New and poorly known Middle Jurassic larger benthic foraminifera from the Karst Dinarides of Croatia

Some new and poorly known larger benthic foraminifera are described from Middle Jurassic (Upper Aalenian–Bajocian) shallow-water limestones of the Croatian Karst Dinarides. Cymbriaella lorigae FUGAGNOLI is reported for the fi rst time outside its type-locality, the Upper Pliensbachian of the Southern Alps. New taxa described include Bosniella bassoulleti n. sp. and Dubrovnikella septfontainei n. gen., n. sp. (family Biokovinidae). Both Cymbriaella lorigae FUGAGNOLI and Everticyclammina praevirguliana FUGAGNOLI are reported for the fi rst time from Middle Jurassic strata. The new fi ndings enlarge the stratigraphic ranges and allow further insights into the phylogenetic evolution of the larger benthic foraminifera during the Early to Middle Jurassic period.


GEOLOGICAL SETTING
During the early Middle Jurassic, two sedimentary provinces with signifi cantly different environments may be distinguish ed within the area of the Adriatic Carbonate Platform (VLA-HOVIĆ et al., 2005, for details).One encompasses platform marginal and peri-marginal dynamics, areas strongly infl uenced by currents and waves, with the occasional de-velopment and destruction of patch reefs.The second area belongs to the inner platform characterized by protected environments with a steady and uniform sedimentation of thick layers of carbonate mud.In marginal areas, the most common deposits are thick-bedded ooid-intraclast-bioclastic-ske letalpelletal grainstones and packstones, in places rud stones, rich in fossil remains.In contrast, thick-layered mudstones, ra re ly wackestones and packstones with very few fos sil remains were deposited in the inner platform area.
In the area of southern Croatia, southeast of Split, as well as in Montenegro and Northern Albania, Middle Jurassic se diments were deposited in the fi rst, marginal province.This area includes the localities in Biokovo Mt. and in the vicinity of Dubrovnik (Fig. 1) where the investigated foraminiferal fauna has been found, including the new taxa described in the present paper.Aalenian-Bajocian carbonates of Southern Croatia crop out in the frontal part of the sequence of Mesozoic deposits uplifted and thrusted over the Palaeogene clastic sediments.Carbonates are in direct contact with clastic deposits, stratigraphically ranging from the Late Triassic to the Middle Jurassic.The reversed-overthrust contact extends from the northwestern part of Biokovo Mt. to the east of Konavle, along the border with Montenegro.It is morphologically expressed by the steep slopes of Biokovo Mt., the hills around Dubrovnik and Montene-

The Osojnik section
The Middle Jurassic deposits of the Osojnik section were investigated along the road connecting the village of Osojnik with Dubrovnik (Figs. 1-2).About 2 km south-southeast of the village, the road crosses the tectonic boundary between the folded Lower Jurassic (Pliensbachian) Lithiotis limestones, (about 30 m in thickness), and massive Aalenian ooid grainstones.The latter are overlain by bioclastic-oncoidal and intraclast grainstones, in places rudstones with recrystallized skeletons and debris of calcareous algae, molluscs

The Slano section
The Middle Jurassic part of the Slano section is measured northeast of Slano on the slopes of the Kosmatovica and Kolomnić hills (Fig. 4).As in the Osojnik profi le, the Middle Jurassic succession starts with Aalenian ooid limestone, who se thickness is unknown due to the tectonic contact with the Lower Jurassic carbonates.The ooid limestones are followed by approximately 20 m of layered mudstones and wacke stones.In the next 270 m of the profi le, massive latediagenetic dolomites, and oncoid-intraclastic-bioclastic grainstones prevail, with rudstones in places in alternation with skeletal-intraclastic grainstones/rudstones.These deposits are mostly thick layered (0.6 m to 1.5 m) and rarely even massive.There are interbeds or thin intervals of mud-supported intraclastic-skeletal limestones, prevailing after the aforementioned massive late-diagenetic dolomites at the end of the Bajocian and beginning of the Bathonian limestones.Fossil remains and debris belong mostly to molluscs (predominantly gastropods), corals, echinoderms, stromatoporoids, calcareous algae, and foraminifera.In the initial part of the sequence and within a further ~60 m thick interval, a rich assemblage of index foraminifera has been observed.In addition to Bosniella croatica, which starts within the ooid limestones, Gutnicella cayeuxi, Cymbriaella lorigae, Dubrovnikella septfontainei n. gen., n. sp., Pseudocyclammina may n ci HOTTINGER and Bosniella bassoulleti n. sp.appear almost in the same horizon (Fig. 5).Most of the cited species have a rather narrow stratigraphic range, so that above this biohorizon, for another 30 m, only G. cayeuxi extends until the appearance of Spiraloconulus perconigi (ALLEMAN & SCHROEDER).After a further 20 m, Timidonella sarda was determined, and around 70 m above that, the ran ge of the dasycladalean alga Selliporella donzellii begins.In the next 30 m of the column, together with S. donzellii, the only foraminifera occurring is B. croatica (also indicating its last appearance) and B. bassoulleti n. sp., the last appearance of which is determined after a further 30 m.In the next 70 m, there are no signifi cant microfossils except S. don zellii.After 15-20 m of massive late-diagenetic dolomite, skeletal-intraclastic grainstones and skeletal wackestones con taining Pfenderella arabica occur; 20 m above Pseudoeggerella elongata and fi nally an additional 20 m or so, a Bathonian as semblage with Paleopfenderina salernitana, etc. occurs (Fig. 5).The thickness of the Upper Aalenian and Bajocian carbonates in the Slano cection between the Aalenian ooid limestones and the Bathonian limestones is about 350 m.

