Lower Cretaceous dinoflagellate cyst and acritarch stratigraphy of the Cismon APTICORE (Southern Alps, Italy)

Review of Palaeobotany and Palynology 108 (2000) 213–266 www.elsevier.nl/locate/revpalbo

Lower Cretaceous dinoflagellate cyst and acritarch stratigraphy of the Cismon APTICORE (Southern Alps, Italy) S. Torricelli * ENI S.p.A., Agip Division, STIG, P.O. Box 12069, 20100 Milan, Italy Received 6 October 1998; accepted for publication 6 September 1999

Abstract A pelagic sedimentary succession, virtually complete from the Upper Hauterivian to the Upper Aptian and unconformably overlain by the Middle-Upper Albian p.p., was continuously cored in the Belluno Basin (southern Alps, NE Italy) as part of the APTICORE Program. APTICORE at Cismon Valley penetrated 131.8 m of limestones, marls and black shales, with 100% recovery of good quality cored material. One hundred and forty-six samples recovered from the marl and shale beds of the Cismon core were processed and analyzed for palynomorphs. Most of them yielded relatively rich and fairly well preserved assemblages of marine and terrestrially-derived palynomorphs. The results of a qualitative study of dinoflagellate cysts and acritarchs are presented and discussed. The distributions of 150 taxa are tabulated against the chronostratigraphy independently established on the basis of original litho-, bio-, chemo-, magnetostratigraphic investigations and of correlations with extensively studied sections outcropping in the vicinity of the Cismon drill site. The acritarch Pinocchiodinium erbae gen. et sp. nov. is described. Due to its distinctive morphology and extremely constant occurrence also in the black shales of the Selli Level, it is proposed as a marker species for the Aptian sediments of the Tethys. The dinoflagellate cysts Kallosphaeridium dolomiticum sp. nov. and Nexosispinum hesperus brevispinosum subsp. nov. are described from the Upper Hauterivian. Additional taxonomic remarks are made about other dinoflagellate cyst species, including the emendations of Tanyosphaeridium magneticum Davies 1983 and Bourkidinium granulatum Morgan 1975. The biostratigraphic value of selected taxa is discussed and compared with data known both from the Tethyan and Boreal realms. In particular, the extinction of Bourkidinium granulatum emend. is proposed as the best dinoflagellate cyst event for the delimitation of the Hauterivian–Barremian boundary in the Northern Hemisphere. The first appearance datums of Prolixosphaeridium parvispinum and Odontochitina operculata, and the slightly younger last appearance datum of Nexosispinum vetusculum are confirmed as useful biohorizons for recognition of the lower part of the Upper Barremian and hence for the approximation of the Lower–Upper Barremian boundary. The last occurrences of Rhynchodiniopsis aptiana and Phoberocysta neocomica are calibrated in the basal Aptian. © 2000 Elsevier Science B.V. All rights reserved. Keywords: acritarchs; new taxa; biostratigraphy; dinoflagellate cysts; Italy; Lower Cretaceous; Southern Alps

* E-mail address: [email protected] 0034-6667/00/$ - see front matter © 2000 Elsevier Science B.V. All rights reserved. PII: S0 0 3 4 -6 6 6 7 ( 9 9 ) 0 0 04 1 - X

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1. Introduction Within the APTICORE Program (Larson et al., 1993), which aims to document the Earth’s responses to a period of major global climate change in the mid-Cretaceous, the coring and logging of the Lower Cretaceous section in the Cismon Valley (southern Alps, NE Italy, Fig. 1) were carried out with the purpose of recovering a reference section for the Hauterivian–Aptian interval at low latitudes ( Erba and Larson, 1998; Erba et al., 1999). The Cismon core penetrated 131.8 m of pelagic sediments corresponding to a total stratigraphic thickness of 116.7 m ( Fig. 2), extending from the upper portion of the Biancone Formation, which is the local equivalent of the tethyan Maiolica ( Upper Hauterivian–Barremian), to the lower portion of the Scaglia Variegata (Aptian–Albian). A major gap within the Scaglia Variegata, eliding most of the Upper Aptian, the Lower Albian and

