergeri species had their maximum frequency within this partial-range zone biozone, although their stratigraphic range is wider than the Late Cenomanian. Within the foraminiferal Boundaries: From the first occurrence of D. schlum­ assemblage, cited index forms have variable strati­ bergeri to the first occurrence of M. lata. graphic ranges, and only V. radoicicae has a strati­ Important taxa: S. samnitica, P sphaeroidea, M. apen­ graphic range throughout the zone. However, on the ninica, M. compressa, D. schlumbergeri, Accordiel­ basis of the former investigation it seems that this la conica and Idalina antiqua. fossil had a relatively poor distribution over the plat­ Stratigraphic range: Coniacian. form, and so was not used to define this zone as a Discussion: This zone represents the most of Conia­ taxon-range zone. A similar situation occurs for the cian deposits without forms of narrow stratigraphic other index fossils, including C. fraasi, P. laurinen­ range, but which lies between two important bio­ sis, P. casertana, P. murgiana and P. parvus. horizons, between the first occurrences of D. sch­ lumbergeri and M. lata, two frequent larger benthic 3.2.6.5. Chrysalidina gradata–Pseudocyclammina sphaeroidea interval zone foraminifera on the AdCP. Boundaries: from the last occurrence of C. gradata to 3.2.6.9. Murgella lata partial-range zone or Murgella the first occurrence of P. sphaeroidea. lata–Calveziconus lecalvezae interval zone Important taxa: The foraminiferal assemblage is sig­ Boundaries: From the first occurrence of M. lata to the nificantly reduced due to temporary drowning of the first occurrence of C. lecalvezae. platform, so there are no index fossils. Only taxa Index and important taxa: Murgella lata, Pseudo­ of wider stratigraphic range occurred, cited in the rhapydionina mediterranea, Keramosphaerina terg­ previous section (and shown on Fig. 6), as well as estina, K. ?sarda, S. samnitica, Reticulinella fleuryi, planktonic foraminifera in pelagic deposits. R. reicheli, D. schlumbergeri, A. conica, Broekinella Stratigraphic range: Latest Cenomanian–Early Turo­ neumannae, M. apenninica, M. compressa, Pseudo­ nian. cyclammina massiliensis, P. sphaeroidea, Pseudol­ Discussion: Drowning of the platform also interrupted ituonella mariae, Archiacina munieri, Dictyopsella the evolution of the platform benthic foraminifera. kiliani, Dictyopselloides cuvillieri, I. antiqua and Consequently, this zone comprises only forms of a Orbitoides gr. hottingeri. wider stratigraphic range. However, its boundaries Stratigraphic range: Early Santonian–Early Campa­ are clearly defined by two important biohorizons, nian. the last occurrence of C. gradata and the first occur­ Discussion: Deposits within this zone contain one of rence of P. sphaeroidea and Moncharmontia apen­ the richest, and biostratigraphically and palaeo­ ninica. geographically most important Upper Cretaceous foraminiferal assemblages. The most important 3.2.6.6. Pseudocyclammina sphaeroidea–Scandonea species is M. lata, accompanied by almost all the samnitica interval zone or Pseudocyclammina sphaeroidea partial-range zone forms cited in the previous two zones, together with newly occurring index forms – P. mediterranea, K. Boundaries: From the first occurrence of P. sphaeroi­ tergestina, P. massiliensis, P. mariae, R. fleuryi, R. dea to the first occurrence of S. samnitica. reicheli, D. kiliani, D. cuvillieri, A. munieri, B. neu­ Important taxa: P. sphaeroidea and M. apenninica. mannae and O. gr. hottingeri. Stratigraphic range: Late Turonian–earliest Coniacian The palaeogeographic significance of this zone is Discussion: There are no index fossils; P. sphaeroidea that it represents the youngest Cretaceous deposits occurs from the beginning of the zone, while M. covered by Eocene deposits over the major part of apenninica and M. compressa appear in younger the Adriatic Carbonate Platform in NE Istria, S and levels. E Lika, Dalmatia and Herzegovina. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 37

3.2.6.9.1. Keramosphaerina tergestina Important taxa: M. apenninica, M. compressa, D. sch­ abundance subzone lumbergeri, P. sphaeroidea, P. douvillei, P. vidali, Boundaries: stratigraphic range of the maximum abun­ O. douvillei, O. media and N. cretacea. dance of K. tergestina. Stratigraphic range: Latest Campanian–Early Maas­ Index and important taxa: Most of the taxa cited in trichtian. the previous zone (Fig. 11). Discussion: Deposits of this zone have been found near Stratigraphic range: Late Santonian–Early Campa­ the platform margins and slopes of the AdCP, while nian. the major part of it was emergent. These depos­ Discussion: In the younger part of this zone the level its crop out e.g. in S Croatia, on the island of Brač, with frequent and/or abundant occurrences of larger where GUŠIĆ & JELASKA (1990) determined K. tergestina is especially important, enabling sepa­ foraminiferal assemblages with predominant orbi­ ration of these deposits in the subzone. toids from detailed studied sections. There are no index fossils specific for this zone. 3.2.6.10. Calveziconus lecalvezae taxon-range zone 3.2.6.12. Murciella cuvillieri and Rhapydionina Boundaries: Stratigraphic range of C. lecalvezae. liburnica assemblage zone Index and important taxa: C. lecalvezae, most of the Boundaries: from the first occurrence of M. cuvillieri taxa cited in the previous zone (Fig. 11) and Num­ to the last occurrence of R. liburnica. mofallotia cretacea, Orbitoides tissoti, O. douvillei, O. media, P. douvillei and Pseudosiderolites vidali. Index and important taxa: Murciella cuvillieri, M. renzi, Rhapydionina liburnica, Cuneolina ketini, Stratigraphic range: Campanian. Fleuryana adriatica, Orbitoides media, Siderolites Discussion: Index fossils are C. lecalvezae and O. tis­ calcitrapoides, Omphalocyclus macroporus and soti. Within this zone, many of the important Late Hellenocyclina beotica. Santonian–Campanian species disappeared: K. terge­ Stratigraphic range: Maastrichtian. stina, M. lata, A. munieri, P. mariae, B. neumannae, and O. gr. hottingeri (Fig. 13). Most of the orbitoids Discussion: Deposits of this zone were studied on the continued into the next zone, and O. media occurs island of Brač (GUŠIĆ & JELASKA, 1990), where near the end of this zone. in the studied sections Murciella gr. cuvillieri occurs before Rhapydionina liburnica. An orbitoidid and There are only a few areas on the AdCP with con­ siderolitid assemblage was determined in slope tinuous depositional sequences and foraminiferal facies of the platform margins, and also locally in assemblages which could enable detailed biostrati­ the area of intraplatform troughs (e.g. VELIĆ, 1973; graphic zonation within these youngest Cretaceous VELIĆ et al., 1982; GUŠIĆ & JELASKA, 1990; deposits. Different environments existed within VESELI, 1999; DRAGIČEVIĆ & VELIĆ, 2002). these spatially restricted depositional areas, repre­ senting an additional difficulty for biostratigraphic 3.2.6.12.1. Fleuryana adriatica taxon-range subzone zonation and correlation. Therefore this zone con­ Boundaries: Stratigraphic range of F. adriatica. tains foraminifera from different biotopes, e.g. O. tissoti, O. douvillei, O. media, P. douvillei and P. Index and important taxa: F. adriatica, R. liburnica, vidali from platform margin and slopes and R. libur­ D. schlumbergeri and Laffitteina? mengaudi. nica, P. mediterranea, S. samnitica, P. massiliensis Stratigraphic range: Late Maastrichtian. and P. sphaeroidea from subtidal environments. It Discussion: Deposits of this subzone were observed in is difficult to precisely correlate the stratigraphic S Croatia (island of Brač), in the central Adriatic in ranges of taxa from both groups. GUŠIĆ & JELAS­ the Kate–1 well, as well as in the NW part of the KA (1990) discussed in detail the difficulties with platform in SW Slovenia, and in the vicinity of Tri­ facies interpretation with a simultaneous occurrence este in Italy (DE CASTRO et al., 1994). Within the of hyaline and imperforate foraminifera. This was deposits with abundant F. adriatica, DROBNE et based on the relationship between the Campanian al. (1989) discovered two very important species of Pučišća formation and the Maastrichtian Sumartin larger foraminifera, R. liburnica and D. schlumber­ formation on the island of Brač. Each attempt to geri, while JURKOVŠEK et al. (1996) only cited R. produce a more detailed biozonation in cases like liburnica. At the type locality of F. adriatica within this would be uncertain and speculative, and would the Upper Maastrichtian deposits on the island of only increase possible mistakes. Therefore, the best Brač, DE CASTRO et al. (1994) mentioned only or the least erroneous choice would be to unite taxa Laffitteina sp. and Bolivinopsis sp. R. liburnica and in assemblages within assemblage zones. Laffitteina mengaudi were discovered earlier in the 3.2.6.11. Calveziconus lecalvezae–Murciella cuvillieri same deposits (GUŠIĆ & JELASKA, 1990). interval zone Boundaries: from the last occurrence of C. lecalvezae to the first occurrence of M. cuvillieri. 38 Geologia Croatica 60/1

4. PALAEOBIOGEOGRAPHY OF MESOZOIC assic deposits GRGASOVIĆ (2003) mentioned the BENTHIC FORAMINIFERA OF THE KARST beginning of the stratigraphic ranges of some species of DINARIDES involutinids – Aulotortus sinuosus from the Late Ani­ sian and A. praegaschei from the Late Ladinian. The limited platform area and incomplete micropalae­ During a considerable part of the Late Triassic, i.e. ontological investigation are two important factors that during the Carnian and Early Norian, the major part of hinder construction of a complete and detailed pal­ the investigated area was emergent. Therefore, depos­ aeobiogeography of benthic foraminifera on the AdCP. its containing shallow-marine benthos occurred only Taking into account that the AdCP was about 700 km within marginal areas, e.g. around Karlovac speci­ long and probably 300–400 km wide, it is clear that mens of Lamelliconus – L. multispirus and L. procerus it represented a relatively small part of large shallow- occured (GRGASOVIĆ, 1997), as well as L. procerus marine areas of Tethys. Within the continuous dynamic in the area of Komiža on the island of Vis (BELAK et succession of events, the Adriatic Carbonate Platform al., 2005). evolved from a position in the southern, marginal area Assemblages within Upper Norian deposits contain of Tethys during the Early Jurassic, through an indi­ the following involutinids: Aulotortus friedli, A. tenuis, vidualized oceanic platform in the Central Tethyan area A. sinuosus, A. praegaschei and A. tumidus. These were with a probable connection to the Albanian and Hellenic accompanied in the Rhaetian by Triasina hantkeni and platforms, to its disintegration in the Northern Tethyan Auloconus permodiscoides. Localities in the Croatian area during the Late Cretaceous. Its palaeogeographic part of the Karst Dinarides where these are observed position controlled climatic changes, sedimentary envi­ include the area of Karlovac (BUKOVAC et al., 1974, ronments and ecological conditions, which directed the 1984; GRGASOVIĆ, 1997, 2003), the Gorski Kotar palaeobiogeography of the foraminiferal taxa, groups or (SAVIĆ & DOZET, 1985), Ogulin (VELIĆ et al., 1982), complete assemblages. Given the aforementioned lim­ vicinity of Slunj (KOROLIJA et al., 1981), S Velebit ited extent and moderate knowledge, the distribution of Mt. and Lika, Svilaja Mt. (BUCKOVIĆ et al., 2003) mostly larger foraminifera, especially index forms, will and the vicinity of Ston (RAIĆ & PAPEŠ, 1982). In be discussed and presented here. This review will cover Slovenia they have been found in the vicinity of Tolmin Mesozoic assemblages, becoming more detailed after (BUSER, 1986). In Bosnia and Herzegovina and Mon­ the Early and Middle Triassic, through the Late Triassic tenegro the aforementioned assemblages were found and Early Jurassic (when the AdCP became separated along NE and SE margin of the AdCP (MOJIČEVIĆ et from the Apulian and Apenninic platforms), to the end al., 1977, 1979; MARINKOVIĆ & ĐORĐEVIĆ, 1981; of Cretaceous. The process of platform separation prob­ VRHOVČIĆ & MOJIČEVIĆ, 1983; VUJNOVIĆ, ably started in the Late Triassic, and was completed 1981; PAPEŠ, 1975; SOFILJ et al., 1980; MIRKOVIĆ, during the Pliensbachian and Toarcian. Since then, the 1980; VUJISIĆ, 1975; ŽIVALJEVIĆ et al., 1971, AdCP represented a specific palaeogeographic entity 1989; KALEZIĆ et al., 1973; ĐOKIĆ et al., 1976; until its final disintegration by the end of Cretaceous. MIRKOVIĆ et al., 1978). Therefore, it may be con­ cluded that this assemblage was very widespread in the 4.1. Triassic Karst Dinarides deposits in the immediate basement of the Adriatic Carbonate Platform. Among Early Triassic foraminifera, Meandrospira pusila was the most widespread and the most frequent, together with abundant, though often poorly studied 4.2. Jurassic specimens of other smaller foraminifera, including There are no specific issues with the palaeobiogeo­ Glomospira sygmoidalis, Glomospirella triphonensis, graphy of the Early Jurassic foraminifera (Fig. 12). Ammodiscus parapriscus, Arenovidalina sp., etc. These Assemblages of the Early Jurassic lituolids are known forms were discovered in all areas with outcrops of from all areas where lower and middle Lower Jurassic Lower Triassic rocks, e.g. SW Slovenia, around Kar­ deposits crop out, so it may be concluded that these lovac and Slunj, in Velebit Mt., Lika, and Dalmatia in foraminifera were spread throughout the area of the Croatia, SW Bosnia and Herzegovina and S Montene­ Karst Dinarides. The most widespread were renowned gro. assemblages of Early Jurassic lituolids with Mesoen­ Middle Triassic index forms were only observed dothyra sp. and Orbitopsella praecursor, O. primaeva, within Anisian deposits. These aresmaller foraminif­ Lituosepta recoarensis, Haurania deserta and Amijiella era Meandrospira dinarica, Pilammina densa and Pil­ amiji. Representatives of the biokovinids in the Karst amminella semiplana. These forms were widespread Dinarides (GUŠIĆ, 1977) were restricted to local occur­ throughout the Dinarides, including areas that repre­ rences, e.g. Bosniella oenensis in W Bosnia and the sented the basement of the future AdCP. As in other vicinity of Dubrovnik, Biokovina gradacensis in Biok­ Periadriatic regions (e.g. the Alps and the Carpathians), ovo Mt. and Dubrovnik area. Pseudocyclammina lias­ there are no index foraminifera for the Ladinian in the sica was found at several localities in Croatia (W Gor­ Dinarides, since all forms (mostly the smaller foraminif­ ski Kotar, Velika Kapela, Velebit, Biokovo and vicinity era) had wider stratigraphic ranges. Within Middle Tri­ of Dubrovnik), but it may be proposed that it has been Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 39

Fig. 12 Distribution of important foraminiferal taxa within the Lower Jurassic deposits of the AdCP. more widespread, since sampling of this level was very and frequency was controlled by palaeogeography. restricted. Distribution of Lituolipora termieri, shows More variable and rich assemblages were found in the a similar pattern as it is common in the middle part of marginal, mostly peritidal parts of the platform, less in the Lower Jurassic deposits of Velika Kapela Mt. (W the slightly deeper subtidal inner platform environments Croatia). (Fig. 13). Toarcian and Early Aalenian deposits of the Karst Mesoendothyra croatica was the most widespread Dinarides contain no larger foraminifera, as a conse­ Middle Jurassic foraminifera, with a stratigraphic range quence of unfavourable ecological conditions, since the from the Late Toarcian to the Early Bathonian. It spread very shallow marginal parts of the platform were cov­ all over the AdCP, and in places is very abundant (e.g. ered by ooid–intraclastic bars, and the inner parts had in the vicinity of Dubrovnik). For years this foraminif­ lagoonal environments with low sedimentation rates era was the only index fossil in Middle Jurassic rocks of fine-grained deposits. Involutina liassica seems older than the Bathonian, especially in the central and restricted to the NE marginal areas of the platform western parts of the platform. Gutnicella cayeuxi had towards the Slovenian and Bosnian basin, in Karlo­ been described (ANIĆ, 1962; RADOIČIĆ, 1966) prior vac area (RADOIČIĆ, 1966; GUŠIĆ & BABIĆ, 1970; to the description of M. croatica (GUŠIĆ, 1969a). This DRAGIČEVIĆ & VELIĆ, 2002), northern Herzegovina foraminifera is much rarer than M. croatica, being com­ (SOFILJ et al., 1980), Montenegro (RADOIČIĆ, 1966; mon only in the SE part of the platform (S Croatia and MIRKOVIĆ, 1980; VUJISIĆ, 1975; ŽIVALJEVIĆ et Montenegro; Fig. 13). Aalenian–Bajocian foraminifera, al., 1989), and on the slopes towards the Budva–Cukali including Timidonella sarda, Spiraloconulus gigan­ trough (MIRKOVIĆ et al., 1978; ANTONIJEVIĆ et al., teus and S. perconigi are new discoveries on the AdCP. 1973). They were observed in two areas: the NW part of the The palaeobiogeography of the Middle Jurassic platform, SE of Ljubljana (Krka, Slovenia), and in the foraminifera is much more interesting. Their occurrence SE part in Biokovo Mt., and the vicinity of Dubrovnik 40 Geologia Croatica 60/1