The Biokovo Mount section
The Biokovo Mt. profi le was measured east of Tučepi along the Staza-Lemišini Doci road in the Biokovo Nature Park (Fig. 6).As in the case of the Osojnik and Slano sections, the oldest Middle Jurassic carbonates are thick layered and massive ooid limestones of Aalenian age.The lithological features of the Upper Aalenian-Bajocian carbonates of the Biokovo section, in continuous succession over ooid limestones, are very similar to those in the Osojnik and Slano sections.In the older parts, bioclastic-skeletal-oncoid and intraclastic grainstones to rudstones prevail, in places with interbedded oolites and skeletal wackestones and mudstones.Among the skeletal remains there is fossil debris and sections of molluscs, corals, stromatoporoids, algae, and for-aminifers.Skeletal and bioclastic grainstones/mudstones are most common in the younger part of the column, and they often alternate with intraclastic grainstones.Local emergen ce ho rizons rarely occur.Among the foraminifera, Bosniella croatica and Gutnicella cayeuxi were already present in the ooid limestones, over lain by about 50 m of thick-layered skeletal-oncoidal grainstones with interbedded skeletal wackestones, followed by another, but thinner package, i.e. about 10 m thick, of ooid limestones.Within the interval between the base and these thinner ooid limestone beds there are bioclastic oncoid and skeletal grainstones/packstones with Everticyclammina praevir guliana FUGAGNOLI (Fig. 7).Above the thin ooid limestones there is a sequence of thick-bedded, oncoidal, skeletal grainstones with rare G. cayeuxi and more frequent Ti mi donella sarda.The latter is found in the younger strata, beneath the local emergence horizon, characterized by the fi rst occurrences of Bosniella bassoulleti n. sp. and Spiraloconulus giganteus.Among the cited species, B. croatica, B. bassoulleti n. sp. and S. giganteus continue after emergence, while 15 m above the emergence the alga Selliporella donzellii appears.In the middle part of the Bajocian column, about 80 m above the mentioned emergence horizon, Pseudodictyopsella jurassica and Marzoella fi ccarellii occur, these being up to now their fi rst, and only occurrences in the Karst Dinarides.Approximately 40 m above the last occurrence of S. giganteus, after a further 60 m of the column, B. croatica disappears, and after another 30 m also B. bassoulleti n. sp.disappears.In the following 40 m of the column to the fi rst occurrence of Pfenderella arabica, there are no important foraminiferal species.About 10 m above, in the topmost Bajocian beds, Paravalvulina complicata and Pseudoeggerella elongata appear, and after a further ca.10 m there is an emergence breccia with black pebbles (originating from marsh deposits on the tidal fl at).About 15 m above the emergence horizon the next emersion level follows, and then layers with Paleopfenderina salernitana and other Bathonian foraminifera.The thickness of the Upper Aalenian and Bajocian carbonates in the Biokovo profi le between the Aalenian ooid limestones and the Bathonian limestones cannot be accurately determined due to tectonic disintegration and gentle folding, but is estimated at about 350 m.