most of the Middle Albian, was detected in core 9 at 7.80 m below the well head. One of the main lithostratigraphic features of the cored section is the occurrence of the Selli Level immediately above the base of the Scaglia Variegata. This unit is characterized by organic carbon-rich black shales and siliceous beds, and is thought to represent the Lower Aptian Oceanic Anoxic Subevent 1a of Arthur et al. (1990). Detailed multidisciplinary studies of the Cismon core, including integrated biostratigraphy, magnetostratigraphy, cyclostratigraphy and organic, inorganic and stable isotope geochemistry, were carried out by an international team of specialists and final results were summarized in a comprehensive paper ( Erba et al., 1999). The aim of the present study is to describe and illustrate the dinoflagellate cyst and acritarch assemblages recovered from the Cismon core, with particular emphasis on the biostratigraphical value and taxonomy of selected forms.

Fig. 1. Location map of the Cismon drill site in the southern Alps.

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2. Site location and geological setting The Cismon drill site is located in the Venetian Prealps, west of Feltre (Belluno Province), in the Valley of the River Cismon, northeastern Italy (Fig. 1). The Venetian Prealps are part of the southern Alps, a west–east structural unit with a general south vergence delimited by the Insubric Line to the north and by the Po Plain to the south, representing a portion of the southern margin of the Mesozoic Tethys. The southern Alps display an inherited Mesozoic background, consisting of a horst and graben morphology, related to the rifting which led to the opening of the Ligurian Ocean. In Jurassic–Cretaceous times, the main palaeostructural–palaeogeographic elements were, from west to east, the Lombardy Basin, the Trento Plateau, the Belluno Basin and the Friuli Platform. The Cismon drill site is located on the eastwarddeepening slope between the Trento Plateau and the Belluno Basin, at an estimated palaeodepth, during the Lower Cretaceous, of 1000–1500 m ( Erba and Larson, 1998).

3. Material and methods The Cismon core was drilled to a total depth of 131.8 m below well head (mbwh) with near 100% recovery of limestones and marly limestones, with subordinate marls, radiolarian beds and black shales. The total stratigraphic thickness of the cored section is 116.7 m (Fig. 2). A total of 146 rock samples, mostly recovered from the marl and shale beds of the cored succession, were processed for palynomorphs. The precise location of each sample is given in Figs. 3 and 4 (cm measured from the top of each core). The standard processing technique involved chemical treatment of 15–20 g of the sample with HCl to remove the calcareous fraction and with HF to remove silicates, sieving with a 15 mm nylon mesh, heavy liquid separation with ZnCl and centrifug2 ing to concentrate the residues. A first slide was prepared for each sample using part of the residue >15 mm (mounting medium: depex). A second slide was made after oxidation

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of the residues, except for samples with a very low organic content. The oxidation was carried out with Schulze’s solution (a solution of KClO in 3 HNO ) in order to clean the residue of minor 3 amorphous kerogen particles and pyrite. Whole slides of oxidized residues, which proved to be the most fossiliferous and were not affected by selective preservation, were counted in order to obtain the relative abundances of palynomorph taxa. However, non-oxidized residues were also examined to check for the presence of rare species. The palynological assemblages are composed of bisaccate and inaperturate pollen grains, trilete spores, dinoflagellate cysts, foraminiferal test linings and acritarchs in fluctuating relative abundances. In general the preservation is ranging from moderate to good. Within the scope of the present study only the dinoflagellate cyst and acritarch record will be qualitatively discussed. Light photomicrographs were taken using a Zeiss Axioplan microscope and interferencecontrast illumination. For illustrated specimens England Finder Coordinates ( E.F.C.) are provided. For taxonomic citations reference is made to Williams et al. (1998). Previously undescribed taxa and those not included in Williams et al. (1998) are treated in Appendices A and B. All the preparation slides examined in this study are housed in the palynological slide collection at the Stratigraphic Department (STIG) of ENI S.p.A., Agip Division, San Donato Milanese, Italy.

4. Dinoflagellate cyst and acritarch biostratigraphy Out of the 146 samples prepared for palynomorphs, 108 proved to be fossilferous. The stratigraphical distributions of 150 dinoflagellate cyst and acritarch taxa encountered in the Cismon core are shown in the range chart of Fig. 3 with the counting results plotted against the chronostratigraphy independently established on the basis of original litho-, bio-, chemo-, magnetostratigraphic investigations ( Erba et al., 1999) and of correlations with adjacent sections (Channell et al., 1979; Erba, 1994; Cecca et al., 1996; Erba and Larson, 1998). Unidentifiable dinoflagellate cysts were not

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Fig. 2. Litho-chronostratigraphic log of the Cismon core after Erba et al. (1999). The column on the left represents the total penetration below well head. On the right, lithostratigraphic and chronostratigraphic boundaries are converted according to the bed dip and represent the real stratigraphic thickness of the cored succession.