Fig. 13 Distribution of important foraminiferal taxa within the Middle Jurassic deposits of the AdCP. in Croatia (Fig. 13), together with G. cayeuxi and M. mesojurassica and Pseudoeggerella elongata in Istria croatica. Isolated occurrences of G. cayeuxi were and Velebit Mt. (Fig. 13). These occur together with also observed in Gorski Kotar and SW of Karlovac in the somewhat more frequent Kilianina blancheti and Croatia, the vicinity of Jajce in Bosnia, as well as in Pfenderella arabica. However, within this assemblage Montenegro – in the vicinity of Nikšić and the coastal the most important and most widespread forms on the area. Within Middle Jurassic deposits, Haurania deser­ AdCP are those that are also very frequent in the Medi­ ta and Amijiella amiji occur in places. These were once terranean area – Paleopfenderina salernitana, P. tro­ considered to be index fossils of the middle part of the choidea and Satorina apuliensis. Early Jurassic. Pseudodictyopsella jurassica and Mar­ Forms having wider stratigraphic ranges through zoella ficcarellii are first documented here for the AdCP either the entire or a larger part of the Middle Jurassic in the area of the Biokovo Mt. This represents the sec­ frequently occur. These include Redmondoides lugeoni, ond occurrence of these forms in the World, after initial Siphovalvulina variabilis, Pfenderella arabica and Pro­ discovery of P. jurassica in Oman (SEPTFONTAINE topeneroplis striata, and Praekurnubia crusei from the & DE MATOS, 1998), and M. ficcarellii in Italy (CHI­ Bathonian, and Mohlerina basiliensis and Kurnubia OCCHINI & MANCINELLI, 2000). P. maynci has jurassica from the Callovian. previously only been observed at the Aalenian–Bajo­ Discovery of species previously unknown on the cian transition on the AdCP, while in other parts of both AdCP are especially important for the palaeobiogeo­ major Neotethyan provinces it was described as being graphy of the Middle Jurassic foraminifera. Of these, of Bathonian and Callovian age (BASSOULLET et al., Spiraloconulus giganteus and Orbitammina elliptica, l985; BASSOULLET, 1997). are particularly interesting, as they had been consid­ Orbitammina elliptica occurs in an assemblage ered as forms inhabitating the Western Mediterranean with other Bathonian species, previously unknown palaeobiogeographic province (W Alps, Jura Mt., N on the AdCP, including Alzonella cuvillieri, Satorina Sardinia, Aquitaine and Paris basins), as well as some Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 41

Fig. 14. Distribution of important foraminiferal taxa within the Upper Jurassic deposits of the AdCP.

Eastern provinces (Caucasus and Turkmenistan). These iensis, Nautiloculina oolithica, Praekurnubia crusei and western and eastern provinces represented parts of the Protopeneroplis striata, while Labyrinthina mirabilis northern Neotethyan margin (BASSOULLET et al., and Parurgonina caelinensis are somewhat rarer (Fig. l985). Occurrences of S. giganteus (Fig. 13) in Krka 14). area (Slovenia), Biokovo Mt. and Dubrovnik, together Trocholina species are frequent, especially T. alpi­ with observations of O. elliptica (Fig. 13) in Istria and na, with a stratigraphic range from the Bajocian to the Velebit Mt. extend their distribution to the southern Late Hauterivian, and T. elongata from the Oxfordian. marginal areas of Neotethys, indicating the necessary They were the most abundant in the Oxfordian tem­ reinterpretation of traditional Middle Jurassic palaeobio­ pestites and carbonate beach sands of W Istria, Gorski geography. K. blancheti, S. mesojurassica and P. elon­ Kotar and Velebit Mt., in an assemblage with Chablai­ gata also occur in smaller areas of the western part of sia chablaisensis. In the central parts of the platform the platform, between W Istria and S Velebit Mt., while Ch. chablaisensis had only been reported at one locality Paravalvulina complicata was found for the first time in Biokovo Mt. (but was probably of Bathonian/Callo­ on the AdCP in the vicinity of Dubrovnik, and later in vian age), and in the Oxfordian of Lastovo island. Tro­ Biokovo Mt. and vicinity of Karlovac. cholina gigantea was only found in the Oxfordian of The most widespread Late Jurassic foraminif­ Istria and Gorski Kotar (W Croatia). era was Kurnubia palastiniensis, occurring in all parts Alveosepta jaccardi and Conicokurnubia orbito­ of the platform. Two other forms of Kurnubia were liniformis were discovered in the Kimmeridgian of the frequently cited under this name: K. wellingsi and K. Velika Kapela, Velebit and Biokovo Mts. in Croatia, jurassica. There is no consensus among micropalaeon­ so, at this point they can be considered as index fos­ tologists as to the validity of these citations. All three sils of that stage in the area of the AdCP. Three addi­ species are widespread on the AdCP. In addition to kur­ tional infrequent foraminifera, previously unknown in nubias, the most frequent species were Mohlerina basil­ the AdCP area, are Kilianina? rahonensis from the Late 42 Geologia Croatica 60/1

Kimmeridgian and Tithonian of W Croatia, and the Aptian), S. auruncensis, S. capitata and S. dinapolii Late Tithonian Anchispirocyclina lusitanica and/or A. range from the Late Aptian–Early Albian. neumannae, found only in S Croatia (Fig. 14). This list During the Late Barremian an orbitolinid assem­ could be completed by Everticyclammina virguliana, blage occurred on the AdCP. Rectodictyoconus gigan­ usually found within Upper Jurassic peri-reefal deposits teus is the oldest orbitolinid genera on the AdCP, pre­ of marginal parts of the platform. viously known only in the vicinity of Jajce and Banja Luka near the northern margin of the platform. 4.3. Cretaceous Comparison with the palaeogeography of orbi­ The Adriatic Carbonate Platform and neighbouring tolinids found on Perimediterranean platforms indicates more or less isolated Central Mediterranean carbonate differences between the AdCP and Western Mediterra­ platforms (e.g. Apenninic, Apulian, Gavrovo–Tripolit­ nean platforms. Orbitolinid assemblages in the Western za, etc.) were all located within Southern Tethyan bio­ Mediterranean, i.e. Northern Tethyan provinces, devel­ provinces. They were characterized by the reduced oped in the earliest Cretaceous. Examples include the occurrence of larger foraminifera from the Berriasian to Valanginian of the Pyrenees (Valdanchella, Paracoski­ the Late Barremian. In contrast, in the Western Mediter­ nolina etc., according to PEYBERNES, 1976), Haute­ ranean, carbonate platforms belonging to the Northern rivian of SE France (e.g. evolutionary lines Praedicty­ Tethyan bioprovinces (Cantabria, Pyreneans, Sardinia, orbitolina–Dictyorbitolina and Valserina–Palorbitolina Vercors, Chartreuse, Jura Mt., etc.) larger foraminifera (according to CLAVEL et al., 1995 and ARNAUD et were present in the Berriasian, Valanginian and Haute­ al., 1998) and the Barremian of the aforementioned are­ rivian. Species include Keramosphaera allobrogensis as (Eopalorbitolina–Palorbitolina after SCHROEDER STEINHAUSER, BRÖNNIMANN & KOEHN-ZANI­ et al., 2002 or Paracoskinolina, Urgonina, Alpillina, NETTI, Eclusia moutyi SEPTFONTAINE, E. decastroi Cribellopsis and diverse species of genus Orbitolinop­ CHERCHI & SCHROEDER and Valdanchella miliani sis according to, e.g. ARNAUD et al., 1998). (SCHROEDER), as well as other orbitolinids which A rich and explosive development of orbitolinids were completely missing in the contemporaneous on the Adriatic Carbonate Platform started in the latest deposits in the AdCP area. Barremian/earliest Aptian, when varied assemblages In the Neocomian and Early Barremian deposits of of taxa especially abundant in the Northern Tethyan the AdCP foraminiferal assemblage including Trocholi­ provinces (like palorbitolinids and praeorbitolinids, as na gr. alpina–elongata, Charentia cuvillieri, Mayncina well as mesorbitolinids in the Late Aptian and Albian), bulgarica and Praechrysalidina infracretacea is char­ became distributed throughout the platform. In contrast acterized by a modest stratigraphic value, and the first to R. giganteus, only discovered in the central part of index fossils occur in the orbitolinid assemblages of the NE AdCP margin in Upper Barremian deposits (Fig. the Late Barremian. However, many small foraminif­ 16), Palorbitolina lenticularis was found all over the era represent index fossils, e.g. for the Valanginian Ver­ AdCP. Some authors (VELIĆ & SOKAČ, 1978a) indi­ corsella camposaurii, V. tenuis, Montsalevia salevensis, cate that this species occurred on the AdCP at the very Haplophragmoides joukowskyi, or Late Hauterivian end of the Barremian, i.e. somewhat later than in the Campanellula capuensis. All these taxa were distributed Western Mediterranean, where it occurred in the Middle throughout the AdCP. Barremian (e.g. SCHROEDER, 1964c; SCHROEDER Protopeneroplis ultragranulata is the index spe­ et al., 1974; PEYBERNES, 1976; ARNAUD-VAN­ cies of transitional Tithonian–Berriasian and Berria­ NEAU, 1980; JAFFREZO, 1980; CLAVEL et al., 1995; sian deposits, and was previously only known from the ARNAUD et al., 1998). On the Apenninic Carbonate coastal part of Southern Croatia (Biokovo Mt., vicinity Platform (which should have been, according to the of Dubrovnik). Representatives of the genus Trocholi­ palaeogeographic interpretations, located between the na, for example, T. gr. alpina–elongata, as well as T. Western Mediterranean platforms and the AdCP dur­ sagittaria, T. delphinensis, and probably also T. molesta ing the Aptian), deposits with palorbitolinids are either occur throughout the platform area. In the Barremian of Upper Barremian–Lower Aptian age (CHIOCCHINI and Early Aptian of the Western parts of the platform, et al., 1994) or Lower Aptian age (CHERCHI et al., T. lenticularis was locally found, and mostly undeter­ 1978). minable small involutinids are also frequent. The palaeobiogeography of Early Aptian foramini­ The entire platform area is also characterized by the fera on the AdCP was very controlled by the palaeoen­ occurrence of representatives of Vercorsella, Sabaudia vironments. Generally, in this area, this period was and Novalesia with V. camposaurii, V. tenuis (the index characterized by sea-level rise and pelagic influences, fossil of the Valanginian and Early Hauterivian), V. represented by common findings of Hedbergella sp., scarsellai, V. laurentii, V. arenata and V. wintereri, N. Saccocoma sp. etc. (VELIĆ & SOKAČ, 1978a). These cornucopia, N. distorta, N. producta and S. minuta. All forms only occur in very restricted subtidal facies and of these have a wider stratigraphic range through the lagoons together with Choffatella decipiens, in the cen­ Early Cretaceous. S. briacensis is always found together tral and western parts of the platform. with palorbitolinids (and is the index fossil of the Early Dominant Early Aptian foraminifera are orbitoli­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 43

Fig. 15 Distribution of orbitolinids within the Cretaceous deposits of the AdCP. nids from the evolutionary line Praeorbitolina cormyi Mesorbitolinid assemblages continued into the Ear­ – P. wienandsi – Mesorbitolina lotzei. They occur in ly Albian. They were widespread, and known from all the W, central and SE parts of the platform (VELIĆ & parts of the platform. Therefore, with the Early Aptian SOKAČ, 1978b; VELIĆ et al., 1979; HUSINEC et al., palorbitolinids, they represent not only the most fre­ 2000) (Fig. 15). Accompanying forms are Early Aptian quent orbitolinids, but also some of the most extensive index fossils found throughout the platform area: foraminifera on the entire AdCP. In addition, in the Late Voloshinoides murgensis (the most frequent), Sabau­ Aptian, M. parva and M. texana occur, continuing into dia briacensis, Derventina filipescui, Pseudolituonella the Early Albian, where they are accompanied by Mes­ conica, Neotrocholina aptiensis and N. friburgensis, as orbitolina subconcava and Mesorbitolina pervia. well as rare Pfenderina globosa. Earliest representa­ In addition to the Early Albian mesorbitolinids, tives of the Archalveolina genus (VELIĆ & SOKAČ, specifically undeterminable forms of the genera Orbito­ 1978a), Pseudonummoloculina aurigerica, as well as linopsis and Paracoskinolina, as well as other, previous­ all the aforementioned species of Sabaudia, Vercorsella, ly cited species of Vercorsella, Sabaudia, Novalesia etc. Novalesia, Nautiloculina, etc. occur. were also widespread throughout the platform. How­ In the Late Aptian, orbitolinid evolutionary trends ever, the predominant species in foraminiferal assem­ continued with Mesorbitolina parva and Mesorbitolina blages of the youngermost Lower Cretaceous deposits texana, found in all parts of the platform. These forms were Cuneolina pavonia, C. parva, Pseudonummolocu­ were accompanied by Archalveolina reicheli, locally lina heimi and Nezzazatinella picardi, with stratigraphic very abundant Haplophragmoides globosus (or Troc­ ranges through the entire Albian and a larger part of the hamminoides coronus according to CHIOCCHINI et Late Cretaceous. al., 1994), and, close to the end of Aptian, Orbitolinop­ In the Late Albian orbitolinid assemblages with sis aquitanica. All these forms were, previously only ‘Valdanchella dercourti’, Simplorbitolina broenni­ known from the coastal, Adriatic part of Croatia. manni, Neoiraquia insolita, Paracoskinolina fleuryi, 44 Geologia Croatica 60/1

Fig. 16 Distribution of alveolinids within the Cretaceous deposits of the AdCP. and Paracoskinolina sunnilandensis were frequent. dle Cenomanian deposits, where it is accompanied by Among other rare foraminifera were Scandonea phoe­ P. balcanica and other Middle Cenomanian species. nissa and S. pumila (e.g. at Biokovo Mt. and the island A group of index fossils of the Middle and Late Ceno­ of Korčula). For the very end of the Albian, Vraconian manian, comprising Chrysalidina gradata, Nummolo­ or the base of Cenomanian, occurrences of Neoiraquia culina? regularis, Pseudolituonella reicheli, Pastrikella convexa (in the vicinity of Dubrovnik and Biokovo Mt.) balcanica and Pseudorhapydionina dubia, was very and Protochrysalidina elongata (island of Cres), as well abundant in all parts of the platform (Fig. 17), as well as the more frequent Ovalveolina maccagnoae are typi­ as other taxa of a wider stratigraphic range cited within cal, though relatively rarely observed. the Cenomanian assemblages (cuneolinas, nezzaza­ During the Cenomanian, there was a successive tids, pseudonumoloculinas, soritids, peneroplids). The occurrence of alveolinids from Ovalveolina maccag­ Late Cenomanian Pseudorhapydionina laurinensis and noae and ?Sellialveolina viallii, through O. crassa, Cis­ Pseudorhipidionina casertana (POLŠAK et al., 1982; alveolina lehneri and Praealvelina iberica in the Early VELIĆ & VLAHOVIĆ, 1994) (Fig. 17), found in Istria, Cenomanian, P. simplex in the Middle, to Cisalveo­ Lika, and Dalmatia are rare. lina fraasi in the Late Cenomanian. These forms were The Early Turonian was characterised by restricted found, although not frequently, in different parts of foraminiferal assemblages due to platform drowning, so the AdCP (Fig. 16). The situation with the orbitolinids in a palaeobiogeographic sense it is not very interesting. Orbitolina gr. concava, Conicorbitolina cuvillieri, C. During the Late Turonian, the entire platform (except conica, C. gr. corbarica–conica and Orbitolina gr. sefi­ for minor parts emergent since at least the Late Ceno­ ni–concava was similar. These however, were extinct manian), was again covered by shallow-marine envi­ before the late Cenomanian, but were widespread all ronments with assemblages of cuneolinas, nezzazatids, over the platform (Fig. 15). Pastrikella biplana occurs miliolids and other groups of foraminifera known since in younger Lower Cenomanian beds and in older Mid­ the Albian and through the Cenomanian. New forms Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 45