SYSTEMATIC PALAEONTOLOGY
The systematics follows the "year 2000 classifi cation" of KAMINSKI ( 2004).For the diagnosis of genera, we essentially follow LOEBLICH & TAPPAN (1987).The description main ly follows the terminological compendium of HOT-TIN GER (2006).Description: Large test consisting of a more or less plani spirally coiled initial portion (1.5 to 2 whorls) and a rather large uncoiled portion with 4 to 5 chambers, separated by thick septa.In the uncoiled portion, chamber width increases moderately.Test wall thick, alveolar with incorporated foreign material (e.g., peloids, small foraminifera).Single foramen in the initial part; in longitudinal sections rather broad multiple (up to 4-5) foramina are discernible in the uncoiled portion (Fig. 8d).Dimensions see Table 1.

Class Foraminiferida
Remarks and comparisons: Cymbriaella lorigae was described from the Pliensbachian of the Southern Alps (FU-GAGNOLI, 1999) and the stratigraphic range can now be extended to include the Upper Aalenian-Lower Bajocian interval.Besides the Italian and Croatian records, the species was recently also reported from the Liassic of Turkey (ÖZ-KAYMAK et al., 2010).The Middle Jurassic specimens from Croatia agree closely with the Lower Jurassic material described by FUGAGNOLI (1999).
Microfacies: Intraclastic pack-to rudstones with debris of molluscs, and gastropods.The associated foraminifera in-

Suborder Biokovinina KAMINSKI, 2004 Superfamily Biokovinacea GUŠIĆ, 1977 Family Biokovinidae GUŠIĆ, 1977
Remarks: Bosniella was removed from the Biokovinidae by MIKHALEVICH ( 2004) due to the lack of endoskeletal structures as reported from Bosniella (see Tab. 2).This view was corrected in the "year 2000 classifi cation" of KAMINSKI ( 2004) with the creation of the suborder Biokovinina KAMIN-SKI and the maintenance of Bosniella within the Bio kovinidae by giving the wall structure a higher hierarchy than the presence/absence of the endoskeleton.The absence/presence of endoskeletal structures can instead be taken into consideration to differentiate the two subfamilies within the Biokovinidae.Holotype: Specimen in equatorial section fi gured in (Fig. 10d).Thin section Os 10.