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Fig. 3. Stratigraphical distribution of dinoflagellates and acritarchs in the Cismon core. Numbers refer to counted specimens in the whole slide. Undetermined taxa are not tabulated but constitute <15% of the overall assemblages. Samples missing with respect to progressive numbering from PAT BIS 1 to PAT BIS 11 and from PAT 1 to PAT 135 proved to be barren or yielded only undetermined palynomorphs and/or extremely poor and hence not diagnostic assemblages. Vertical scale not proportional to stratigraphic thicknesses.

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Fig. 3. (continued )

counted, but apparently constitute <15% of the overall assemblages. Reworked specimens of Jurassic dinoflagellate cysts constitute a minor percentage of the examined material and are included in the counting. In general, they were not

identified by their different state of preservation but by their known stratigraphical ranges. By contrast, reworking of earliest Cretaceous material was very difficult to identify and is probably almost absent, as extremely rare specimens of Berriasian–

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Fig. 3. (continued )

Valanginian–Lower Hauterivian species were detected. The stratigraphical ranges of 35 age diagnostic taxa, recorded in the Cismon core and selected from Fig. 3, are shown in Fig. 4. Since dinoflagel-

late cysts document paleoenvironmental changes, most of these taxa display significantly differing ranges compared with those known from the literature on Lower Cretaceous, due to the paleoenvironmental control on their geographical and

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Fig. 4. The total stratigraphic ranges recorded in the Cismon core for 35 taxa selected from the composite range-chart of Fig. 3, ordered according to first occurrences. Index numbers as in Fig. 3. Vertical scale not proportional to stratigraphic thicknesses.

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temporal distributions. In exploring synchronism of dinoflagellate cyst events (first and last occurrences), we realize that most of the events recorded in the present study are not directly related to evolutionary inceptions (oldest occurrence or first appearance datum) or extinctions (youngest occurrence or last appearance datum), but possess only regional significance. For some species, however, synchronism exists between events recorded in this study and those known from other Tethyan and Boreal sections with a good chronostratigraphic control. These key events can be regarded as true first and last appearance datums ( FADs and LADs) useful for delineating some Lower Cretaceous stage and substage boundaries on a wide geographic extent. The Hauterivian-Barremian boundary was placed in the Cismon core by Erba et al. (1999) at 69.40 m within core 34, immediately above the Faraoni Level equivalent (Fig. 2), a lithostratigraphic marker attributed to the uppermost Hauterivian on the basis of the ammonite content recovered from two outcrop sections close to the Cismon drill site (Cecca et al., 1996). In investigating the dinoflagellate cyst events approximating at this boundary, it must be considered that samples collected from the uppermost Hauterivian, within the reddish-pinkish pseudonodular limestones of cores 36–40 (from 72.54 to 79.57 m), were barren of palynomorphs, probably due to the unfavorable lithology. Furthermore, at the top of core 35, within the Faraoni Level equivalent (uppermost Hauterivian), the dinoflagellate cyst assemblage is not diagnostic due to the abundant amorphous organic matter persisting both in the oxidized and non-oxidized residue. In this frame it is reasonable to suppose that the LAD of Bourkidinium granulatum emend. marks the Hauterivian–Barremian boundary, since the occurrence of this species is consistent and often abundant throughout the lower portion of the Cismon core and its range top lies at 80.60 m in core 41, in the youngest fossiliferous sample of Hauterivian age. This datum confirms the biostratigraphical importance of the extinction of B. granulatum pointed out in the Tethyan Realm by Leereveld (1995), who correlated the last occurrence of Bourkidinium spp. to the top of the angulicostata ammonite Zone in