Fig. 17 Distribution of several index and the more important foraminiferal species within the older Upper Cretaceous deposits (Cenoma- nian–Lower Santonian) of the AdCP. which were widespread include Moncharmontia apen­ mannae, Nummofallotia cretacea, Archiacina munieri ninica and Pseudocyclammina sphaeroidea. Cuneolina and Abrardia mosae were discovered in central Bosnia compressa was only found in the deep off-shore well (near Jajce). Along the platform slopes, as well as in the near the island of Palagruža. deeper parts of the inner platform, the previously men­ The composition of Coniacian and Early Santonian tioned forms were accompanied by the orbitoid species assemblages was very similar to those from the Late Orbitoides gr. hottingeri. Turonian, with the exception of newly occurring forms: Late Santonian species also continued into the Early Scandonea samnitica, Dicyclina schlumbergeri and Campanian, when they were accompanied by new ones. Moncharmontia compressa. Important changes took Calveziconus lecalvezae and Reticulinella reicheli were place in the Late Santonian with the occurrence of new found over the major part of the platform. However, species of larger foraminifera – Murgella lata, Pseudor­ they are most abundant along the Adriatic coast and hapydionina mediterranea, Idalina antiqua, Keramos­ on the islands. Along the platform margins and on the phaerina tergestina and Pseudocyclammina massilien­ slopes or in deeper areas within the inner parts of the sis, as well as several species of smaller foraminifera platform, the aforementioned assemblage was accom­ – Accordiella conica, Murgeina apula, Reticulinella panied by Praesiderolites douvillei, Orbitoides douvil­ fleuryi, Dictyopsella kiliani, Dictyopselloides cuvil­ lei, O. tissoti, O. media and Pseudosiderolites vidali lieri, Spirolina cretacea, Minouxia lobata, Pseudol­ (GUŠIĆ & JELASKA, 1990; Fig. 18). ituonella marie, Rotorbinella scarsellai and Stensiöina During the Maastrichtian margins, slopes and surrentina. All these forms were widespread over the inner platform deeper areas were inhabitated by abun­ AdCP (Fig. 18). Raadshovenia salentina occurs in the dant Orbitoides media, Siderolites calcitrapoides and Santonian deposits of the NW part of the platform (S Omphalocyclus macroporus. Murciella cuvillieri, M. Istria – POLŠAK et al., 1982, and in the area of Trieste renzi, Fleuryana adriatica and Laffitteina mengaudi Karst – VENTURINI, 2005), while Broekinella neu­ occur on the island of Brač (GUŠIĆ & JELASKA, 46 Geologia Croatica 60/1

Fig. 18 Distribution of several index and the more important foraminiferal species within the younger Upper Cretaceous deposits (Upper Santonian–Upper Maastrichtian) of the AdCP.

1990; DE CASTRO et al., 1994) and vicinity of Makar­ l985). This can be shown by comparison of the strati­ ska. Rhapydionina liburnica was found in areas contain­ graphic ranges of foraminifera and biozones within ing Maastrichtian deposits across the AdCP, accompa­ Lower Jurassic deposits of the High Atlas in Morocco nied by species with wider stratigraphic ranges through and Velebit Mt. in Croatia (SEPTFONTAINE, 1984; the Late Cretaceous, like C. pavonia, N. picardi, D. sch­ TIŠLJAR et al., 1991; SEPTFONTAINE et al., 1991), lumbergeri, M. apula, M. apenninica, A. conica and R. which are almost identical. The other Perimediterranean scarsellai. platforms in Italy, Greece and Turkey also belonged to South-Tethyan bioprovince during the Early and Mid­ 4.4. Correlation dle Jurassic. This palaeogeographic data on the AdCP can be corre­ A significant new result is the co-occurrence of lated with data from other Periadriatic carbonate plat­ Orbitammina elliptica and Satorina apuliensis in forms, as well as with previous knowledge on palaeobio­ the Bathonian deposits of Istria and Velebit. Namely, geographic provinces in Neotethys during the Jurassic these two forms are typical for two major Neotethyan and the Cretaceous. Results of these comparisons show provinces, O. elliptica for the Northern province, and important similarities, but also certain specificities of S. apuliensis for the Southern one (BASSOULLET the AdCP, which will not change the general concept of et al., l985). The discovery of O. elliptica within the the position of the AdCP in the Tethyan realm during foraminiferal assemblage of the Southern Neotethyan the younger Mesozoic. province (VELIĆ, 2005), as well as the very serious The composition of foraminiferal assemblages indications of its presence in the Bathonian deposits of definitely indicates the location of the AdCP within the Apenninic Carbonate Platform (LUPERTO-SINNI, the southern marginal part of this palaeoocean during 1968, pl./fig.: 7/1, 8/3, 9/2,3, 10/2, 11, 12), indicate that the Early and Middle Jurassic (BASSOULLET et al., its area of distribution included the northernmost parts Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 47 of the Southern Neotethyan bioprovince during the alveolinids Praealveolina cretacea (D’ARCHIAC), P. Bathonian. During the Late Cretaceous both provinces brevis REICHEL, Ovalveolina ovum (D’ORBIGNY) had more or less similar foraminiferal assemblages. and Pseudedomia drorimensis REISS, HAMAOUI & Abundant occurrences of Chablaisia chablaisensis in ECKER are also absent. However, it cannot be affirmed the Oxfordian of the AdCP, but only in its NW part are that the AdCP was still part of the Southern Neotethys interesting, because S of Velebit Mt. it has only been bioprovince during the Cenomanian, in spite of the sig­ found at one locality in the Bathonian of Biokovo Mt. nificant similarity of foraminiferal assemblages. For this (VELIĆ, 2005) and the island of Lastovo. Present data province, typical Near East species of larger foraminif­ also indicate the location of the AdCP within Southern era, including Cycledomia iranica HENSON, Taberina Neotethys during the Late Jurassic. bingistani HENSON, P. drorimensis and Thomasinella From the beginning of the Cretaceous to the Late punica SCHLUMBERGER (e.g. SAINT-MARC, 1974, Barremian, shallow-marine environments of the North­ 1977; HAMAOUI, 1979) have never been found on ern Neotethyan bioprovince were gradually inhabitated the AdCP. On the basis of the presented data, it may be by orbitolinids and other larger foraminifera (genera concluded that during the Cenomanian and the Turoni­ an, the AdCP was probably biogeographically isolated, Valdanchella, Praedictyorbitolina, Dictyorbitolina, i.e. that it was not part of either of the two major bio­ Valserina, Eygallerina, Eopalorbitolina, Montseciella, geographical provinces. This is indicated by the compo­ Eclusia, Keramosphaera, Pfenderina etc. – see e.g. sition of the foraminiferal assemblage, which is similar BASSOULLET et al., l985; CLAVEL et al., 1995; to assemblages of both bioprovinces, but lacking forms SCHROEDER et al., 2002). Contemporaneous depos­ characteristic of either. A similar view was proposed its of the AdCP contain no foraminifera. This indicates by FLEURY et al. (1985) for the Periadriatic–Aegean that at that time, the AdCP was still a part of the South­ provinces during the Late Cretaceous, i.e. the Adriatic ern bioprovince, and that there was still no connection Carbonate Platform, South Apenninic platforms (most between the two major Neotethyan bioprovinces. In likely Apulian and Apenninic), Hellenic platforms contrast, assemblages of small foraminifera found on (Gavrovo–Tripolitza) and Tauridic platforms (probably the AdCP containing index forms for the Valanginian Menderes). and Early Hauterivian, including Montsalevia salev­ ensis, Haplophragmoides joukowskyi, Meandrospira A Late Cenomanian–Early Turonian sea-level rise favrei, Vercorsella tenuis and V. camposaurii, are also caused temporary drowning of the AdCP (GUŠIĆ & known from the areas along the northern Neotethyan JELASKA, 1990, 1993; JENKYNS, 1991; JELASKA margin (where the first three cited species were origi­ et al., 1994; TIŠLJAR et al., 2002; VLAHOVIĆ et al., nally described by CHAROLLAIS et al., 1966). 2003, 2005), resulting in the temporary disappearance of shallow-marine environments favourable for biotopes In the Late Barremian and Early Aptian, orbitolinid of benthic foraminifera. They were later re-established assemblages rapidly spread over the Southern Neotethy­ due to shallowing in the Late Turonian, and lasted until an bioprovince, led by Palorbitolina lenticularis, which the end of the Cretaceous in parts of the platform cov­ was accompanied by praeorbitolinids – Praeorbitoli­ ered by the sea. na cormyi and P. wienandsi, as well as Mesorbitolina On the basis of foraminiferal assemblage composi­ lotzei during the Early Aptian. Therefore, since the Late tion in Upper Cretaceous deposits of Western Neotethys Barremian through the Aptian, Albian and Early Ceno­ (Upper Turonian to Maastrichtian), two bioprovinces manian, biofacies and biotopes of the larger foraminif­ can be clearly distinguished: Central Mediterranean, era on the AdCP gradually became more and more simi­ composed of the Periadriatic carbonate platforms – lar to those from the Northern Neotethyan bioprovince. Adriatic, Apenninic, Apulian and Gavrovo–Tripolitza, BASSOULLET et al. (1985) acknowledged that dur­ and Western Mediterranean, comprising carbonate plat­ ing the Late Aptian and Albian it was still possible to forms and shallow-marine areas of the present Pyrenees, clearly differentiate two bioprovinces within Neotethys: Aquitaine, Provence and Sardinia. Within foraminiferal a northern one with Simplorbitolina manasi and Neor­ assemblages of the Central Mediterranean bioprovince, bitolinopsis conulus, and a southern one with Archaeal­ on the AdCP and neighbouring platforms (e.g. LUPER­ veolina and Simplorbitolina ?brönnimanni. TO-SINNI, 1976; LUPERTO-SINNI & RICCHETTI, During the Cenomanian, composition of orbi­ 1978; FLEURY, 1980; FLEURY et al., 1985; GUŠIĆ & tolinid and alveolinid assemblages on the AdCP (e.g. JELASKA, 1990; CHIOCCHINI et al., 1994; CVETKO VELIĆ & VLAHOVIĆ, 1994; HUSINEC et al., 2000; TEŠOVIĆ et al., 2001, and Section 2 of this paper), TIŠLJAR et al., 2002) was similar, but not identical, there are no presently known occurrences of taxa typi­ to contemporaneous assemblages in SW Europe, i.e. cal for the Western Mediterranean bioprovince. Such areas belonging to the Northern Neotethyan province absent species include Orbitolinopsis senonicus GEN­ (e.g. SCHROEDER, 1962, 1964a, 1975; BILOTTE, DROT, Peneroplis giganteus GENDROT, Lacazina 1985; CASTRO et al., 2001). In the area of the AdCP, elongata SCHLUMBERGER, L. compressa MUNIER- there are presently no known occurrences of Mesorbi­ CHALMAS, Pseudolacazina cantabrica HOTTINGER, tolina aperta (ERMAN) and Orbitolina durandelgai DROBNE & CAUS, Ilerdorbis decussatus HOTTING­ SCHROEDER among the orbitolinids, while among the ER & CAUS, Broeckina dufrenoyi (D’ARCHIAC), 48 Geologia Croatica 60/1