Depository
Diagnosis: A representative of the genus Bosniella showing a planispiral discoidal test, varying from partly involute and widely umbilicate to involute, often with slightly undulating growth direction and a prominent uncoiled portion.Chambers numerous in the coiled part, becoming moderately broader during growth, in the uncoiled portion either constant in width or expanding.Foramen single, interiomarginal in the coiled part becoming multiple in the uncoiled part.Wall microgranular with keriothecal texture.
Description: Test free, lenticular, more or less planispirally coiled initially, with 2-2½ whorls and a maximum of up to 14 chambers in the last whorl in adult specimens; the adult part may be uncoiling.The coiling is often not exactly planispiral but may show a slight inclination towards the plane of the previous whorl or an oscillating inclination that may result in an s-or sigmoid shape in axial sections (Figs.11b,  i, k).In most cases, the umbilicus is well pronounced (e.g.Fig. 10g, upper specimen, Fig. 11c, m).The fi rst chamber (protoconch) is subsphaerical (diameter up to 0.11 mm), enclosed by a thin wall (membrane) with a thickness of about 0.005 mm (Fig. 12a).In the coiled portion, chamber width gradually increases, often distinctly so in the last half of the fi nal whorl, resulting in a more or less asymmetry in axial sections (Fig. 10g, upper specimen, Figs.11c, e).In equatorial sections, the chambers are becoming higher during ontogeny but also show a fl attened top with angular peripheries; in the older parts, chamber peripheries are more rounded (Figs.10a-b, d, 11a).In the same way, the diameter of the single foramen near the chamber base also increases.Sutures between subsequent chambers are poorly developed (e.g.Figs.10a, 11a).The ultimate chambers often show upward growth towards the previous whorl resulting in a massive thickening of the chamber wall (Figs. 10a,11a).This seems to be the preparation of the test to uncoil, in broadening the junction between the coiled and uncoiled parts (Figs.10e-f).
The planispiral stage is often followed by an uncoiled, more or less rectilinear, stage.The shape may vary from cy-   lindrical with chamber width remaining more or less constant (e.g.Fig. 10f), to expanding (cylindroconical), with chamber width increasing successively (Fig. 10o).Occasionally, there are specimens displaying chambers not rectilinearly arrang ed, but with varying growth directions (with respect to the previous chamber).Successive change of chamber growth direction may result in an overall sigmoid appearance (Fig. 10n).Chamber shape of the uncoiled part in axial sections is variable, mostly hemispherical with fl at or concave base, in cases also with fl attened top (trapezoid, e.g.third ultimate chamber in Fig. 10e).The distal end of the uncoiled part is mostly broadly rounded (Fig. 10f, o) or more rarely acutely rounded (Fig. 10e).The septa of the uncoiled portion are pierced by several ("multiple") foramina (Fig. 10d), often not detectable.In oblique sections, the chamber lumina are in direct contact to each other giving the impression of a broad single foramen in the centre (Figs.10o, q).The test wall is fi nely agglutinating; the fi ne keriothekal structure is indistinct and not always detectable (Fig. 11d), presumably due to diagenetic processes.
Remarks and comparisons: The differences of Bosniella bassoulleti n. sp. to the other three species of the genus Bosniella can be summarized as follows: B. oenensis GUŠIĆ (Pliensbachian of Bosnia): As axial sections were not available in the material on which the original description was based, information on the type of coil- Both Bosniella croatica and Bosniella fontainei display a test dimorphism (GUŠIĆ, 1977;BASSOULLET, 1994;FU GAGNOLI & LORIGA BROGLIO, 1998).As discussed by FUGAGNOLI (2004, Fig. 5), the occurrence/absence of tests showing di-or trimorphism in complex large benthic Lower Jurassic foraminifers is triggered by the trophic regime.The genera Cymbriaella, Bosniella and Lituolipora are interpreted as characterizing oligotrophic conditions.For Bosniella croatica, GUŠIĆ (1977, p. 14) remarks that the dimorphism concerns "mainly the size of the tests and of the proloculus, while the general shape and basic growth pattern is the same for both types".Furthermore, GUŠIĆ states that in microspheric tests "the proloculus could not be observed " and "no uncoiled stage has been observed ".In accordance with this, FUGAGNOLI & LORIGA BROGLIO (1998, p. 64) also noticed "a more developed uncoiled stage of megalospheric forms".In Bosniella fontainei it seems to be just the opposite, with uncoiled megalospheric and coiled microspheric forms.A discrete dimorphism is not well pronounced in Bosniella bassoulleti; in fact, the assumed megalospheric specimens with subsphaerical proloculus are observable in both coiled (Fig. 11f, m) and uncoiled (Fig. 10d) forms.Specimens sectioned in the median plane without a detectable proloculus are assumed to represent microspheric forms.A possible difference in test morphology seems to be that microspheric forms are more fl attened lenticular than the macrospheric ones.
Stratigraphy: Bosniella bassoulleti appears within the upper part of the stratigraphic range of Timidonella sarda and Gutnicella cayeuxi, and has its maximum abundance immediately above the disappearance of these two species (see VELIĆ, 2007 for details).It disappears a little below the level with mass abundance of Selliporella donzellii, indicative of the Bajocian-Bathonian interval (e.g.GRANIER & DE-LOFFRE, 1994).For instance, a Selliporella donzellii cenozone (Bajocian) has been established in the Taurus Mountains of Turkey (ALTINER & SEPTFONTAINE, 1979), later modifi ed to a Timidonella sarda-Selliporella donzelli zone (EKMEKÇI & ALTINER, 2008).According to these data, the stratigraphic range of B. bassoulleti can be considered as Late (or latest) Aalenian-?Early Bajocian.