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SE Spain consistently with unpublished data of ammonite-controlled sections from Switzerland (Pourtoy, 1989), SE France (Londeix, 1990) and NW Germany (Lutat, 1995). In addition, the absence of B. granulatum in the post-Hauterivian interval of the Cismon core ( Fig. 3) indicates that no significant amount of Hauterivian reworking occurs in the Barremian–Aptian interval. The previous records of Bourkinidium granulatum emend. are from the Valanginian to the Upper Hauterivian of the North Sea [documented as aff. B. granulatum by Heilmann-Clausen (1987)], from the Lower Hauterivian of NW Germany [documented as Tanyosphaeridium magneticum by Pro¨ssl (1990)] and from the Hauterivian of the Monte Soro Flysch, Sicily, southern Italy ( Torricelli, personal observation). These represent further data supporting the pre-Barremian stratigraphical range of B. granulatum both in the Tethyan and in Boreal realms. The rare post-Hauterivian occurrences of this species so far documented in the Northern Hemisphere (Singh, 1983; Nøhr-Hansen, 1993), could be related to reworking episodes and/or to the imprecise chronostratigraphical control of the sections. The dinoflagellate cyst species Cymososphaeridium validum has a similar stratigraphical record to Bourkinidium granulatum, although the former becomes extinct in core 42 at 83.50 m in a slightly older horizon than the latter, thus consistently with results of Heilmann-Clausen (1987), Pro¨ssl (1990) and Leereveld (1995, 1997). In particular Leereveld (1997) used the last occurrence of C. validum as an additional criterion for recognition of the top of the A. eilema dinoflagellate cyst Zone in the Upper Hauterivian of the western Mediterranean area. Therefore, it is reasonable to consider the last occurrence of this species in the Upper Hauterivian of the Cismon core as its true LAD. A distinctive element of the dinoflagellate cyst assemblages encountered in the Upper Hauterivian of the Cismon core is the prominent and regular presence of Kallosphaeridium dolomiticum sp. nov. and of Oligosphaeridium totum subsp. totum. The youngest occurrence of the latter in core 42 at 83.50 m corresponds with the range top of Cymososphaeridium validum: this is an unprece-

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dented record for O. totum subsp. totum, as it was mostly reported from the Aptian–Albian (Brideaux, 1971; Singh, 1971; Brideaux and McIntyre, 1975; Bujak and Williams, 1978; Below, ˚ rhus, 1991; Nøhr-Hansen, 1993). 1982, 1984; A Nevertheless, although rare and sporadic, the Upper Berriasian through Lower Barremian occurrence of O. totum subsp. totum was already documented in Switzerland by Pourtoy (1989). Based on the litho-, bio-, chemo-, magnetostratigraphy, Erba et al. (1999) recognized the Lower– Upper Barremian boundary within core 29 at 60.40 m. The first appearances of the distinctive dinoflagellate cyst species Odontochitina operculata and Prolixosphaeridium parvispinum, in core 26 at 53.70 m and in core 27 at 56.0 m respectively, occur within the magnetic chron M1, hence in the lower part of the Upper Barremian. This correlates with the first occurrences reported for these species in SE Spain (Leereveld, 1995, 1997) and SE France (De Rene´ville and Raynaud, 1981; Wilpshaar, 1995) at the base of the A. vandenheckei ammonite Zone. A similar age was documented for the first occurrences of O. operculata and P. parvispinum in some ammonite–belemnite dated sections of the Boreal Realm (Duxbury, 1977, 1980; Harding, 1990; Pro¨ssl, 1990), suggesting them as important biohorizons in the lower Upper Barremian of the Northern Hemisphere. Only a few specimens of Nexosispinum vetusculum were found in this study, but the last occurrence of this taxon in core 23 at 45.60 m within the lower Upper Barremian corresponds with other data from the Boreal and Tethyan realms (Duxbury, 1980; Harding, 1990; Leereveld, 1995, 1997). Therefore, the close FADs of Odontochitina operculata and Prolixosphaeridium parvispinum and the slightly younger LAD of N. vetusculum are proposed as key events for recognition of the lower Upper Barremian and hence for the approximation of the Lower–Upper Barremian boundary on a wide geographic extent. The acme of the dinoflagellate cyst recorded here as Kleithriasphaeridium cf. fasciatum (see remarks in Appendix A) within a narrow interval straddling the Lower–Upper Barremian boundary in core 29 (from 59.50 to 60.90 m), seems to have local significance, since similar forms were pre-