Cyclopsinella steinmanni (MUNIER-CHALMAS), – for the first time the palaeobiogeography of larger Paleodictyoconus senonicus MOULLADE & VIAL­ foraminifera of the AdCP is proposed, which is espe­ LARD, Abrardia catalunica BILOTTE, Lamarmorella cially important for regional Mediterranean correla­ sarda CHERCHI & SCHROEDER, Periloculina zit­ tion, in which the AdCP was previously inadequately teli MUNIER-CHALMAS & SCHLUMBERGER, etc. represented or was completely excluded. (GENDROT, 1968, CHERCHI & SCHROEDER, 1977; BILOTTE, 1985; HOTTINGER et al., 1989). Likewise, The chronostratigraphic succession comprises species typical for the Central Mediterranean bioprov­ foraminiferal assemblages from the Early Triassic to ince, e.g. Pseudorhapydionina mediterranea, Murgella the end of the Cretaceous. Previous knowledge on the lata, Keramosphaerina tergestina, Murciella cuvillieri, assemblages within Lower and Middle Triassic depos­ M. renzi, Rhapydionina liburnica and Fleuryana adri­ its of the Adriatic Carbonate Platform indicate a prob­ atica, have never been observed in the Western Medi­ ably rich assemblage of genera and species, but reliable terranean bioprovince. Therefore, it is clear that these data could only be provided after detailed investigations two provinces were, according to the larger foraminif­ which have not yet been performed. At this moment it era, ‘isolated’ from each other. Each had their typical can be stated that smaller foraminifera prevailed within species and assemblages, in places with even endemic these deposits, mostly ammodiscids and endothyrids. characteristics (e.g. Catalonian subprovince, after HOT­ Index forms are: Cyclogyra mahayeri, Ammodiscus TINGER et al., 1989). The palaeogeographic position paraprisca, Meandrospira pusila, Glomospira sygmoi­ of these provinces was from the Turonian to the end of dalis and Glomospirella triphonensis for the Scythian, Cretaceous in Central Neotethys, because both prov­ and Meandrospira dinarica, Pilammina densa and Pil­ inces were placed between 23° and 35° N of equator amminella semiplana for the Anisian. Within Ladinian after DERCOURT et al. (2000). During this period, the deposits there are no index species of foraminifera, but Gondwanian coast of Neotethys (from Ethiopia/Erithrea the beginning of the stratigraphic ranges of some spe­ to Morocco), was placed between 5° S and 18° N in the cies of Aulotortus (A. praegaschei and A. sinuosus) can Cenomanian, and from the equator to the 20° N in the be presumed, as stated by GRGASOVIĆ (2003). The Maastrichtian. The European coastline was more stable, maximum abundance of these was in the Norian and and during the Late Cretaceous its position was between Rhaetian. An assemblage of involutinids, with Aulo­ 35° and 45° N (after DERCOURT et al., 2000). The tortus friedli, A. praegashei, A. sinuosus, A. tenuis, A. position of the AdCP, as the northernmost Periadriatic tumidus, was generally predominant in Upper Triassic platform in the Central Mediterranean bioprovince and deposits, and Aulotortus pokornyi, Trocholina crassa in the Late Cretaceous Neotethys, was between 23° and and T. acuta in the Norian and Rhaetian. Besides the 30° N. In such a palaeogeographic constellation, when cited species, discovery of the index forms Lamelli­ Neotethys was at its widest and most complex, includ­ conus multispirus and L. procerus in Carnian, Turrispi­ ing the beginning of its closure, it was characterized by rillina minima in the Norian, and Triasina hantkeni in the occurrence of more biogeographical provinces than the Rhaetian are very important. at any other time. Therefore, it may be concluded that in Within the Jurassic deposits, lituolid foraminifera the Late Cretaceous, the area of the Adriatic Carbonate predominate, which were not equally distributed in Platform represented part of the Central Mediterranean time. During the Early Jurassic, mesoendothyrids were bioprovince in the Central Neotethyan area. predominant with index forms including Mesoendothyra sp., Lituosepta recoarensis, Orbitopsella primaeva and Paleomayncina termieri in the Sinemurian, Mesoen­ 5. REVIEW OF THE RESULTS dothyra sp., L. recoarensis, O. primaeva, O. praecursor, P. termieri, Planisepta compressa, Lituolipora polymor­ The most important results of the investigation pre­ pha, Biokovina gradacensi and Bosniella oenensis in sented in this paper can be summarised in the following the Early Pliensbachian (Carixian), and O. praecursor, conclusions: O. ?dubari, Pseudocyclammina liassica, P. termieri, P. – this is the first paper synthesizing and summarizing compressa, B. gradacensis and B. oenensis in the Late the results of Mesozoic benthic foraminiferal study in Pliensbachian (Domerian). As a consequence of unfa­ the Karst Dinarides undertaken by the author during vourable palaeoecological conditions during the Toar­ almost 50 years of research, cian, a significant decline took place in the foraminif­ eral assemblage, and there were no index forms. Mid­ – numerous new results regarding foraminiferal assem­ dle Jurassic deposits contain the richest foraminiferal blages were achieved, including discovery of many assemblages with numerous taxa, including well known taxa previously unknown in the Karst Dinarides, index species – Gutnicella cayeuxi, Pseudocyclammina – a chronostratigraphic position is provided for forami­ maynci, Spiraloconulus perconigi, S. giganteus, Timi­ niferal assemblages, donella sarda in the Aalenian and Early Bajocian, S. – a completely new biostratigraphic zonation was pro­ giganteus, Pseudodictyopsella jurassica and Marzoella duced, respecting the results of previous investiga­ ficcarellii in the Bajocian, Paravalvulina complicata tions, at the end of Bajocian and beginning of the Bathonian, Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 49 as well as Pseudoeggerella elongata, Paleopfenderina bitolina conica, C. cuvillieri, Praealvelina iberica and salernitana, P. trochoidea, Alzonella cuvillieri, Orbita­ P. simplex in Early Cenomanian, Chrysalidina gradata, mmina elliptica, Kilianina blancheti, Satorina apuli­ Nummoloculina? regularis, Pseudolituonella reicheli, ensis and S. mesojurassica in the Bathonian. Among Pastrikella balcanica, Conicorbitolina conica, C. gr. the cited species P. maynci, S. perconigi, S. giganteus, corbarica–conica, Orbitolina gr. sefini–concava, Pseu­ T. sarda, P. complicata, P. elongata, A. cuvillieri and dorhapydionina dubia and Peneroplis turonicus in O. elliptica had not been previously observed on the the Middle Cenomanian, C. gradata, N.? regularis, P. AdCP. Occurrences of P. jurassica and M. ficcarellii reicheli, P. balcanica, P. dubia, Cisalveolina fraasi, are not only the first on the AdCP, but probably the first Vidalina radoicicae, V. hispanica, Pseudorhapydionina observations outside the type localities. Upper Jurassic laurinensis, Pseudorhipidionina casertana and Pen­ deposits are characterized by numerous specimens of eroplis parvus in the Late Cenomanian, Pseudorhapy­ some taxa, but diversification was less pronounced, as dionina mediterranea, Murgella lata, Keramosphaerina well as there being a low number of index species. The tergestina, Pseudocyclammina massiliensis, Pseudol­ most frequent are kurnubias – Kurnubia jurassica, K. ituonella marie, Spirolina cretacea, Minouxia lobata, palastiniensis and K. wellingsi, and index fossils were Broekinella neumannae, Reticulinella fleuryi, Dicty­ Labyrinthina mirabilis, Alveosepta jaccardi, Parur­ opsella kiliani, Dictyopselloides cuvillieri and Orbi­ gonina caelinensis and Kilianina? rahonensis. Conico­ toides gr. hottingeri in the Late Santonian and Early curnubia orbitoliniformis, Anchispirocyclina lusitanica Campanian, P. mediterranea, K. tergestina, P. mas­ and/or A. neumannae are documented for the first time siliensis, Reticulinella reicheli, R. fleuryi, D. kiliani, on the AdCP. D. cuvillieri, Calveziconus lecalvezae, Nummofallotia Cretaceous assemblages are much richer than Trias­ cretacea, Praesiderolites douvillei, Orbitoides douvil­ sic and Jurassic ones. Taxa of wider stratigraphic rang­ lei, O. tissoti, O. media and Pseudosiderolites vidali in es through the Cretaceous are predominant. From the the Late Campanian, and Murciella cuvillieri, M. renzi, beginning of the Early Cretaceous to the Late Barremi­ Rhapydionina liburnica, Fleuryana adriatica, Laffittei­ an the most important index forms were Protopenerop­ na mengaudi, O. media, Siderolites calcitrapoides and lis ultragranulata in the Early Berriasian, Montsalevia Omphalocyclus macroporus in the Maastrichtian. salevensis, Haplophragmoides joukowskyi, Meandros­ pira favrei, Vercorsella tenuis and V. camposaurii in the Biostratigraphic zonation was performed on depos­ Valanginian and Early Hauterivian, and Campanellula its ranging from the Carnian to the Maastrichtian. Low­ capuensis in the Late Hauterivian. From the Late Bar­ er and Middle Triassic rocks have not been investigated remian and Early Aptian to the Middle and Late Ceno­ sufficiently in order to propose a reliable subdivision. manian, the most important species were orbitolinids With respect to previous knowledge on the biostratig­ and alveolinids, accompanied by taxa from different raphy of shallow-marine, platform carbonate deposits foraminiferal groups. Index forms include Rectodic­ of the Mediterranean area, a new, more detailed bios­ tyoconus giganteus and Palorbitolina lenticularis in tratigraphic subdivision is proposed. The following bio­ the Late Barremian, P. lenticularis, Paleodictyoconus zones were defined: actinostoma, Praeorbitolina cormyi, P. wienandsi, Mes­ – within Upper Triassic deposits: orbitolina lotzei, Voloshinoides murgensis, Sabaudia briacensis, Choffatella decipiens, Derventina filipescui, Lamelliconus procerus taxon-range zone, Neotrocholina friburgensis, N. aptiensis, Trocholina Lamelliconus procerus–Triasina hantkeni interval lenticularis, Pfenderina globosa and Pseudolituonella zone, conica in the Early Aptian, Mesorbitolina parva, M. Triasina hantkeni taxon-range zone, and texana, Archalveolina reicheli, Sabaudia capitata and Triasina hantkeni–Mesoendothyra sp. interval zone; Orbitolinopsis aquitanica in the Late Aptian, M. parva, M. texana, M. pervia, M. subconcava, Paracoskinolina – within Lower Jurassic deposits: sunnilandensis, A. reicheli and S. capitata in the Early Triasina hantkeni–Mesoendothyra sp. interval zone, Albian, as well as ‘Valdanchella’ dercourti, Neoiraquia Mesoendothyra sp. lineage zone, insolita, N. convexa, Simplorbitolina broennimanni, Lituosepta recoarensis lineage zone, Paracoskinolina fleuryi, Protochrysalidina elongata and Ovalveolina maccagnoae in the Late Albian. In Orbitopsella primaeva lineage zone, addition to orbitolinids and alveolinids, Upper Creta­ Orbitopsella primaeva–Lituosepta recoarensis con­ ceous deposits also contain nezzazatids, chrysalinids, current-range subzone, miliolids and soritids. However, despite relatively rich Lituosepta recoarensis–Orbitopsella praecursor fossil assemblages within certain sequences, which interval subzone, comprise all or part of some epochs, e.g. a stratigraphic Orbitopsella praecursor taxon-range zone, range of Turonian–Early Santonian, there are no index forms of short stratigraphic range. The most important Orbitopsella praecursor–Orbitopsella primaeva con­ index forms are: Ovalveolina maccagnoae, O. crassa, current-range subzone, Sellialveolina viallii, Orbitolina gr. concava, Conicor­ Orbitopsella praecursor abundance subzone, 50 Geologia Croatica 60/1

Orbitopsella praecursor–Pseudocyclammina liassica Praeorbitolina wienandsi lineage zone, interval zone, Mesorbitolina lotzei lineage zone, Pseudocyclammina liassica taxon-range zone, and Mesorbitolina parva lineage zone, Pseudocyclammina liassica–Gutnicella cayeuxi Mesorbitolina texana lineage zone, interval zone; Mesorbitolina subconcava taxon-range zone, – within Middle Jurassic deposits: ‘Valdanchella dercourti’ partial-range zone, and Pseudocyclammina liassica–Gutnicella cayeuxi Neoiraquia convexa taxon-range zone; interval zone – within Upper Cretaceous deposits: Gutnicella cayeuxi partial-range zone, Conicorbitolina conica/Conicorbitolina cuvillieri Timidonella sarda taxon-range zone, partial-range zone, Timidonella sarda–Pseudodictyopsella jurassica Chrysalidina gradata superzone, interval zone, Chrysalidina gradata partial-range zone, Pseudodictyopsella jurassica and Marzoella ficcarel­ lii assemblage zone, Pastrikella balcanica–Conicorbitolina conica con­ current-range zone, Pseudodictyopsella jurassica–Paravalvulina compli­ cata interval zone, Vidalina radoicicae–Chrysalidina gradata concur­ rent-range zone, Paravalvulina complicata taxon-range zone, Chrysalidina gradata–Pseudocyclammina sphaeroi­ Paleopfenderina salernitana taxon-range zone, dea interval zone, Satorina apuliensis taxon range subzone, Pseudocyclammina sphaeroidea–Scandonea samni­ Paleopfenderina salernitana–Kurnubia jurassica tica interval zone, interval zone, and Scandonea samnitica–Dicyclina schlumbergeri inter­ Kurnubia jurassica lineage zone; val zone, – within Upper Jurassic deposits: Dicyclina schlumbergeri–Murgella lata interval Kurnubia palastiniensis superzone, zone, Kurnubia palastiniensis lineage zone, Murgella lata partial-range zone, Kurnubia wellingsi lineage zone, Keramosphaerina tergestina abundance subzone, Chablaisia chablaisensis–Alveosepta jaccardi inter­ Calveziconus lecalvezae taxon-range zone, val zone, Calveziconus lecalvezae–Murciella cuvillieri inter­ Alveosepta jaccardi partial-range zone, val zone, Parurgonina caelinensis–Conicokurnubia orbitolini­ Murciella cuvillieri and Rhapydionina liburnica formis concurrent range-zone, assemblage zone, and Conicokurnubia orbitoliniformis–Kurnubia palasti­ Fleuryana adriatica taxon-range subzone. niensis interval zone (highest occurrence zone), With a total of 64 biozones in the stratigraphic range Kurnubia palastiniensis–Protopeneroplis ultragranu­ from the Carnian to Maastrichtian this is up to now lata interval zone, and not only the most detailed subdivision of the shallow- Protopeneroplis ultragranulata taxon-range zone; marine carbonate deposits of the Karst Dinarides, but also in the wider Tethyan/Mediterranean area. – within Lower Cretaceous deposits: Protopeneroplis ultragranulata taxon-range zone, Palaeobiogeographic analysis of the Adriatic Car­ Protopeneroplis ultragranulata–Vercorsella campo­ bonate Platform deposits indicated similarities and dif­ saurii interval zone, ferences of biotopes and foraminiferal assemblage com­ positions between the AdCP and other Tethyan shallow- Vercorsella camposaurii taxon-range zone, marine areas. Close similarities with neighbouring Peri­ Vercorsella camposaurii–Campanellula capuensis adriatic platforms are especially important, as well as interval zone, changes in the platform position within the Neotethyan Campanellula capuensis taxon-range zone, realm, from individualization in the Triassic and Ear­ Campanellula capuensis–Palorbitolina lenticularis ly Jurassic to disintegration in the Late Cretaceous. It interval zone may be concluded, that from its individualization to the Late Albian the AdCP represented part of the Southern Palorbitolina lenticularis superzone, Neotethyan palaeobiogeographic province, while during Rectodictyoconus giganteus and Palorbitolina lenti­ the Cenomanian it was probably palaeobiogeographi­ cularis assemblage zone, cally isolated from both main Neotethyan bioprovinces, Praeorbitolina cormyi lineage zone, and together with the neighbouring Periadriatic carbon­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 51 ate platforms represented part of the Central Mediterra­ largest foraminifera (Orbitopsella, Rectodictoconus, nean province until its final disintegration in the Latest Murgella, Keramosphaerina, etc.); Cretaceous. – Ivo SUŠA, HGI–CGS technician, for preparation of During a span of almost 280 MY, i.e. from the figures; Moscovian to the end of the Eocene in the area of the – Miroslav KLADNIČKI, HGI–CGS technician, for present Karst Dinarides, approximately 8000 m of preparation of tables; almost pure carbonates was deposited, most of them – Blanka CVETKO TEŠOVIĆ (University of Zagreb, Mesozoic in age. Deposits of the Adriatic Carbonate Faculty of Natural Sciences), for providing thin- Platform, representing a succession from the Toarcian sections with Murciella cuvillieri; to the end of the Cretaceous, are up to 6500 m thick, and they had only been partially adequately investi­ – Katica DROBNE (Palaeontological institute I. gated. They were mostly relatively poorly subdivided, Rakovac of the Slovenian Academy of Sciences and and some sequences were almost completely undivid­ Arts, Ljubljana, Slovenia) for photomicrographs of ed. Therefore, this paper will contribute to the more Fleuryana adriatica and microspheric Rhapydionina detailed stratigraphic subdivision of the Late Triassic, liburnica. Jurassic and Cretaceous. The results presented here may I am especially grateful to the reviewers – Blanka represent a sound foundation for future investigation of CVETKO TEŠOVIĆ and Ivan GUŠIĆ (both Univer­ these deposits in the Karst Dinarides, especially those sity of Zagreb, Faculty of Natural Sciences) and dealing with chronostratigraphy. Vladimir VESELI (INA Oil Indutry Plc., Research and Development Sector) – for their reviews enabling the improvement of this paper, but also for long- Acknowledgement lasting cooperation and discussions on taxonomy and stratigraphy of the Mesozoic benthic foraminifera, as I would like to thank so many of my colleagues and well as to Felix SCHLAGINTWEIT (University of collaborators for various forms of help during the Leoben, Austria) for his very valuable suggestions and writing and finalization of this paper, including many correction of some very important data, resulting in the discussions and suggestions on stratigraphic ranges of significant improvement of this paper. Also, I would some foraminiferal species within different parts of the like to thank Julie ROBSON (Open University Milton Karst Dinarides. I would like to thank my colleagues Keynes, UK) for the language review. who collaborated with me during the last four decades, and especially to: This paper represents the results of investigations – Branko SOKAČ (Croatian Geological Survey – HGI– performed with the support of the Ministry of Science, CGS) for his long-lasting collaboration, support Education and Sports of the Republic of Croatia and countless discussions on the stratigraphy of through the project 181–1953068–0242: ‘Microfossil Mesozoic benthic foraminifera, their assemblages, Assemblages in Carbonate Deposits of the Karst and relationships with calcareous algae; Dinarides’. – Igor VLAHOVIĆ (formerly HGI–CGS, now with the University of Zagreb, Faculty of Mining, Geology 6. REFERENCES and Petroleum Engineering) for continuous support, discussions on the importance of foraminifera for ANIĆ, D. (1962): Prilog poznavanju jure i krede istočnog Mesozoic stratigraphy and palaeogeography of the dijela Biokova (Beiträge zur Kenntnis der Jura- und Krei­ Adriatic Carbonate Platform and Karst Dinarides, as deschichten des östlichen Teiles des Biokovo Gebirges).– well as for the translation of this paper; Geol. vjesnik, 15/1, 161–176. – Tonći GRGASOVIĆ (HGI–CGS) for discussions on ANTONIJEVIĆ, R., PAVIĆ, A., KAROVIĆ, J., DIMITRIJE­ Triassic foraminifera; VIĆ, M., RADOIČIĆ, R., PEJOVIĆ, D., PANTIĆ, S. & – Antun HUSINEC (formerly HGI–CGS, now with ROKSANDIĆ, M. (1973): Osnovna geološka karta SFRJ the St. Lawrence University, Department of Geology, 1:100.000. Tumač za listove Kotor i Budva K34–50 i K34–62 (Basic Geological Map of SFRY 1:100.000. Geo­ Canton, NY, USA) for discussions on Late Jurassic logy of the Kotor and Budva Sheets).– Zavod za geološka and Early Cretaceous foraminifera; i geofizička istraživanja Beograd (1969), Savezni geološki – Ladislav FUČEK (HGI–CGS) for discussion on zavod Beograd, 64 p. Cretaceous, especially Late Cretaceous foraminifera; ARNAUD, H., ARNAUD-VANNEAU, A., BLANC-ALÉ­ – Milka SARKOTIĆ-ŠLAT, head of the HGI– TRU, M.C., ADATTE, T., ARGOT, M., DELANOY, G., CGS Laboratory and technicians T. ŠKRTIĆ, D. THIEULOY, J.P., VERMEULEN, J., VIRGONE, A., VIR­ GALOVIĆ and Đ. NOVOSELEC for thin-sections; LOUVET, B. & WERMEILLE, S. (1998): Répartition stratigraphique des orbitolinidés de la plate-forme urgo­ – Goran MIKŠA (INA Oil Indutry Plc., Research and nienne subalpine et jurassienne (SE de la France).– Géol. Development Sector) for microphotographs of the Alpine, 74, 3–89. 52 Geologia Croatica 60/1