Origin of the name:
The genus refers to the city of Dubrovnik on the Adriatic coast of Croatia.
Diagnosis: Test free, irregularly planispirally to streptospirally coiled in early stage, nautiloid in form; later a slight tendency to uncoil may be present.Wall thin, fi nely agglutinating, with inner alveolar layer and outer imperforate epidermis.Alveoli simple, more or less parallel and more or less of equal diameter.Septa short and not alveolar.No en- doskeletal structures present.Aperture single, basal in the early stage, later becoming cribrate with only a few large openings.Most likely dimorphic.Megalospheric embryonic stage complex, bilocular, consisting of a large thin-walled subglobular megalosphere and a deuteroconch enveloping the former ½ to ¾.
Comparisons: The characteristic features enable the inclusion of Dubrovnikella n. gen.into the suborder Biokovinina KAMINSKI, defi ned as follows: "Test free or attached, may be coiled in the early stage, later uncoiled or branched.Wall fi nely agglutinated, traversed by pores, or with a coarsely perforate or canaliculated inner layer and an outer imperforate layer" (KAMINSKI, 2004, p. 250).In contrast to other representatives of the Biokovinidae, such as Bosniella GUŠIĆ and Biokovina GUŠIĆ, the pores in the wall of Dubrovnikella are distinctly coarser; it is noteworthy that in the diagnosis of the family Biokovinidae provided by LOEBLICH & TAPPAN (1987, p. 91) the wall is said to be "coarsely perforate".For the superfamily Biokovinacea, LOEBLICH & TAPPAN (1985, p. 94) characterize the wall as "fi nely agglutinated, with outer imperforate layer and coar sely perforate or canaliculated inner layer in later stage".Biokovina is unique within the biokovinids by possessing endoskeletal pillars.Such elements are lacking in Dubrovnikella n. gen.as well as in Bosniella.Both Bosniella and Biokovina may develop a straight uncoiled test portion, whereas in Dubrovnikella only in rare specimens, a slight tendency to uncoil was observed.A third genus that has been described as belonging to the Biokovinidae GUŠIĆ is Trochamijiella ATHERSUCH, BANNER & SIMMONS, 1992 (see also KAMINSKI, 2004, p. 250).As this genus has a test wall that "is solid and is not known to become protocanaliculate or canaliculated" (ATHERSUCH et al. 1992, p. 7) it cannot be included in the Biokovinidae sensu GUŠIĆ 1977 (see also LOEBLICH & TAPPAN, 1987).The complex embryonal apparatus consisting of a protoconch and deuteroconch or spheroconch (Fig. 12b) is reported from other Low er Jurassic foraminifera, e.g.Orbitopsella MUNIER-CHAL- MAS, 1902(e.g. HOTTINGER, 1967, 2006, Fig. 41h) or Cym briaella FUGAGNOLI, 1999 and Middle Jurassic larger benthic foraminifera, e.g.Bostia BASSOULLET, 1998.Dubrovnikella can generally be compared to some extent with Lituolipora GUŠIĆ & VELIĆ and Cymbriaella FUGAG-NOLI.Lituolipora with a more irregular coiling and a discoidal test form, has a microgranular wall "traversed by large pores that open both to exterior and interior" (LOEBLICH & TAPPAN, 1987, p. 91) whereas in Dubrovnikella the pores are covered to the exterior by a thin imperforate layer.Cymbriaella has a prominent uncoiled test portion, thick septa and an exoskeleton of beams and rafters; short beams are also present in the deuteroconch (see genus diagnosis of FU-GAGNOLI, 1999, p. 103).
Dubrovnikella septfontainei n. gen.n. sp.(Figs. 12b, Origin of the name: Dedicated to Michel Septfontaine for his numerous outstanding contributions to the knowledge of Jurassic larger benthic foraminifera. Holotype: Slightly oblique equatorial section of a macrospheric specimen fi gured in (Fig. 13c).Thin section SL 2.
Diagnosis: Being monotypic so far, the diagnosis is that for the genus.
Description: Test involute, subglobular to nautiloid in form, initially planispirally later may be slightly streptospirally coiled.Test usually higher than wide (height/thickness up to 1.55); thickness ranges from 0.6 mm to 1.15 mm, height 0.8 to 1.45 mm.Chambers broad with crescent-shaped chamber lumina; test periphery broadly rounded.Early coiling planispiral involute and biumbilicate, later with a slight tendency to be streptospiral, consisting of 1½ to 2 whorls with few chambers each (Figs.13b-c).The fi rst whorl comprises 4 to 5 chambers.During ontogeny, chambers continuously increase in size, becoming distinctly higher than long.In equ a torial section they fi rst appear rhombic later becoming more crescent-shape.An uncoiling of the last 1 to 2 chambers is rarely observable (Figs.13e-f).Sutures are depressed and recurved.The megalosphere (proloculus) is globular to subglobular, mostly laterally fl attened and enclosed by a thin microcrystalline (?originally organic) membrane (Fig. 12b).Its inner diameter (d) ranges from 0.15 mm to 0.23 mm, most values about 0.2 mm.The height (h) of the proloculus ranges from 0.13 mm to 0.21 mm, most frequently about 0.16 mm; the d/h ratio is 1.0 to 1.54, mostly about 1.25.The proloculus is enveloped by a semi-spherical deuteroconch of reduced height.The wall is thin and fi nely agglutinating with an inner alveolar layer with simple parallel pores (diameter 0.01 to 0.02 mm) and a thin outer imperforate epidermis (thickness 0.006 to 0.015 mm).The septa are solid and not alveolar.The aperture is basal single in the early stage, later becoming cribrate with only a few broad openings.No endo skeletal structures are present.
Remarks and comparisons: Being so far monospecifi c, see comparisons for the genus.
Stratigraphy: According to the foraminiferal assemblage at the type locality (Fig. 4), the stratigraphic range of Dubrovnikella septfontainei n. gen.n. sp.can be considered as latest Aalenian-earliest Bajocian.15a-d)