viously reported to be restricted to the Upper Hauterivian of SE Spain (Leereveld, 1995, 1997). The Barremian interval of the Cismon core is characterized by the continuous and sometimes abundant occurrence of Rhynchodiniopsis aptiana. The last common occurrence of this taxon was recorded at 28.10 m in the basal part of core 16, immediately above the Barremian–Aptian boundary placed by Erba et al. (1999) at 29.17 m at the base of magnetic chron M0, following the recommendation of the Aptian Working Group ( Erba, 1996). Only rare and sporadic specimens of R. aptiana occur in the lowermost Aptian interval of the Cismon core succession. Therefore, this species, as previously discussed by Davey and Verdier (1974) and Heilmann-Clausen (1987), appears to be typical of the Barremian and basal Aptian, although it was sometimes reported also from the Lower and Upper Aptian. The occurrence of Phoberocysta neocomica is consistent in the Upper Hauterivian, but becomes discontinuous in the Barremian and the range top is at 28.10 m near the bottom of core 16, in the basal Aptian. The last occurrence of this species was reported in the lowermost Aptian of the Boreal Realm by Williams and Bujak (1985), Habib and Drugg (1987), Heilmann-Clausen (1987), Lister and Batten (1995). In more detail, Duxbury (1983), in an ammonite-controlled section from the Isle of Wight ( England), correlated the extinction of P. neocomica to the top of the D. forbesi ammonite Zone in the Lower Aptian. Some workers calibrated the extinction of P. neocomica in the lowermost Aptian also in the Tethyan Realm. In describing the microplankton assemblages from the Aptian type localities at La Be´doule and Gargas (SE France), Davey and Verdier (1974) explained the absence of P. neocomica by the fact that the oldest Aptian samples at La Be´doule ( limestone facies) were organically barren and that this species, which was previously recorded by Millioud (1969) in the Angles section, probably disappeared during the earliest Aptian with Rhynchodiniopses aptiana. Nevertheless, the use of the LAD of Phoberocysta neocomica for the recognition of the lowermost Aptian, and hence for the delimitation of the Barremian–Aptian boundary, has still to be

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confirmed with the evaluation of data published during the last decade, due to the different approach to the taxonomy of the Ceratioid group followed by the authors after the publication of Monteil (1991). It is noticeable, for instance, that Leereveld (1995) did not mention this species. In conclusion, the dinoflagellate cyst events discussed by Erba (1996) for the approximation of the Barremian–Aptian boundary, that is the last occurrence of Pseudoceratium pelliferum and the first occurrence of Cribroperidinium ?tenuiceras (reported as Tehamadinium tenuiceras) were not confirmed by the results of the present study, since in the Cismon core P. pelliferum has only a sporadic record in the Upper Hauterivian, whereas C. ?tenuiceras was not identified. The FAD of Pinocchiodinium erbae gen. et sp. nov. was recorded at 25.50 m in core 15, 1.82 m below the base of the Lower Aptian Selli Level. Due to its extremely distinctive morphology, consistent stratigraphic record even in the black shales of the Selli Level and occurrence in the Aptian member of the Monte Soro Flysch of Sicily ( Torricelli, personal observation), the total distribution of P. erbae is tentatively proposed as a major event for the Aptian at low paleolatitudes. The Aptian strata of the Cismon core represent only a part of the Aptian Stage, since the Upper Aptian G. ferreolensis planktonic foraminifer Zone is unconformably overlain by the Middle Albian T. primula planktonic foraminifer Zone ( Erba et al., 1999). This unconformity, located at 7.80 m near the bottom of core 9 (Fig. 2), is marked by a turnover in the dinoflagellate cyst assemblages from sample PAT 4 upwards ( Figs. 3 and 4), determined by the abrupt disappearance of Cassiculosphaeridia reticulata, Hystrichodinium pulchrum, Palaeoperidinium cretaceum, Pinocchiodinium erbae and by the first occurrences of Damassadinium chibane, Hystrichosphaeridium atlasiense, Ovoidinium diversum and Prolixosphaeridium conulum. Due to the presence of such an important hiatus (most of the Upper Aptian, the whole Lower Albian and most of the Middle Albian are missing), it is not possible to calibrate the chronostratigraphic position of these events. The only record of Litosphaeridium arundum and Litosphaeridium conispinum was in core 2 at

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2.10 m, in the Upper Albian. As six samples collected from the reddish sediments extending from core 4 to the upper part of core 9 resulted barren of palynomorphs, it was not possible to achieve sufficient information for calibrating the inceptions of these two species in the Cismon core.