ARNAUD-VANNEAU, A. (1980): Micropaléontologie, paléo­ Barilović and Skradska gora).– Geol. vjesnik, 27, 59–77. écologie et sédimentologie d’une plate-forme carbonatée BUKOVAC, J., POLJAK, M., ŠUŠNJAR, M. & ČAKALO, de la marge passive de la Téthys: L’Urgonien du Vercors M. (1984): Osnovna geološka karta SFRJ 1:100.000. septentrional et de la Chartreuse (Alpes occidentales).– Tumač za list Črnomelj L33–91 (Basic geological map Géol. Alpine, Mem., 11, 874 p. of SFRY 1:100.000. Geology of the Črnomelj sheet).– ARNAUD-VANNEAU, A., CONRAD, M.A., DELOFFRE, Geološki zavod Zagreb, Geološki zavod Ljubljana (1983), R., LUPERTO-SINNI, E., MASSE, J.P., PEYBERNÉS, Savezni geološki zavod Beograd, 63 p. B., RADOIČIĆ, R., SOKAČ, B. & VELIĆ, I. (1991): Distribution of Lower Cretaceous foraminifera and dasy­ BUSER, S. (1986): Osnovna geološka karta SFRJ 1:100.000. cladal algae of the Adriatic plate.– In: VLAHOVIĆ, I. Tolmač listov Tolmin in Videm L33–64, L33–63 (Basic & VELIĆ, I. (Eds.): Some Aspects of the Shallow Water geological map of SFRY 1:100.000. Geology of the Sedimentation on the Adriatic Carbonate Platform (Permi­ Tolmin and Videm sheets).– Geološki zavod Ljubljana an to Eocene). The Second Int. Symp. on the Adriatic Car­ (1985), Savezni geološki zavod Beograd, 103 p. bonate Platform, Zadar, 1991, Inst. of Geology Zagreb, CASTRO, J.M., COMPANY, M., DE GEA, G.A. & AGUA­ Abstracts, 22–26. DO, R. (2001): Biostratigraphy of the Aptian–Middle BASSOULLET, J.P. (1997): Les grands Foraminifères.– In: Cenomanian platform to basin domain in the Prebetic CARIOU, E. & HANTZPERGUE, P. (Coord.): Biostra­ Zone of Alicante, SE Spain: calibration between shallow tigraphie du Jurassique ouest-européen et méditerranéen. water benthonic and pelagic scales.– Cretaceous Research, Bull. Centre Rech. Elf. Explor. Prod., Mem, 17, 293–304. 22, 145–156. BASSOULLET, J.P., CHABRIER, G. & FOURCADE, E. CHAROLLAIS, J., BRÖNNIMANN, P. & ZANINETTI, L. (l976): Données complémentaires sur la morphologie, la (1966): Troisième note sur les Foraminifères du Crétacé structure interne et la position stratigraphique de Timido­ inférieur de la région genevoise. Remarques stratigra­ nella sarda (Foraminifère, Lituolidae).– Revue de Micro­ phiques et description de Pseuditextulariella salevensis, paléontologie, 19/1, 3–18. n.sp.; Haplophragmoides joukowskyi, n.sp.; Citaella? favrei, n.sp.– Arch. Sc. Genève, 19/1, 23–48. BASSOULLET, J.P., FOURCADE, E. & PEYBERNÈS, B. (l985): Paléobiogéographie des grands Foraminifères ben­ CHERCHI, A. & SCHROEDER, R. (1977): La forma micro­ thiques des marges néotéthysiennes au Jurassique et au sferica di Lamarmorella sarda CHERCHI e SCHROE­ Crétacé inférieur.– Bull. Soc. géol. France, 85, 699–713. DER, 1975 (Foraminiferida).– Boll. Soc. Paleont. Ital., 16/2, 239–243. BELAK, M., KOCH, G., GRGASOVIĆ, T., VLAHOVIĆ, I., VELIĆ, I., SOKAČ, B. & BENČEK, Đ. (2005): Novi CHERCHI, A. & SCHROEDER, R. (1980): Broeckina (Pas­ prinosi stratigrafiji evaporitno–karbonatno–klastično–vul­ trikella) biplana n.sp., noveau grand Foraminifère du kanogenog kompleksa Komiškog zaljeva (otok Vis, Hrvat­ Cénomanien basal de la Montagna de Tauch (Corbières, ska) (New contribution to the stratigraphy of evaporitic– Pyrénées françaises).– C. r. Acad. Sc. Paris, (D), 290, carbonate–clastic–volcanogenic complex of the Komiža 319–322. bay, Island of Vis, Croatia).– In: VELIĆ, I., VLAHOVIĆ, CHERCHI, A. & SCHROEDER, R. (1985): Broeckina (Pas­ I. & BIONDIĆ, R. (Eds.): 3. Hrv. geol. Kongres – 3rd trikella) biplana.– In: SCHROEDER, R. & NEUMANN, Croatian geological congress, Opatija 2005, Knjiga M. (eds., 1985): Les grands Foraminifères du Crétacé sažetaka – Abstracts book, Zagreb, 13–14. moyen de la région méditerranéenne. Geobios, mém. BILOTTE, M. (1985): Le Crétacé supérieur des plates-formes spéc., 7, 97–98. Est-Pyrénéennes.– Strata (2), 5, 1–438. CHERCHI, A., DE CASTRO, P. & SCHROEDER, R. (1978): BUCKOVIĆ, D., CVETKO TEŠOVIĆ, B., JELASKA, V. & Sull’età dei livelli a Orbitolinidi della Campania e del­ GUŠIĆ, I. (2003): The Jurassic succession of Mt. Svila­ le Murge Baresi (Italia meridionale).– Boll. Soc. Natur. ja.– In: VLAHOVIĆ, I. & TIŠLJAR, J. (Eds.): Evolution Napoli, 87, 1–17. of Depositional Environments from the Palaeozoic to the Quaternary in the Karst Dinarides and Pannonian Basin. CHIOCCHINI, M. & MANCINELLI, A. (2000): Marzoel­ 22nd IAS Meeting of Sedimentology, Opatija 2003, Field- la ficcarellii n.gen., n.sp. (Foraminiferida) nel Dogger trip Guidebook, Zagreb, 73–81. del Lazio nord-orientale (Italia).– Bolletino della Società Paleontologia Italiana, 39/1, 21–28. BUCUR, I.I. (1993): Les reprépresentants du genre Proto­ peneroplis WEYNSCHENK dans le dépôts du Crétacé CHIOCCHINI, M., FARINACCI, A., MANCINELLI, A., inférieur da la zone de Reşiţa-Moldova Nouǎ (Carpathes MOLINARI, V. & POTETTI, M. (1994): Biostratigrafia Méridionales, Roumanie).– Revue de Micropaléontologie, a foraminiferi, dasicladali e calpionelle delle successioni 36/3, 213–223. carbonatiche mesozoiche dell’Appennino centrale (Ita­ lia).– Studi Geologici Camerti, vol. spec., 1994, “Biostra­ BUCUR, I.I., KOCH, R., KIRMACI, Z.M. & TASLI, K. tigrafia dell’Italia centrale”, 9–129. (2004): Foraminifères du Jurassique supérieur et du Créta­ cé inférieur (Calcaire de Berdiga) de Kircaova (région de CLAVEL, B., CHAROLLAIS, J., SCHROEDER, R. & BUS­ Kale-–Gümüshane, NE Turquie).– Revue de Paléobiolo­ NARDO, R. (1995): Réflexions sur la biostratigraphie du gie, 23/1, 209–225. Crétacé inférieur et sur sa complémentarité avec l’analyse séquentielle: exemple de l’Urgonien jurassien et subal­ BUKOVAC, J., VELIĆ, I. & SOKAČ, B. (1974): Stratigraf­ pin.– Bull. de la Soc. géol. de France, 166, 663–680. ski, tektonski i paleogeografski odnosi u području Duga­ rese, Barilovića i Skradske gore (Stratigraphy, tectonics CVETKO TEŠOVIĆ, B. (2000): Stratigrafija donje krede and paleogeography of the region between Dugaresa, odabranih lokaliteta zapadne Hrvatske (Lower Creta­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 53

ceous stratigraphy at selected localities in western Croa­ FUČEK, L., GUŠIĆ, I., JELASKA, V., KOROLIJA, B. & tia).– Unpubl. PhD Thesis (in Croatian with Summary in OŠTRIĆ, N. (1990): Stratigrafija gornjokrednih naslaga English), University of Zagreb, Faculty of Natural Sci­ jugoistočnog dijela Dugog otoka i njihova korelacija s ences, 263 p. istovremenim naslagama otoka Brača (Upper Cretaceous stratigraphy of the SE part of the Dugi otok Island and its CVETKO TEŠOVIĆ, B., GUŠIĆ, I., JELASKA, V. & BUC­ correlation with the corresponding deposits of the Brač KOVIĆ, D. (2001): Stratigraphy and microfacies of the Island, Adriatic Carbonate Platform).– Geol. vjesnik, 43, Upper Cretaceous Pučišća Formation, Island of Brač, Cro­ 23–33. atia.– Cretaceous Research, 22/5, 591–613. FUČEK, L., VELIĆ, I., VLAHOVIĆ, I., OŠTRIĆ, N., KORO­ DE CASTRO, P. (1966): Contributo alla conoscenza delle LIJA, B. & MATIČEC, D. (1995): Novi podaci o strati­ Alveoline albiano–cenomaniane della Campania.– Boll. grafiji donje krede otoka Cresa.– In: VLAHOVIĆ, I., Soc. Natur. Napoli, 75, 219–275. VELIĆ, I. & ŠPARICA, M. (eds.): 1. hrv. geol. kongres DE CASTRO, P. (1980): Le Alveoline aptiano–cenomaniane Zbornik radova, 1 (First Croat. Geol. Congress, Procee­ del Mediterraneo centrale e orientale: sintesi e problemi.– dings, 1), 167–172. Universita degli studi di Napoli, Istituto di Paleontologia, GENDROT, C. (1968): Stratigraphie et micropaléontologie du 1–86. Sénonien de la région des Martigues près Marseille (Bou­ DE CASTRO, P. (1988): Les Alvéolinidés du Crétacé d’Ita­ ches-du-Rhône).– Eclogae geol. Helv., 61/2, 657–694. lie.– Revue de Paléobiologie, Vol. spéc. 2, Benthos ‘86, GRADSTEIN, F.M., OGG, J.G., SMITH, A.G., AGTER­ 401–416. BERG, F.P., BLEEKER, W., COOPER, R.A., DAVY­ DE CASTRO, P., DROBNE, K. & GUŠIĆ, I. (1994): Fleury­ DOV, V., GIBBARD, P., HINNOV, L., HOUSE, M.R., ana adriatica n.gen., n.sp. (Foraminiferida) from the LOURENS, L., LUTERBACHER, H.-P., McARTHUR, J., Uppermost Maastrichtian of the Brač island (Croatia) and MELCHIN, M.J., ROBB, L.J., SHERGOLD, J., VILLE­ some other localities on the Adriatic carbonate platform.– NEUVE, M., WARDLAW, B.R., ALI, J., BRINKHUIS, Razprave IV razreda SAZU Ljubljana, 35/8, 129–149. H., HILGEN, F.J., HOOKER, J., HOWARTH, R.J., KNOLL, A.H., LASKAR, J., MONECHI, S., POWELL, DERCOURT, J., GAETANI, M., VRIELYNCK, B., BAR­ J., PLUMB, K.A., RAFFI, I., RÖHL, U., SANFILIPPO, RIER, E., BIJU-DUVAL, B., BRUNET, M.F., CADET, A., SCHMITZ, B., SHACKLETON, N.J., SHIELDS, J.P., CRASQUIN, S. & SANDULESCU, M. (Eds., 2000): G.A., STRAUSS, H., VAN DAM, J., VEIZERM J., VAN Atlas Peri-Tethys, Palaeogeographical maps.– CCGM/ KOLFSCHOTEN, Th. & WILSON, D. (2004): A Geolo­ CGMW, Paris: 24 maps and explanatory notes: I–XX; gic time scale.– Cambridge University Press, 611 p. 269 p. GRGASOVIĆ, T. (1997): Upper Triassic biostratigraphy and DRAGIČEVIĆ, I. & VELIĆ, I. (2002): The northeastern mar­ algae from Žumberak (Croatia).– Geol. Croatica, 50/2, gin of the Adriatic Carbonate Platform.– Geol. Croatica, 201–214. 55/2, 185–232. GRGASOVIĆ, T. (2003): Revizija rodova Physoporella DROBNE, K., OGORELEC, B., PLENIČAR, M., BARAT­ STEINMANN i Oligoporella PIA (Dasycladales) (Revisi­ TOLO, F., TURNŠEK, D. & ZUCCHI-STOLFA, M.L. on of genera Physoporella STEINMANN and Oligoporel­ (1989): The Dolenja Vas section, a transition from Cre­ la PIA (Dasycladales)).– Unpubl. PhD Thesis (in Croatian taceous to Paleocene in the NW Dinarides, Yugoslavia.– with English Summary), University of Zagreb, Faculty of Mem. Soc. Geol. It., 40 (1987), 73–84. Natural Sciences, VIII+484 p. ĐOKIĆ, V., ŽIVALJEVIĆ, M. & PEROVIĆ, Z. (1976): Osno­ GRGASOVIĆ, T., HALAMIĆ, J., GORIČAN, Š., SLOVE­ vna geološka karta SFRJ 1:100.000 list Gusinje K34–52 NEC, D. & KOLAR-JURKOVŠEK T. (2000): Trijaske (Basic geological map of SFRY 1:100.000. Geology of the dubokomorske naslage odabranih lokaliteta u sjevero­ Gusinje sheet).– Zavod za geološka istraživanja Titograd zapadnoj Hrvatskoj (Triassic deep-water sediments of (1968), Savezni geološki zavod Beograd, 49 p. selected localities in northwestern Croatia).– 2. Hrv. FARINACCI, A. & RADOIČIĆ, R. (1964): Correlazione fra geol. kongres Cavtat–Dubrovnik, Zbornik radova Zagreb serie giuresi e cretacee dell’Appennino centrale e delle (Second Croat. Geol. Congress, Proceedings), 181–188. Dinaridi esterne.– La ricerca scientifica, 34/2, 269–300. GUŠIĆ, I. (1969a): Some new and inadequately known Juras­ FLEURY, J.J. (1980): Les zones de Gavrovo–Tripolitza et du sic foraminifers from central Croatia.– Geol. vjesnik, 22, Pinde–Olnos (Grèce continentale et Péloponèse du Nord). 55–87. Evolution d’une plate-forme et d’un bassin dans leur cadre GUŠIĆ, I. (1969b): Biostratigraphic and micropaleontologic alpin.– Publ. Soc. Géol. Du Nord, 4, 1–651. characteristics of some Jurassic cross-sections in central FLEURY, J.J., BIGNOT, G., BLONDEAU, A. & POIGNANT, Croatia.– Geol. vjesnik, 22, 89–97. A. (1985): Biogéographie de Foraminifères benthiques GUŠIĆ, I. (1977): A new foraminiferal family Biokovinidae téthysiens du Sénonien à l’Éocène supérieur.– Bull. Soc. from the Jurassic of the Dinarids and its phylogenetic rela­ géol. France, (8), 1/5, 757–770. tionships.– Palaeont. Jugosl., 18, 1–31. FRIJIA, G. & PARENTE, M. (2007) : Isotope-stratigraphy GUŠIĆ, I. (1981): Variation range, evolution, and biostrati­ in Turonian–Campanian shallow-water carbonates of graphy of Palorbitolina lenticularis (BLUMENBACH) southern Apennines (Italy).– Geophysical Research Abs­ (Foraminiferida, Lituolacea) in the Lower Cretaceous of tract, 9, SRef-ID: 1607-7962/gra/EGU2007-A-06819, the Dinaric Mountains in Yugoslavia.– Paläont. Z., 55, European Geoscience Union 2007. 3/4, 3–4, 191–208. 54 Geologia Croatica 60/1