Remarks:
The Croatian specimens of Everticyclammina praevirguliana agree closely with the Late Sinemurian-Pliens bachian material of the Southern Alps of Italy described by FUGAGNOLI (2000).For other Liassic occurrences see BOU DA GHER-FADEL et al. (2001).In addition, E. praevirguliana was also reported (without illustration) from the Sinemurian-Pliensbachian of East Timor (HAIG et al., 2007).The stratigraphic range can now be extended to include the Upper Aalenian-Lower Bajocian interval.BOUDAGHER-FADEL et al. (2007) introduced a mid Sinemurian Everticyclammina praevirguliana biozone for different sections in the Mediterranean realm.
Stratigraphy: Everticyclammina praevirguliana illustrated in this paper is of Alenian age -it is accompanied by Gutnicella cayeuxi, but somewhat earlier of the fi rst occurences of Timidonella sarda.

REMARKS ON THE PHYLOGENETIC EVOLUTION OF LOWER TO MIDDLE JURASSIC LARGER BENTHIC FORAMINIFERA
The phylogenetic and morphostructural evolution of Jurassic larger benthic foraminifera has been a topic of research and discussion for decades (HOTTINGER, 1967;SEPTFON-TAI NE, 1980, 1988;SEPTFONTAINE et al., 1991;KUZNET-SO VA et al., 1996;BOUDAGHER-FADEL, 2008;KAMIN-SKI et al., 2010).In 1980, SEPTFONTAINE presented a use ful hierarchical grouping, starting with the outer morphology (type of coiling), presence/absence of exoskeletal and/ or endoskeletal elements and other features (mainly aperture).The temporal evolution of these elements has been the basis for phylogenetic considerations (SEPTFONTAINE, 1988).With respect to the end-Triassic mass extinction (e.g.TANNER et al., 2004), the calculated diversity curve for agglutinated foraminifera of KAMINSKI et al. ( 2010) only shows a "small dip" at the Triassic/Jurassic boundary.Within the Jurassic period, KUZNETSOVA et al. (1996, p. 121) recognized two major evolutionary stages (megaphases): early to middle Jurassic and the late Jurassic to early Cretaceous.
Concerning the appearance of "many species", the authors recognize "obvious markers" at the base of the Bathonian, the upper Bathonian, the beginning of the Oxfordian and at the base of the Tithonian.More detailed information concerning this topic is provided by the data compilation of KA-MINSKI et al. (2010, Fig. 5) showing that the origination rate of agglutinated foraminiferal genera had its Phanerozoic maximum in the Hettangian.This prominent peak was na med the "Hettangian origination" by KAMINSKI et al. (2010), actually covering the Hettangian-Pliensbachian interval and "characterized by the fi rst appearance of some larger agglutinated foraminiferal genera, derived from Lituola, such as Cymbriaella, Haurania, Orbitopsella, Everticyclammina, Lituosepta, Biokovina etc." Considering the fi rst appearances of these genera, however, there are differing data obtainable from the literature: Cymbriaella FUGAGNOLI: So far monospecifi c Cymbriaella lorigae described by FUGAGNOLI (1999) from the Domerian (= Upper Pliensbachian) of the Southern Alps, Ita ly.Actually we don´t know of any other record in the literature.The fi rst appearance at the base of the Hettangian as indicated by BOUDAGHER-FADEL (2008, p. 171 and Fig. 4.6), cannot be understood.It is noteworthy that FUGAG-NOLI (2004, p. 114) again reinstated the restriction to the Upper Pliensbachian.Our data allows the enlargement of the stratigraphic range into the Lower Bajocian.
Lituosepta CATI: So far monospecifi c with Lituosepta recoarensis described by CATI (1959) from the Pliensbachian of the Southern Alps.According to BASSOULLET (1997) and VELIĆ (2007) it is known from the Latest Early Sinemurian to the earliest Lower Pliensbachian .
Summarizing these data, there is a more or less parallel appearance of different genera in the Late Sinemurian, whereas in the Hettangian-Lower Sinemurian there is a lack of larger agglutinated benthic foraminifera.In addition to the 5 genera mentioned above, (except Haurania/Amijiella), Lituo lipora GUŠIĆ & VELIĆ also appears at the beginning of the Late Sinemurian (VELIĆ, 2007).These data do not show a Hettangian but instead a (Late) Sinemurian origination event.This fi ts better with the data of other invertebrates, showing a minimum number of genera in the Hettangian, the "sur-ance of favourable shallow-water facies, e.g., the Trento plateau in the southern Alps (e.g.FUGAGNOLI & LORIGA BROGLIO, 1998) or the Saharan Platform (e.g.WILMSEN & NEUWEILER, 2007).It is worth mentioning, that this extinction event in platform-dwelling larger benthic foraminifera didn´t happen exactly at the Pliensbachian-Toarcian boundary, but covered the whole Pliensbachian (BASSOUL-LET, 1997, Tab.LI).Biokovina and Orbitopsella did not survive the intra-Pliensbachian extinction as was the general assumption also for Cymbriaella prior to our new data.Then in the Late Aalenian, Cymbriaella lorigae and Everticyclammina praevirguliana are recorded again and the new genus Dubrovnikella appeared.Until now, there were no records of Everticyclammina known from the Late Pliensbachian un til the Callovian/Oxfordian (FUGAGNOLI, 2000, Fig. 4).The Late Aalenian Croatian fi ndings fi ll this gap.
After the Toarcian crisis, some new genera arose such as  vival phase" in the aftermath of the end-Triassic mass-extinction, then increasing steadily during the Sinemurian ("recovery phase") (e.g.WILMSEN & NEUWEILER, 2007, Fig. 17;ROMANO et al., 2008, Fig. 6).For the Hettangian to Low er Sinemurian Tethyan shallow-water carbonates, assemblages of small-sized textularids and miliolids together with opportunistic taxa such as Duotaxis, are recorded (e.g.BOUDAGHER-FADEL & BOSENCE, 2007), interpreted as refl ecting eutrophic conditions (FUGAGNOLI, 2004).Especially in the Late Sinemurian to Early Pliensbachian interval, larger benthic foraminifera show a progressive diversifi cation (e.g.BOUDAGHER-FADEL & BOSENCE, 2007).There is a signifi cant reduction of larger benthic foraminifera in the Toarcian, following a step-by-step extinction of several species and genera during the Pliensbachian (see BAS-SOULLET, 1997, Tab. 51;VELIĆ, 2007, Fig. 7).This extinction event can be seen in connection with a general biological crisis of other groups such as ammonoids that have been linked with the Toarcian oceanic anoxic event (OAE) (e.g.JENKYNS, 1988;WIGNALL & HALLAM, 1997;CEC CA & MACCHIONI, 2004;BILOTTA et al., 2010).Many platforms were drowned, resulting in the disappear- kella n. gen.Previously, the Middle Jurassic shallow-water carbonates of the Karst Dinarides were considered to be poor in benthic foraminifera.New investigations, however, have shown that these are distinctly richer in taxa than the Lower Jurassic strata; many of these representing index forms (VE-LIĆ, 2007).Our new fi ndings can be considered as further evidence of this observation.