5. Conclusions The distribution of 150 dinoflagellate cyst and acritarch taxa, recovered from 146 samples collected from an expanded pelagic succession continuously cored in the Lower Cretaceous sediments of the Belluno Basin (NE Italy), was tabulated against a reliable chronostratigraphy independently established on the basis of original litho-, bio-, chemo-, magnetostratigraphic investigations and of correlations with ammonite-dated sections in the immediate vicinity. Some of the key dinoflagellate cyst events discussed in the present paper ( Fig. 5) are consistent with published data obtained from ammonite– belemnite controlled sections and are therefore confirmed and/or proposed as useful tools for marking some chronostratigraphic boundaries in the Lower Cretaceous: (1) The LAD of Bourkidinium granulatum emend. nov. is a reliable biohorizon for the approximation of the Hauterivian–Barremian boundary in the Northern Hemisphere. It is slightly younger than the LAD of Cymososphaeridium validum. (2) The prominent occurrences of Oligosphaeridium totum subsp. totum and of Kallosphaeridium dolomiticum sp. nov. are distinctive elements of the Upper Hauterivian dinoflagellate cyst assemblages of the Cismon core, thus possessing local significance. (3) The near simultaneous FADs of Odotochitina operculata and Proloxosphaeridium parvispinum, and the slightly higher LAD of Nexospinum vetusculum represent events suitable for the recognition of the lower Upper Barremian on a wide geographic extent. (4) The acme of Kleithriasphaeridium cf. fasciatum characterizes a narrow interval of the Cismon core close to the Lower–Upper Barremian boundary. This event seems to possess local significance.

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Fig. 5. Calibration of selected dinoflagellate cyst events in the Cismon core. Magnetic stratigraphy after Erba et al. (1999).

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(5) Rhychodinopses aptiana is a typically Barremian species in the Tethyan Realm and displays its range top in the basal Aptian. (6) Despite problems with the taxonomy of the Ceratioid group, the LAD of Phoberocysta neocomica is most likely calibrated in the lowermost Aptian both in the Tethyan and Boreal realms. (7) Pinocchiodinium erbae gen. et sp. nov. is restricted to the Aptian and its total distribution is proposed as a major event at low palaeolatitudes. (8) Damassadinium chibane, Hystrichosphaeridium atlasiense, Oviodinium diversum, Prolixosphaeridium conulum, Litosphaeridium arundum and Litosphaeridium conispinum occur only in the Albian portion of the studied section. Since the Aptian–Albian transition in the Cismon core is not continuous but marked by a hiatus (most of the Upper Aptian, the whole Lower Albian and most of the Middle Albian are missing), it was not possible to calibrate the inceptions of these taxa and the extinction of Pinocchiodinium erbae gen. et sp. nov.

Acknowledgements The author would like to express his gratitude to U. Biffi (ENI, Milan) for his support to the study. E. Erba ( University of Milan) is warmly acknowledged for giving the author the opportunity to sample and study the Cismon core, for coordinating the project and for providing the location map and the lithostratigraphic log of Figs. 1 and 2. R.J. Davey, P. Hochuli, J.B. Riding, H. Visscher and K. Zonneveld are gratefully acknowledged for providing comments and suggestions which substantially improved the original version of the manuscript. The author is endebted to ENI-Agip Division management for permission to participate in the project and to publish the results.

Appendix A: Annotated species list Alphabetical listing of dinoflagellate cyst and acritarch taxa recovered from the Cismon core. Taxa dealt with in the systematic section are asteri-