GUŠIĆ, I. & BABIĆ, Lj. (1970): Neke biostratigrafske i lito­ Yugoslavia.– AAPG Bull., 75, 1007–1017. genetske osobine jure Žumberka (Some biostratigraphic JURKOVŠEK, B., TOMAN, M., OGORELEC, B., ŠRIBAR, and lithogenetic characteristics of the Jurassic in Žumbe­ L., DROBNE, K., POLJAK, M. & ŠRIBAR, Lj. (1996): rak, Northwest Croatia).– Geol. vjesnik, 23, 39–54. Formacijska geološka karta južnega dela Tržaško– GUŠIĆ, I. & JELASKA, V. (1990): Stratigrafija gornjokred­ komenske planote: kredne in paleogenske karbonatne nih naslaga otoka Brača u okviru geodinamske evolucije kamnine 1:50000 (Geological map of the southern part Jadranske karbonatne platforme (Upper Cretaceous Strati­ of the Trieste–Komen plateau: Cretaceous and Paleogene graphy of the Island of Brač within the Geodynamic Evo­ carbonate rocks).– Inštitut za geologijo, geotehniko in lution of the Adriatic Carbonate Platform).– Djela JAZU, geofiziko Ljubljana, 143 p. Zagreb, 69, 1–160. KABAL, Y. & TASLI, K. (2003): Biostratigraphy of the Low­ GUŠIĆ, I. & JELASKA, V. (1993): Upper Cenomanian–Low­ er Jurassic carbonates from the Aydincik area (Central er Turonian sea-level rise and its consequences on the Taurides, S Turkey) and morphological analysis of Lituoli­ Adriatic–Dinaric carbonate platform.– Geol. Rundsch., pora termieri (HOTTINGER, 1967).– Journal of Forami­ 82/4, 676–686. niferal Research, 33/4, 338–351. GUŠIĆ, I. & VELIĆ, I. (1978): Lituolipora polymorpha KALEZIĆ, M., MIRKOVIĆ, M. & ŠKULETIĆ, D. (1973): n.gen., n.sp. (Foraminiferida, Lituolacea?) from the Osnovna geološka karta SFRJ 1:100.000. Tumač za Middle Liassic of the Outer Dinarids in Croatia and the list Šavnik K34–39 (Basic geological map of SFRY establishment of a new family Lituoliporidae (Lituolipo­ 1:100.000. Geology of the Šavnik sheet).– Zavod za ra polymorpha n.gen., n.sp. (Foraminiferida, Lituolacea?) geološka istraživanja Titograd (1966), Savezni geološki iz srednjeg lijasa Vanjskih Dinarida Hrvatske i uspostav­ zavod Beograd, 65 p. ljanje nove porodice Lituoliporidae).– Geol. vjesnik, 30/1, KOCHANSKY-DEVIDÉ & PANTIĆ, (1966): Meandrospira 73–93. u donjem i srednjem trijasu i neki popratni fosili u Dina­ GUŠIĆ, I., NIKLER, L. & SOKAČ, B. (1971): The Jurassic ridima (Meandrospira in der unteren und mittleren Trias in the Dinaric mountains of Croatia and the problems of sowie einige begleitende Fossilien in den Dinariden).– its subdivision.– Ann. Inst. Geol. Publ. Hungar., 54/2, Geol. vjesnik, 19, 15–28. 165–173. KORBAR, T., FUČEK, L., HUSINEC, A., VLAHOVIĆ, I., GUŠIĆ, I., JELASKA, V. & VELIĆ, I. (1988): Foraminiferal OŠTRIĆ, N., MATIČEC, D. & JELASKA, V. (2001): assemblages, facies and environments in the Upper Cre­ Cenomanian carbonate facies and rudists along shallow taceous of the island of Brač.– Revue de Paleobiologie, intraplatform basin margin – the island of Cres (Adriatic Vol. spec. 2, Benthos ‘86, 447–456. sea, Croatia).– Facies, 45, 39–58. HAMAOUI, M. (1979): Contribution a l’étude du Cénoma­ KOROLIJA, B., ŽIVALJEVIĆ, T. & ŠIMUNIĆ, An. (1981): no–Turonien d’Israel comparaisons micropaléontologi­ Osnovna geološka karta SFRJ 1:100.000. Tumač za list ques avec quelques régions mésogéennes.– Unpubl. PhD Slunj L33–104 (Basic geological map of SFRY 1:100.000. Thesis, L’Université Pierre et Marie Curie Paris, 121 p. Geology of the Slunj sheet).– Institut za geološka istraživanja Zagreb (1972), Geološki zavod Sarajevo HOTTINGER, L. (1967): Foraminifères imperforés du Méso­ (1968), Savezni geološki zavod Beograd, 47 p. zoïque marocain.– Notes Mém. Serv. géol. Maroc, 209, 1–168. LUPERTO SINNI, E. (1968): Microfauna bathoniana del M. Alpi.– Boll. Soc. Natur. Napoli, 77, 181–206. HOTTINGER, L., DROBNE, K. & CAUS, E. (1989): Late Cretaceous, larger, complex miliolids (Foraminifera) ende­ LUPERTO SINNI, E. (1976): Microfossili senoniani delle mic in Pyrenean faunal province.– Facies, 21, 99–134. Murge.– Riv. Ital. Paleont., 82/2, 293–416. HUSINEC, A., VELIĆ, I., FUČEK, L., VLAHOVIĆ, I., LUPERTO SINNI, E. & RICCHETTI, G. (1978): Studio MATIČEC, D., OŠTRIĆ, N. & KORBAR, T. (2000): micropaleontologico–stratigrafico di una successione car­ Mid Cretaceous orbitolinid (Foraminiferida) record from bonatica del Cretaceo superiore rilevata nel sottosuolo del­ the islands of Cres and Lošinj (Croatia).– Cretaceous le Murge sud-orientali.– Riv. Ital. Paleont., 84/3, 561–666. Research, 21, 155–171. MAMUŽIĆ, P. & GRIMANI, M. (1970): Mikrofosilne zajed­ JAFFREZO, M. (1980): Les formations carbonatées des Cor­ nice gornje krede otočnog i obalnog dijela Hrvatske (pre­ bières (France) du Dogger à l’Aptien: micropaléontologie, liminarna podjela) (Microfossil assemblages of the Upper stratigraphique, biozonation, paléoécologie. Extension des Cretaceous of the Croatian island and coastal region; in résultats à la Mésogée.– Unpubl. PhD Thesis, Univ. Pierre Croatian).– VII kongres geologa SFRJ Zagreb, 1, 177– et Marie Curie Paris VI, XVII+614+23 p. 182. MAMUŽIĆ, P., POLŠAK, A., GRIMANI, M. & MAGAŠ, JELASKA, V., GUŠIĆ, I., JURKOVŠEK, B., OGORELEC, N. (1976): Biostratigrafske i litofacijelne karakteristi­ B., ĆOSOVIĆ, V., ŠRIBAR, Lj. & TOMAN, M. (1994): ke gornje krede sjeveroistočnih padina Biokova u južnoj The Upper Cretaceous geodynamic evolution of the Adri­ Hrvatskoj (Biostratigraphic and lithofacial characteristics atic–Dinaric carbonate platform(s).– Geol. Mediterr., of the Upper Cretaceous of northeastern slopes of Bioko­ 21/3–4, 89–91. vo in southern Croatia).– VIII jugosl. geol. kongres Bled JENKYNS, H.C. (1988): The Early Toarcian (Jurassic) anoxic (1974), Ljubljana, 2, 167–180. event: stratigraphic, sedimentary, and geochemical evi­ MAMUŽIĆ, P., POLŠAK, A., GRIMANI, M., ŠIMUNIĆ, dence.– Am. J. Sci., 288, 101–151. Al. & KOROLIJA, B. (1980): Detaljni geološki stup kroz JENKYNS, H.C. (1991): Impact of Cretaceous sea-level rise naslage cenomana sjeverno od Vela Luke na otoku Korču­ and anoxic events in the Mesozoic carbonate platform of li (Detailed geological column through Cenomanian depo­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 55

sits north of Vela Luka, Island of Korčula, Dalmatia, sout­ sheet).– Zavod za geološka istraživanja Titograd (1968), hern Croatia).– Geol. vjesnik, 31, 91–103. Savezni geološki zavod Beograd, 67 p. MAMUŽIĆ, P., POLŠAK, A., GRIMANI, M., ŠIMUNIĆ, Al. MOJIČEVIĆ, M., VILOVSKI, S., TOMIĆ, B. & PAMIĆ, & KOROLIJA, B. (1981a): Detaljni geološki stup kroz J. (1977): Osnovna geološka karta SFRJ 1:100.000 list naslage cenomana u zapadnom dijelu otoka Hvara (Collo­ Banjaluka L 33–119 (Basic geological map of SFRY ne stratigraphique détaillée du cenomanien et du turonien 1:100.000, geology of the Banjaluka sheet).– Institut za dans la partie occidentale de ľile de Hvar).– Geol. vjesnik, geološka istraživanja Sarajevo (1976), Savezni geološki 33, 49–57. zavod Beograd, 50 p. MAMUŽIĆ, P., KOROLIJA, B., GRIMANI, M. & ŠIMU­ MOJIČEVIĆ, M., PAMIĆ, J., MAKSIMČEV, S. & PAPEŠ, J. NIĆ, Al. (1981b): Detaljni geološki stupovi kroz naslage (1979): Osnovna geološka karta SFRJ 1:100.000. Tumač krede (gornji otriv–donji turon) središnjeg dijela poluo­ za list Bosanska Krupa L 33–117 (Basic geological map of toka Pelješca (Collones géologiques à travers les couches SFRY 1:100.000. Geology of the Bosanska Krupa sheet).– Cretacées (Hauterivien supérieur–Turonien inférieur) de Geoinženjering Sarajevo (1971), Savezni geološki zavod la partie centrale de la presque île de Pelješac).– Geol. Beograd, 50 p. vjesnik, 34, 19–25. NEAGU, T. (1994): Early Cretaceous Trocholina group and MAMUŽIĆ, P., POLŠAK, A., GRIMANI, M. & KOROLIJA, some related genera from Romania (Part I).– Revista B. (1982): Detaljni geološki stup kroz naslage cenomana Española de Micropaleontologia, 26/3, 117–143. i donjega turona u središnjem dijelu otoka Cresa (Collo­ NIKLER, L. & SOKAČ, B. (1968): Biostratigraphy of the ne stratigraphique détaillée à travers du cenomanien et du Jurassic of Velebit (Croatia).– Geol. vjesnik, 21, 161–176. turonien inférieur dans la partie centrale de ľile de Cres).– Geol. vjesnik, 35, 65–70. PANTIĆ, S. (1965): Pilammina densa, n.gen., n.sp. and other Ammodiscidae from the Middle Triassic in Crmnica MAMUŽIĆ, P., KOROLIJA, B., GRIMANI, M. & ŠIMU­ (Montenegro).– Geol. vjesnik, 18/1, 189–192. NIĆ, Al. (1983): Geološki stup kroz naslage krede (raspo­ na gornji cenoman–donji senon) središnjega dijela poluo­ PANTIĆ, S. (1966/67): Turrispirillina minima n.sp. iz trijas­ toka Pelješca (Collone stratigraphique détaillée à travers kih sedimenata Dinarida (Turrispirillina minima n.sp. des les couches cretacées (cénomanien supérieur–sénonien sédiments triasiques des Dinarides externes).– Vesnik (A) inférieur) dans la partie centrale de la presqu’ile de Pel­ Beograd, 24/25, 255–258. ješac).– Geol. vjesnik, 36, 73–77. PANTIĆ-PRODANOVIĆ, S. (1975): Trijaske mikrofacije MAMUŽIĆ, P., GRIMANI, M. & ŠIMUNIĆ, Al. (1984): Geo­ Dinarida (Les microfacies triasiques des Dinarides).– Druš­ loški stup kroz naslage gornjeg turona i donjeg senona tvo za nauku i umjetnost Crne Gore (Soc. Sci. Arts Mon­ kod Primoštena u Dalmaciji (Collone géologique à travers ténégro) Titograd, 4, Odj. prir. nauka (Classe Sci. Natur.), les couches du Turonien supérieur–Sénonien inférieur 4, 1–257. près de Primošten en Dalmatie centrale).– Geol. vjesnik, PAPEŠ, J. (1975): Osnovna geološka karta SFRJ 1:100.000. 37, 55–58. Tumač za list Livno K33–11 (Basic geological map of MARINČIĆ, S. (1997): Tectonic structure of the island of SFRY 1:100.000. Geology of the Livno sheet).– Institut za Hvar (southern Croatia) (Tektonska struktura otoka Hva­ geološka istraživanja Sarajevo (1967), Savezni geološki ra).– Geol. Croatica, 50/1, 57–77. zavod Beograd, 64 p. MARINKOVIĆ, R. & ĐORĐEVIĆ, D. (1981): Osnovna geo­ PEYBERNÈS, B. (1976): Le Jurassique et le Crétacé inféri­ loška karta SFRJ 1:100.000. Tumač za list Jajce L33–131 eur des Pyrénées franco–espagnoles entre la Garonne et (Basic geological map of SFRY 1:100.000. Geology of la Méditerranée.– Unpubl. PhD Thesis, Univ. Tolouse, the Jajce sheet).– Geoinženjering Sarajevo (1975), Savez­ 459 p. ni geološki zavod Beograd, 52 p. PFENDER, J. (1938): Les Foraminifères du Valanginien pro­ vençal.– Bull. Soc. Géol. France, 5/8, 231–242. MARITAN, L., MAZZOLI, C. & MELIS, E. (2003): A mul­ tidisciplinary approach to the characterization of Roman POLŠAK, A., BAUER, V. & SLIŠKOVIĆ, T. (1982): Stratigra­ gravestones from Aquileia (Udine, Italy).– Archaeometry, phie du Crétacé supérieur de la plate-forme carbonatée 45/3, 363–374. dans les Dinarides externes.– Cretaceous Research, 3, 125–133. MARKOVIĆ, B. (1975): Osnovna geološka karta SFRJ 1:100.000. Tumač za list Dubrovnik K34–49 (Basic geo­ RADOIČIĆ, R. (1960): Mikrofacije krede i starijeg tercijara logical map of SFRY 1:100.000. Geology of the Gacko Spoljnih Dinarida Jugoslavije (Microfaciès du Crétacé et sheet).– Zavod za geološka i geofizička istraživanja Beo­ du Paleogène des Dinarides externes de Yougoslavie).– grad (1966), Savezni geološki zavod Beograd, 43 p. Zavod geol. istraž. NR Crne Gore (A) Titograd. Paleonto­ logija jugosl. Dinarida, 4/1, 5–172. MIRKOVIĆ, M. (1980): Osnovna geološka karta SFRJ 1:100.000. Tumač za list Gacko K34–26 (Basic geological RADOIČIĆ, R. (1966): Microfaciès du Jurassique des Dinari­ map of SFRY 1:100.000. Geology of the Gacko sheet).– des externes de la Yougoslavie.– Geologia, 9, 5–378. Zavod za geološka istraživanja Titograd (1974), Savezni RADOIČIĆ, R. (2005): New Dasycladales and microbiota geološki zavod Beograd, 67 p. from the lowermost Valanginian of the Mirdita zone.– Annales géologiques de la pénninsule Balkanique, 66, MIRKOVIĆ, M., KALEZIĆ, M., PAJOVIĆ, M., ŽIVALJE­ 27–53. VIĆ, M. & ŠKULETIĆ, D. (1978): Osnovna geološka karta SFRJ 1:100.000. Tumač za list Bar K34–63 (Basic RAIĆ, V. & PAPEŠ, J. (1982): Osnovna geološka karta SFRJ geological map of SFRY 1:100.000. Geology of the Bar 1:100.000. Tumač za list Ston K33–48 (Basic geological 56 Geologia Croatica 60/1