ACKNOWLEDGMENT
Field and laboratory investigations were supported by the Croatian Geological Survey and the Croatian Ministry of science, education and sports of the Republic of Croatia, projects 1951953068-0241 "Microfossil assemblages in carbonate deposits of the Karst Dinarides", and 181-1811096-1093 "Basic geological map of the Republic of Croatia 1:50.000".We kindly acknowledge the helpful comments of the two reviewers Ivan GUŠIĆ (Zagreb) and Kemal TASLI (Mersin).A warm thank to Julie ROBSON for the English review of the paper.

Figure 3 :
Figure 3: Lithostratigraphy of the Osojnik section and distribution of important larger benthic foraminifera.

Figure 5 :
Figure 5: Lithostratigraphy of the Slano section and distribution of important larger benthic foraminifera.

Figure 7 :
Figure 7: Lithostratigraphy of the Biokovo section and the distribution of important larger benthic foraminifera.

Figure 8 :
Figure 8: Cymbriaella lorigae FUGAGNOLI.Upper Aalenian-Lower Bajocian of the Croatian Karst Dinarides.a) Equatorial section, uncoiled portion with 5 chambers.Note incorporation of small particles (peloids, foraminifera) into the wall and septa.b) Equatorial section, slightly oblique.c) Axial section, slightly oblique.d-e) Equatorial sections with 5 and 3 chambers in the uncoiled portion.Note single aperture in the coiled part becoming multiple in the uncoiled part and both thick wall and septa.f ) Detail of a showing alveolar wall.g) Oblique section through the uncoiled part.Scale bar 1mm, except f 0.5 mm.Thin sections: a SL 3A/3, b SL 3B/2, c SL 3B/1, d SL 5B2, e SL 3B/2, g SL 3A.