map of SFRY 1:100.000. Geology of the Ston sheet).– SEPTFONTAINE, M. (1984): Biozonation (à l’aide des Fora­ Geoinženjering Sarajevo, Geološki zavod Zagreb (1980), minifères imperforés) de la plate-forme interne carbona­ Savezni geološki zavod Beograd, 39 p. tée liasique du Haut Atlas (Maroc).– Rev. Micropaléont., 27/3, 209–229. SALVADOR, A. (Ed., 1994): International stratigraphic guide. A guide to stratigraphic classification, terminology, and SEPTFONTAINE, M. (1988): Vers une clasification évoluti­ procedure.– International Subcommission on Stratigraphic ve des Lituolidés (Foraminifères) jurassiques en milieu de Classification (ISSC), Second Edition, XIX+214 p. plate-forme carbonatèe (Towards an evolutionary classifi­ cation of Jurassic lituolides (Foraminifera), in carbonate SAINT-MARC, P. (1974): Etude stratigraphique et micropa­ platform environment).– Rev. de Paléobiol., Vol. Spec., 2, léontologique de l’Albien, du Cénomanien et du Turonien 229–256. du Liban.– Notes Mémoires Moyen–Orient, 13, 1–342. SEPTFONTAINE, M. & DE MATOS, J.E. (1998): Pseudodic­ SAINT-MARC, P. (1977): Répartition stratigraphique des tyopsella jurassica n.gen., n.sp., a new foraminifera from grands Foraminifères benthiques de l’Aptien, de l’Albien, the early Middle Jurassic of the Musandam Peninsula, N. du Cénomanien et du Turonien dans les régions méditer­ Oman Mountains; sedimentological and stratigraphical ranéennes.– Rev. Esp. De Micropal., 9/3, 317–325. context.– Rev. Micropaléont., 41/1, 71–87. SARTONI, S. & CRESCENTI, U. (1962): Ricerche biostrati­ SEPTFONTAINE, M., ARNAUD-VANNEAU, A., BASSO­ grafische nel mezozoico dell’Appennino meridionale.– ULLET, J.P., GUŠIĆ, I., RAMALHO, M. & VELIĆ, I. Giorn. Geol. (2a), 29, 161–304. (1991): Les Foraminifères imperforés des plate-formes SAVIĆ, D. & DOZET, S. (1985): Osnovna geološka karta carbonatées jurassiques: état des connaissances et perspec­ SFRJ 1:100.000. Tumač za list Delnice L33–90 (Basic tives d’avenir.– Bull. Soc. Vaud. Sc. Nat., 80/3, 255–277. geological map of SFRY 1:100.000. Geology of the Del­ SOFILJ, J., ŽIVANOVIĆ, M. & PAMIĆ, J. (1980): Osno­ nice sheet).– Geološki zavod Zagreb, Geološki zavod vna geološka karta SFRJ 1:100.000. Tumač za list Prozor Ljubljana (1983), Savezni geološki zavod Beograd, 66 p. K33–12 (Basic geological map of SFRY 1:100.000. Geo­ SCHROEDER, R. (1962): Orbitolinen des Cenomans Süd­ logy of the Prozor sheet).– Institut za geološka istraživanja westeuropas.– Paläont. Z., 36, 171–202. Sarajevo (1971), Savezni geološki zavod Beograd, 49 p. SCHROEDER, R. (1964a): Zur evolution der Cenoman-Orbi­ SOKAČ, B. (2001): Lower and Middle Liassic calcareous tolinen. Eine Entgegnung an J. HOFKER jun.– N. Jb. algae (Dasycladales) from Mt. Velebit (Croatia) and Mt. Geol. Paläont., Mh., 1964, 682–693. Trnovski Gozd (Slovenia) with particular reference to the genus Palaeodasycladus (PIA, 1920) 1927 and its spe­ SCHROEDER, R. (1964b): Communication préable sur l’ori­ cies.– Geol. Croatica, 54/2, 133–257. gine des Orbitolines.– C. R. Soc. geol. France, 1964/10, 411–413. SOKAČ, B. & VELIĆ, I. (1978): Biostratigrafska istraživanja donje krede Vanjskih Dinarida (I). Neokom zapadne Istre SCHROEDER, R. (1964c): Orbitoliniden biostratigraphie des (Biostratigraphic investigations of the Lower Cretaceous Urgons nordöstlich von Teruel (Spanien).– Neues Jahrb. of the Outer Dinarids. The Neocomian of Western Istria).– für Geol. U. Paläont., Mh., 462–473. Geol. vjesnik, 30/1, 243–250. SCHROEDER, R. (1975): General evolutionary trends in SOKAČ, B. & VELIĆ, I. (1979a): Triassic, Jurassic and Low­ Orbitolinas.– Revista Española de Micropaleontologia, er Cretaceous of the Karst Dinarids in the western Croa­ num. esp., 117–128. tia.– 16th European micropal. coll. Ljubljana, 79–100. SCHROEDER, R. (1979): Les Orbitolines de l’Aptien: défini­ SOKAČ, B. & VELIĆ, I. (1979b): Lower Cretaceous in tions, origine et évolution (The Aptian Orbitolinas – defi­ western Istria. Excursion J (Limska Draga).– 16th Europe­ nitions, origin and evolution).– Géobios Mém. spéc., 3, an micropal. coll. Ljubljana., 209–216. 289–299. SOKAČ, B., TIŠLJAR, J. & VELIĆ, I. (1978): Model biostra­ SCHROEDER, R., CHERCHI, A., GUELLAL, S. & VILA, tigrafskog raščlanjivanja i analiza sredina taloženja u kar­ J.M. (1974): Biozonation par les grands Foraminifères du bonatnim sedimentima donje krede Biokova (A model of Jurassique supérieur et du Crétacé inférieur et moyen des biostratigraphic subdivision and an interpretation of depo­ séries néritiques en Algérie NE. Considérations palèobio­ sitional environments in the Lower Cretaceous carbona­ gèographiques.– VI Coll. Africain Micropal., Tunis 1974, te sediments of Biokovo mountain, South Croatia).– IX prétirage, 1–8. Kongres geologa Jugoslavije Sarajevo, Zbornik radova, SCHROEDER, R., CLAVEL, B., CHERCHI, A., BUSNAR­ 226–232. DO, R., CHAROLLAIS, J. & DECROUEZ, D. (2002): STEUBER, T., KORBAR, T., JELASKA, V. & GUŠIĆ, I. Lignées phylétiques d’Orbitolinidés de l’intervalle Haute­ (2005): Strontium-isotope stratigraphy of Upper Cretace­ rivien supérieur–Aptien inférieur; leur importance strati­ ous platform carbonates of the island of Brač (Adriatic graphique.– Revue Paléobiologie, 21/2, 853–863. sea, Croatia): implications for global correlation of plat­ SCHROEDER, R. & NEUMANN, M. (Eds., 1985): Les form evolution and biostratigraphy.– Cretaceous Research, grands Foraminifères du Crétacé moyen de la région médi­ 26, 741–756. terranéenne.– Geobios, mém. spéc., 7, 1–160. TASLI, K. (1993): Micropaléontologie, stratigraphie et envi­ SEPTFONTAINE, M. (1980): Les Foraminifères imperforés ronnement de dépôt des séries Jurassiques à faciès de des milieux de plate-forme au Mésozoïque: détermination plate-forme dela région de Kale–Gümüşhane (Pontides pratique, interprétation phylogénétique et utilisation bios­ Orientales, Turquie) (Micropaleontology, stratigraphy and tratigraphique.– Rev. Micropaléont., 23/3–4, 169–203. depositional environment of the Jurassic carbonate plat­ Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 57

form sequence of Kale–Gümüşhane region, Eastern Pon­ Korčule (Biostratigraphic investigations of the Lower tids, Turkey).– Rev. Micropaléont., 36/1, 45–65. Cretaceous of the Outer Dinarids (II). The Upper Albian of the island of Korčula).– Geol. vjesnik (1979), 31, 185– TIŠLJAR, J. & VELIĆ, I. (1993): Upper Jurassic (Malm) 190. shallow water carbonates in the western Gorski Kotar area: facies and depositional environments (western Cro­ VELIĆ, I. & SOKAČ, B. (1983): Stratigraphy of the Lower atia).– Geol. Croatica, 46/2, 263–279. Cretaceous index-fossils in the Karst Dinarides (Yugosla­ via).– Zitteliana, 10, 485–491. TIŠLJAR, J., VLAHOVIĆ, I., SREMAC, J., VELIĆ, I., VESELI, V. & STANKOVIĆ, D. (1991): Excursion “A” Velić, I. & Tišljar, J. (1988): Litostratigrafske jedinice u – Velebit Mt., Permian–Jurassic.– In: VLAHOVIĆ, I. & dogeru i malmu zapadne Istre, zapadna Hrvatska, Jugosla­ VELIĆ, I. (eds.): “Some Aspects of the Shallow Water vija (Lithostratigraphic units in the Dogger and Malm of Sedimentation on the Adriatic Carbonate Platform (Per­ western Istria).– Geol. vjesnik, 41, 25–49. mian to Eocene)”. The Second Int. Symp. on the Adria­ VELIĆ, I. & VLAHOVIĆ, I. (1994): Foraminiferal assem­ tic Carbonate Platform, Zadar, 1991., Inst. of Geology, blages in the Cenomanian of the Buzet–Savudrija area Zagreb, Excursion Guide-Book, 1–49. (northwestern Istria, Croatia).– Geol. Croatica, 47/1, 25– TIŠLJAR, J., VLAHOVIĆ, I., VELIĆ, I. & SOKAČ, B. 43. (2002): Carbonate platform megafacies of the Jurassic and VELIĆ, I., TIŠLJAR, J. & SOKAČ, B. (1979): Stratigraphy Cretaceous deposits of the Karst Dinarides.– Geol. Croati­ and depositional environments of the Lower Cretaceous ca, 55/2, 139–170. in the Karst region of the Outer Dinarides, Yugoslavia VELIĆ, I. (1973): Stratigrafija krednih naslaga u graničnom (Stratigraphie et paleoenvironnments du Cretace inferieur području Velike i Male Kapele (Stratigraphy of the Cre­ dans la region karstique des Dinarides externes, Yougosla­ taceous deposits in the border region of Velika Kapela vie).– Geobios, Mem. spec., 3, 245–252. and Mala Kapela mountains, Croatia).– Geol. vjesnik, 26, VELIĆ, I., SOKAČ, B. & ŠĆAVNIČAR, B. (1982): Osno­ 93–109. vna geološka karta SFRJ 1:100.000. Tumač za list Ogu­ VELIĆ, I. (1977): Jurassic and Lower Cretaceous assembla­ lin L33–103 (Basic Geologic Map of SFRY 1: 100.000. Geology of the Ogulin sheet).– Geološki Zavod Zagreb ge-zones in Mt. Velika Kapela, central Croatia.– Acta (1980), Savezni geol. zavod Beograd, 46 p. Geologica, 9/2, 15–37. VELIĆ, I., TIŠLJAR, J. & SOKAČ, B. (1989): The variability VELIĆ, I. (1988): Lower Cretaceous benthic foraminiferal of thicknesses of the Barremian, Aptian and Albian carbo­ biostratigraphy of the shallow-water carbonates of the nates as a consequence of changing depositional environ­ Dinarides.– Revue de Paleobiologie, Vol. spec. 2, Benthos ments and emersion in Western Istria (Croatia, Yugosla­ ‘86, 467–475. via).– Mem. Soc. Geol. It., 40 (1987), 209–218. VELIĆ, I. (2005): “Misterij dogera”: foraminiferske zajednice VELIĆ, I., MATIČEC, D., VLAHOVIĆ, I. & TIŠLJAR, J. i stratigrafska raščlamba srednjojurskih naslaga u krškim (1995): Stratigrafski slijed jurskih i donjokrednih karbo­ Dinaridima (Jadranska karbonatna platforma) (Benthic nata (bat–gornji alb) u zapadnoj Istri, ekskurzija A (Stra­ foraminifera assemblages and stratigraphic subdivision of tigraphic succession of Jurassic and Lower Cretaceous the Middle Jurassic in Karst Dinarides, Adriatic Carbonate carbonates (Bathonian–Upper Albian) in western Istria, Platform).– In: VELIĆ, I., VLAHOVIĆ, I. & BIONDIĆ, Excursion A).– In: VLAHOVIĆ, I. & VELIĆ, I. (Eds.): R. (eds.): 3. Hrvatski geološki kongres (3rd Croatian 1. Hrv. geol. kongres, Vodič ekskurzija (First Croat. Geol. Geological Congress), Opatija, 2005, Knjiga sažetaka Congress, Exc. Guide-Book), 31–66. (Abstracts book), Hrvatski geološki institut (Croatian Geological Survey), 159–160, Zagreb. VELIĆ, I., VLAHOVIĆ, I. & MATIČEC, D. (2002): Deposi­ tional sequences and palaeogeography of the Adriatic car­ Velić, I. & Sokač, B. (1974): On the tripartite subdivision bonate platform.– Mem. Soc. Geol. It., 57, 141–151. of the Malm in Mt. Velika Kapela (Croatia).– Geol. vjes­ nik, 27, 143–150. VENTURINI, S. (2005): L’evento a Keramosphaerina ter­ gestina: cosiderazioni bio-cronostratigrafiche (The Kera­ VELIĆ, I. & SOKAČ, B. (1978a): Biostratigrafska analiza mosphaerina tergestina event: bio-chronostratigraphic jure i donje krede šire okolice Ogulina (središnja Hrvat­ aspects).– Natura Nascosta, 31, 15–22. ska) (Biostratigraphic analysis of the Jurassic and Lower Cretaceous in wider region of Ogulin, central Croatia).– VESELI, V. (1999): Facijesi karbonatnih sedimenata mlađeg Geol. vjesnik, 30/1, 309–337. mezozoika i paleogena u pučinskim bušotinama sjevernog Jadrana (Younger Mesozoic and Palaeogene carbonate VELIĆ, I. & SOKAČ, B. (1978b): Zone s orbitolinidama u deposits facies in the off-shore wells of the Northern Adri­ krednim naslagama krškog dijela hrvatskih Dinarida atic).– Unpublished PhD Thesis, University of Zagreb, (Orbitolinid biozones in the Cretaceous deposits of the 306 p. Croatian Karst Dinarides).– IX Kongres geologa Jugosla­ vije Sarajevo, Zbornik radova, 215–222. VLAHOVIĆ, I., TIŠLJAR, J. & VELIĆ, I. (1994): Influence of synsedimentary tectonics and eustatic changes on depo­ VELIĆ, I. & SOKAČ, B. (1979): Younger Lower Cretaceous sition of the Cenomanian carbonates in Istria (Western and Lower Cenomanian of central Croatia. Excursion C Croatia).– Geol. Mediterr., 21/3–4, 189–193. (Karlovac–Ogulin).– 16th European micropal. coll., Ljub­ VLAHOVIĆ, I., TIŠLJAR, J., VELIĆ, I. & MATIČEC, D. ljana, 147–156. (2002): The Karst Dinarides are composed of relics of a VELIĆ, I. & SOKAČ, B. (1980): Biostratigrafska istraživanja single Mesozoic platform: facts and consequences.– Geol. donje krede Vanjskih Dinarida (II). Gornji alb otoka Croatica, 55/2, 171–183. 58 Geologia Croatica 60/1

VLAHOVIĆ, I., TIŠLJAR, J., VELIĆ, I., MATIČEC, D., geološka i geofizička istraživanja Beograd (1967), Savez­ SKELTON, P.W., KORBAR, T. & FUČEK, L. (2003): ni geološki zavod Beograd, 53 p. Main events recorded in the sedimentary succession of VUJNOVIĆ, L. (1981): Osnovna geološka karta SFRJ the Adriatic Carbonate Platform from the Oxfordian to the 1:100.000. Tumač za list Bugojno L33–140 (Basic geolo­ Upper Santonian in Istria (Croatia).– In: VLAHOVIĆ, I. gical map of SFRY 1:100.000. Geology of the Bugojno & TIŠLJAR, J. (eds.): Evolution of Depositional Environ­ sheet).– Geoinženjering Sarajevo (1975), Savezni geološki ments from the Palaeozoic to the Quaternary in the Karst zavod Beograd, 58 p. Dinarides and the Pannonian basin. 22nd IAS Meeting of Sedimentology, Opatija – September 17–19 2003, Field- ŽIVALJEVIĆ, M., PAJOVIĆ, M., ĐOKIĆ, V. & ŠKULETIĆ, Trip Guidebook 19–56, Zagreb. D. (1971): Osnovna geološka karta SFRJ 1:100.000. Tumač za list Titograd K34–51 (Basic geological map of VLAHOVIĆ, I., TIŠLJAR, J., VELIĆ, I. & MATIČEC, D. SFRY 1:100.000. Geology of the Titograd sheet).– Zavod (2005): Evolution of the Adriatic Carbonate Platform: za geološka istraživanja Titograd (1967), Savezni geološki palaeogeography, main events and depositional dyna­ zavod Beograd, 57 p. mics.– Palaeogeography, Palaeoclimatology, Palaeoecolo­ gy, 220, 333–360. ŽIVALJEVIĆ, M., VUJISIĆ, P. & STIJOVIĆ, V. (1989): Osnovna geološka karta SFRJ 1:100.000. Tumač za VRHOVČIĆ, J. & MOJIČEVIĆ, M. (1983): Osnovna geološ­ list Žabljak K34–27 (Basic geological map of SFRY ka karta SFRJ 1:100.000. Tumač za list Ključ L33–130 1:100.000. Geology of the Žabljak sheet).– Zavod za geo­ (Basic geological map of SFRY 1:100.000. Geology of loška istraživanja Titograd (1987), Savezni geološki zavod the Ključ sheet).– Geoinženjering Sarajevo (1976), Savez­ Beograd, 62 p. ni geološki zavod Beograd, 40 p. VUJISIĆ, T. (1975): Osnovna geološka karta SFRJ 1:100.000. Tumač za list Nikšić K34–38 (Basic geological map of SFRY 1:100.000. Geology of the Nikšić sheet).– Zavod za