Figure 10 :
Figure 10: Bosniella bassoulleti n. sp.Upper Aalenian-Lower Bajocian of the Croatian Karst Dinarides.a-c, e-q Isotypes.d Holotype.a-d) Equatorial sections, in part slightly oblique.Holotype specimen (d) with uncoiled part consisting of three chambers; note multiple foramina evolving in the uncoiled portion (arrows).e-f) Specimens with prominent uncoiled part with 6 (?7) chambers in f.In e the chambers of the uncoiled part widen continuously, whereas in f the width remains more or less constant.g-h) Two specimens in diff erent sections.Note distinct biumbonal depression of the upper specimen in g cut in axial section and also the change of growth direction in the ultimate whorl.i-m) Oblique sections.n) Subaxial section; note irregular planispiral coiling resulting in a bent test.o) Specimen with large uncoiled portion.p) Two specimens, small juvenile specimen above in subaxial section, the other one below in oblique section.q) Equatorial section, slightly oblique; coiled portion showing 2 ½ whorls, uncoiled portion with 4 chambers.Scale bar 0.5 mm for all pictures.Thin sections: a, i-j, p O 32b, b, d, g Os10/1, c, h O 32a, e SZ 64/4, f SL 2/1, k O 35, l SL 2/1, m SL 4/1, n, q SL 4a, o SZ 64/5.

Figure 11 :
Figure 11: Bosniella bassoulleti n. sp.Upper Aalenian-Lower Bajocian of the Croatian Karst Dinarides.a-m Isotypes.a) Equatorial section; on the right Gutnicella cayeuxi (Lucas).b-c) (Sub)axial sections, in part slightly oblique.Note distinct biumbonal depression in c and oscillating coiling direction in b. d) Magnifi cation from the holotype specimen of Fig. 10d showing the poorly recognizable keriothecal wall structure.e-i) (Sub)axial sections, in part slightly oblique.Note signifi cant increase in chamber width within the ultimate whorl in e and f. j) Broken uncoiled portion; note the more or less constant chamber width.k-m) (Sub)axial sections.Note sigmoidal test bending of specimen shown in k.Scale bar 0.3 mm in d, all other pictures 0.5 mm.Thin sections: a SL 4a/1, b, i SL 2/1, c SL 3/b/2, d Os 10/1, e, f, g, h, m O 32/b, j SL 3A/3, k SL 4/4, l Os 10.

Figure 13 :
Figure 13: Dubrovnikella septfontainei n. gen., n. sp.Upper Aalenian-Lower Bajocian of the Croatian Karst Dinarides.a) Grainstone with three specimens of Dubrovnikella septfontainei and other small benthic foraminifera.b-d) Equatorial sections, slightly oblique; c = holotype specimen.e) Oblique section.f) Detail from e showing wall structure.g-i) Oblique axial sections.Note two large chamber openings in (h) and embryonic apparatus in (i) detail of which is shown in Figure 12b.j-k) Tangential sections of the test.l) Broken equatorial section.Scale bar 1 mm for a, all other pictures 0.5 mm.Thin sections: a, g, j SL 2-2, b SL 2-5, c-d SL 2, e SL 3A-2, f SL 3A, h SL 4A-1, i SL 5B/2, k SL 2-3, l SL 4a-4.

Figure 15 :
Figure 15: Everticyclammina praevirguliana FUGAGNOLI from the Upper Aalenian-Lower Bajocian of Mount Biokovo section.a-b) Slightly oblique equatorial section of uncoiled specimen.c) Oblique axial section.d) Equatorial section.Due to the size of the proloculus in b-d, these can be referred to megalospheric forms.As the uncoiled portion of the microspheric specimen should not exceed two chambers, fi gure a can also be considered megalospheric.Note that the proloculus is not visible due to oblique sectioning.Scale bars 1 mm.Thin sections: a BST 8-2, b, d BST 8-3, c BST 8.

Table 3 :
Dimensions of Bosniella bassoulleti n. sp., Upper Aalenian-Lower Bajocian of Croatia.ed = equatorial diameter (mm), dup = diameter uncoiled part (mm), ad max = maximum axial diameter (mm), ad min = minimum axial diameter at umbilical region (mm), n = number of chambers in the last whorl, ne = number of chambers in the uncoiled test portion.Mean values were not calculated for those parametres with only a few data available.