Manuscript received December 17, 2006. Revised manuscript accepted April 10, 2007. Velić: Stratigraphy and Palaeobiogeography of Mesozoic Benthic Foraminifera... 59

APPENDIX LIST OF TAXA

1. Foraminifera Glomospira watersi LOEBLICH, 1946 Glomospirella sp. Accordiella conica FARINACCI, 1962 Glomospirella triphonensis BAUD, ZANINETTI & BRONNIMANN, Agerina martana (FARINACCI, 1959) 1971 Alveosepta jaccardi (SCHRODT, 1894) Gutnicella cayeuxi (LUCAS, 1935) Alzonella cuvillieri BERNIER & NEUMANN, 1970 Haplophragmium sp. Amijiella amiji (HENSON), 1948 Haplophragmoides joukowskyi CHAROLLAIS, BRÖNNIMANN & Ammobaculites radstadtensis KRISTAN-TOLLMANN, 1964 ZANINETTI, 1966 Ammodiscus parapriscus (HO, 1959) Haplophragmoides globosus LOZO, 1944 Anchispirocyclina lusitanica (EGGER, 1902) Haurania deserta HENSON, 1948 Anchispirocyclina neumannae BERNIER, FLEURY & RAMALHO, Hellenocyclina beotica REICHEL, 1949 1979 Hoyenella sinensis (HO, 1959) Archaealveolina sp. Idalina antiqua MUNIER-CHALMAS & SCHLUMBERGER, 1885 Archaealveolina reicheli (DE CASTRO, 1966) ‘Involutina’ sp. Archiacina munieri (MARIE, 1958) Involutina liasica (JONES, 1853) Arenovidalina chialingchiangensis HO, 1959 Karaisella uzbekistanica KURBATOV, 1971 Auloconus permodiscoides (OBERHAUSER, 1964) Keramosphaerina tergestina STACHE, 1913 Aulotortus sp. Keramosphaerina sarda CHERCHI & SCHROEDER, 1990 Aulotortus friedli (KRISTAN-TOLLMANN, 1962) Kilianina blancheti PFENDER, 1933 Aulotortus pokornyi (SALAJ, 1967) Kilianina? rahonensis (FOURY & VINCENT, 1967) Aulotortus praegaschei (KOEHN-ZANINETTI, 1968) Kurnubia jurassica (HENSON, 1948) Aulotortus sinuosus (WEYNSCHENK, 1956) Kurnubia palastiniensis HENSON, 1948 Aulotortus tenuis (KRISTAN, 1957) Kurnubia wellingsi (HENSON, 1948) Aulotortus tumidus (KRISTAN-TOLLMANN, 1964) Labyrinthina mirabilis WEYNSCHENK, 1951 Belorussiella textilarioides (REUSS, 1863) Laffitteina mengaudi (ASTRE, 1923) Biconcava bentori HAMAOUI & SAINT-MARC, 1970 Lamelliconus multispirus (OBERHAUSER,1957) Biokovina gradacensis GUŠIĆ, 1977 Lamelliconus procerus (LIEBUS, 1942) Biplanata peneropliformis HAMAOUI & SAINT-MARC, 1970 Lepidorbitoides minor (SCHLUMBERGER, 1901) Bosniella oenensis GUŠIĆ,1977 Lituolipora termieri (HOTTINGER, 1967) Broekinella neumannae GENDROT, 1968 Lituosepta recoarensis CATI, 1959 Calveziconus lecalvezae CAUS & CORNELLA, 1981 Marzoella ficcarellii CHIOCCHINI & MANCINELLI, 2000 Campanellula capuensis DE CASTRO, 1964 Mayncina bulgarica LAUG, PEYBERNÈS & REY, 1980 Chablaisia chablaisensis (SEPTFONTAINE, 1977) Meandrospira dinarica KOCHANSKY-DEVIDÉ & PANTIĆ, 1966 Charentia cuvillieri NEUMANN, 1965 Meandrospira favrei (CHAROLLAIS, BRÖNNIMANN & ZANI­ Choffatella decipiens SCHLUMBERGER, 1904 NETTI, 1966) Chrysalidina gradata D’ORBIGNY, 1839 Meandrospira pusila (HO, 1959) Cisalveolina fraasi (GÜMBEL, 1872) Meandrovoluta asiagoensis FUGAGNOLI, GIANNETTI & RET­ Cisalveolina lehneri REICHEL, 1941 TORI, 2003 Conicokurnubia orbitoliniformis SEPTFONTAINE, 1988 Merlingina cretacea HAMAOUI & SAINT-MARC, 1970 Conicorbitolina conica (D’ARCHIAC, 1837) Mesoendothyra croatica GUŠIĆ, 1969 Conicorbitolina corbarica SCHROEDER, 1985 Mesoendothyra sp. Conicorbitolina cuvillieri (MOULLADE, 1972) Mesorbitolina lotzei SCHROEDER, 1964 Cuneolina compressa SCHLAGINTWEIT, 1988 Mesorbitolina oculata DOUGLASS, 1960 Cuneolina conica D’ORBIGNY, 1846 Mesorbitolina parva DOUGLASS, 1960 Cuneolina ketini INAN, 1988 Mesorbitolina pervia DOUGLASS, 1960 Cuneolina parva HENSON, 1948 Mesorbitolina subconcava LEYMERIE, 1878 Cuneolina pavonia D’ORBIGNY, 1846 Mesorbitolina texana (ROEMER), 1849 Cuneolina sliteri ARNAUD-VANNEAU & PREMOLI SILVA, 1995 Minouxia lobata GENDROT, 1963 Cyclogyra mahayeri (BRÖNNIMANN, ZANINETTI & BOZOR­ Mohlerina basiliensis (MOHLER, 1938) GNIA, 1972) Moncharmontia apenninica (DE CASTRO, 1966) Debarina hahounerensis FOURCADE, RAOULT & VILA, 1972 Moncharmontia compressa (DE CASTRO, 1966) Derventina filipescui NEAGU, 1968 Montsalevia salevensis (CHAROLLAIS, BRÖNNIMANN & ZANI­ Dictyopsella kiliani MUNIER-CHALMAS, 1900 NETTI, 1966) Dictyopselloides cuvillieri (GENDROT, 1968) Murciella cuvillieri FOURCADE, 1966 Dicyclina schlumbergeri MUNIER-CHALMAS, 1887 Murciella renzi FLEURY, 1979 Duostomina alta KRISTAN-TOLLMANN, 1960 Murgeina apula (LUPERTO SINNI, 1968) Duotaxis metula KRISTAN, 1957 Murgeina cenomana (LUPERTO SINNI, 1998) Endotriadella wirtzi (KOEHN-ZANINETTI, 1968) Murgella lata LUPERTO SINNI, 1966 Everticyclammina praevirguliana FUGAGNOLI, 2000 Nautiloculina sp. Everticyclammina virguliana (KOECHLIN, 1942) Nautiloculina broennimanni ARNAUD-VANNEAU & PEYBER­ Fleuryana adriatica DE CASTRO, DROBNE & GUŠIĆ, 1994 NÈS, 1978 Frondicularia woodwardi HOWCHIN,1895 Nautiloculina circularis (SAID & BARAKAT, 1959) Glomospira sp. Nautiloculina cretacea PEYBERNÈS, 1976 Glomospira urgoniana ARNAUD-VANNEAU, 1980 Nautiloculina oolithica MOHLER, 1938 60 Geologia Croatica 60/1

Neoendothyra reicheli REITLINGER,1965 Pseudodictyopsella jurassica SEPTFONTAINE & DE MATOS, 1998 Neoiraquia convexa DANILOVA, 1963 Pseudoeggerella elongata SEPTFONTAINE, 1988 Neoiraquia insolita (DECROUEZ & MOULLADE, 1974) Pseudolituonella conica LUPERTO SINNI & MASSE, 1993 Neotrocholina aptiensis (IOVCHEVA, 1969) Pseudolituonella marie GENDROT,1968 Neotrocholina friburgensis GUILLAUME & REICHEL, 1957 Pseudolituonella reicheli MARIE, 1955 Nezzazata conica (SMOUT, 1956) Pseudonummoloculina sp. Nezzazata germanica OMARA & STRAUCH, 1965 Pseudonummoloculina aurigerica CALVEZ, 1988 Nezzazata gyra (SMOUT, 1956) Pseudonummoloculina heimi (BONET, 1956) Nezzazata isabellae ARNAUD-VANNEAU & SLITER, 1995 Pseudorhapydionina dubia (DE CASTRO, 1965) Nezzazata simplex OMARA, 1956 Pseudorhapydionina laurinensis (DE CASTRO, 1965) Nezzazatinella picardi (HENSON, 1948) Pseudorhapydionina mediterranea (DE CASTRO, 1974) Novalesia sp. Pseudorhipidionina casertana (DE CASTRO, 1965) Novalesia angulosa (MAGNIEZ, 1972) Pseudorhipidionina murgeana (CRESCENTI, 1964) Novalesia cornucopia ARNAUD-VANNEAU, 1980 Pseudosiderolites vidali (DOUVILLE, 1907) Novalesia distorta ARNAUD-VANNEAU, 1980 Raadshovenia salentina (PAPETTI & TEDESCHI, 1965) Novalesia producta (MAGNIEZ, 1972) Rectodictyoconus giganteus SCHROEDER, 1964 Nummofallotia cretacea (SCHLUMBERGER, 1900) Redmondoides lugeoni (SEPTFONTAINE, 1977) Nummoloculina? regularis PHILIPPSON, 1887 Reticulinella fleuryi CVETKO, GUŠIĆ & SCHROEDER, 1997 Omphalocyclus macroporus (LAMARCK, 1816) Reticulinella reicheli CUVILLIER, BONNEFOUS, HAMAOUI & Orbitammina elliptica (D’ARCHIAC, 1843) TIXIER, 1969 Orbitoides douvillei (SILVESTRI, 1910) Rhapydionina liburnica (STACHE, 1889) Orbitoides hottingeri VAN HINTE, 1966 Rotalia mesogeensis TRONCHETTI, 1993 Orbitoides media (D’ARCHIAC, 1837) Rotorbinella scarsellai TORRE, 1966 Orbitoides tissoti (SCHLUMBERGER, 1902 ) Rumanoloculina robusta (NEAGU, 1968) Orbitolina concava (LAMARCK, 1816) Sabaudia auruncensis (CHIOCCHINI & DI NAPOLI ALLIATA, Orbitolina sefini HENSON, 1948 1966) Orbitolinopsis sp. Sabaudia briacensis ARNAUD-VANNEAU, 1980 Orbitolinopsis aquitanica SCHROEDER & POIGNANT, 1964 Sabaudia capitata ARNAUD-VANNEAU, 1980 Orbitopsella ?dubari HOTTINGER, 1967 Sabaudia dinapolii CHIOCCHINI, 1984 Orbitopsella praecursor (GÜMBEL, 1872) Sabaudia minuta (HOFKER, 1965) Orbitopsella primaeva (HENSON, 1948) Satorina apuliensis FUORCADE & CHOROWICZ, 1980 Ovalveolina crassa DE CASTRO, 1966 Satorina mesojurassica (MAYNC, 1972) Ovalveolina maccagnoae DE CASTRO, 1966 Scandonea phoenissa SAINT-MARC, 1974 Paleodictyoconus actinostoma ARNAUD-VANNEAU & SCHROE­ Scandonea pumila SAINT-MARC, 1974 DER, 1976 Scandonea samnitica DE CASTRO, 1971 Paleopfenderina salernitana (SARTONI & CRESCENTI, 1962) Sellialveolina viallii COLALONGO, 1963 Paleopfenderina trochoidea (SMOUT & SUGDEN, 1962) Siderolites calcitrapoides LAMARCK, 1801 Palorbitolina lenticularis (BLUMENBACH, 1805) Simplorbitolina brönnimanni (DECROUEZ & MOULLADE, 1974) Paracoskinolina fleuryi DECROUEZ & MOULLADE, 1974 Siphovalvulina colomi BOU DAGHER-FADEL, ROSE, BOSENCE Paracoskinolina sunniladensis (MAYNC, 1955) & LORD, 2001 Paravalvulina complicata SEPTFONTAINE, 1988 Siphovalvulina gibraltarensis BOU DAGHER-FADEL, ROSE, Parurgonina caelinensis CUVILLIER, FOURY & PIGNATTI- BOSENCE & LORD, 2001 MORANO, 1968 Siphovalvulina variabilis SEPTFONTAINE, 1988 Pastrikella balcanica CHERCHI, RADOIČIĆ & SCHROEDER, Spiraloconulus giganteus CHERCHI & SCHROEDER, 1982 1976 Spiraloconulus perconigi (ALLEMANN & SCHROEDER,1972) Pastrikella biplana CHERCHI & SCHROEDER, 1980 Spirolina cretacea TRONCHETTI & GROSHENY, 1993 Peneroplis parvus DE CASTRO, 1965 Spiroloculina cretacea REUSS, 1854 Peneroplis turonicus SAID & KENAWY, 1957 Spiroloculina robusta BRADY, 1884 Pfenderella arabica REDMOND, 1964 Stensiöina surrentina TORRE, 1966 Pfenderina globosa FOURY, 1968 Timidonella sarda BASSOULLET, CHABRIER & FOURCADE, Pilammina densa PANTIĆ, 1965 1974 Pilamminella semiplana (KOCHANSKY-DEVIDÉ & PANTIĆ, Triasina hantkeni MAJZON, 1954 1966) Trochamminoides coronus LOEBLICH & TAPPAN, 1946 Planisepta compressa (HOTTINGER, 1967) Trocholina sp. Praealveolina iberica REICHEL, 1936 Trocholina acuta OBERHAUSER, 1964 Praealveolina simplex REICHEL, 1936 Trocholina alpina (LEUPOLD, 1935) Praechrysalidina sp. Trocholina arabica HENSON, 1949 Praechrysalidina infracretacea LUPERTO SINNI, 1979 Trocholina crassa KRISTAN, 1957 Praekurnubia crusei REDMOND, 1964 Trocholina delphinensis ARNAUD-VANNEAU, BOISSEAU & Praeorbitolina cormyi SCHROEDER, 1964 DARSAC, 1988 Praeorbitolina wienandsi SCHROEDER, 1964 Trocholina elongata (LEUPOLD, 1935) Praesiderolites douvillei WANNIER, 1983 Trocholina molesta GORBATCHIK, 1959 Protochrysalidina elongata LUPERTO-SINNI, 1999 Trocholina gigantea PELISSIE & PEYBERNÈS, 1983 Protopeneroplis striata WEYNSCHENK, 1950 Trocholina odukpaniensis DESSAUVAGIE, 1968 Protopeneroplis ultragranulata (GORBATCHIK, 1971) Trocholina sagittaria ARNAUD-VANNEAU, BOISSEAU & DAR­ Pseudocyclammina liassica HOTTINGER, 1967 SAC, 1988 Pseudocyclammina lituus (YOKOYAMA, 1890) Trochospira avnimelechi HAMAOUI & SAINT-MARC, 1970 Pseudocyclammina massiliensis MAYNC, 1959 Turrispirillina minima PANTIĆ, 1967 Pseudocyclammina maynci HOTTINGER, 1967 Turriglomina mesotriassica (KOEHN-ZANINETTI, 1968) Pseudocyclammina rugosa (D’ORBIGNY, 1850 ‘Valdanchella’ dercourti DECROUEZ & MOULLADE, 1974 Pseudocyclammina sphaeroidea GENDROT, 1968 Vercorsella arenata ARNAUD-VANNEAU, 1980