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sked. Taxa illustrated are followed by plate and figure references in parentheses. The first number in parentheses refers to the position of the taxon in the range-charts (Figs. 3 and 4). The generic allocation and authorship of dinoflagellate cyst species follow Williams et al. (1998). Achomosphaera neptunii ( Eisenack, 1958) Davey and Williams, 1966 (1, Plate XX, 9). Aprobolocysta neista Duxbury, 1980 (73, Plate XV, 5). Aptea polymorpha Eisenack, 1958 (138, Plate IX, 10). Apteodinium granulatum ( Eisenack, 1958) LucasClark, 1987 (11, Plate XVII, 6). Apteodinium reticulatum Singh, 1971 (84, Plate XVII, 7). Apteodinium cf. A. maculatum subsp. grande (Cookson and Hughes, 1964) Below, 1981 (113). Apteodinium sp. A (33, Plate XVII, 5). Remarks: A species of Apteodinium with a slightly ovoid ambitus, a prominent apical horn and a clearly expressed paracingulum. The autophragm is thick, spongy and foveolate. Cyst length=65–70 mm; cyst width=52–56 mm; length of apical horn=8–9 mm. Apteodinium sp. A most closely resembles Apteodinium spiridoides Benedek, 1972 which displays a thick alveolate autophragm. The apical horn of the latter is, however, very short (<4 mm long) or missing and the paracingulum is not discernible. Stratigraphic occurrence: Rare in the Upper Hauterivian interval of the Cismon core. Similar forms were recorded from the Hauterivian of the Moscow Basin, Russia, by Iosifova (1996); Plate IV, 1 as Apteodinium sp. A of Stover and Helby. However, some undescribed specimens illustrated by Stover and Helby [1987b, p. 253, fig. 25(H–N )] as Apteodinium sp. A exhibit a reticulate rather than a foveolate ornamentation pattern. Atopodinium haromense Thomas and Cox, 1988 (23, Plate III, 6). Avellodinium falsificum Duxbury, 1977 (44, Plate XIV, 6). Avellodinium cf. A. falsificum Duxbury, 1977 (85, Plate XIV, 4 and 5).

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PLATE I

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PLATE II

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PLATE III

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Remarks: Dinoflagellate cysts here recorded as Avellodinium cf. falsificum are larger than the specimens of A. falsificum documented in the type material by Duxbury (1977) from Speeton,

England, and their processes are seldom proximally connected by low sutural crests. On the contrary, specimens from the type material have processes that are usually proximally connected

PLATE I 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

Batioladinium varigranosum ×750, sample PAT 106-ox, E.F.C. ×63/3. Batiacasphaera angularis ×750, sample PAT 3-ox, E.F.C. L 59. Batiacasphaera subtilis ×600, sample PAT 114-ox, E.F.C. S 39/4. Batiacasphaera macrogranulata ×750, sample PAT 106-ox, E.F.C. S 47/2. Batiacasphaera macrogranulata ×600, sample PAT 112-ox, E.F.C. V 20. Batiacasphaera saidensis ×750, sample PAT 7-ox, E.F.C. M 31. Batiacasphaera? sp. ×600, sample PAT 44-ox, E.F.C. K 50/2. Batiacasphaera? sp. ×600, sample PAT 121-ox, E.F.C. H 60. Kallosphaeridium ?helbyi subsp. psilatum ×750, sample PAT 4-ox, E.F.C. M 34/1. Ovoidinium diversum ×750, sample PAT 4-ox, E.F.C. F 55. Ovoidinium diversum ×750, sample PAT 4-ox, E.F.C. D 49/4. Ovoidinium incomptum ×750, sample PAT 3-ox, E.F.C. O 30/4.

PLATE II 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13.

Mendicodinium sp. ×550, sample PAT 16-ox, E.F.C. V 56 Dissiliodinium globulus ×550, sample PAT 13-ox, E.F.C. J 49 Cassiculosphaeridia magna ×550, sample PAT 121-ox, E.F.C. V 23/3 Cassiculosphaeridia reticulata ×680, sample PAT 7-ox, E.F.C. H 46/1 Cassiculosphaeridia reticulata ×500, sample PAT 12-ox, E.F.C. V 34 Cassiculosphaeridia reticulata ×500, sample PAT 11-ox, E.F.C. P 23 Cassiculosphaeridia reticulata ×500, sample PAT 9-ox, E.F.C. U 32 Cassiculosphaeridia reticulata ×500, sample PAT 85-ox, E.F.C. X 68 Cassiculosphaeridia reticulata ×550, sample PAT 13-ox, E.F.C. N 44 Valensiella tazadensis ×500, sample PAT 64-ox, E.F.C. Y 57 Sentusidinium spp. ×550, sample PAT 43, E.F.C. W 65 Sentusidinium spp. ×500, sample PAT 116-ox, E.F.C. J 62 Sentusidinium spp. ×550, sample PAT 128-ox, E.F.C. M 21/4

PLATE III 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

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Lithodinia stoveri ×600, sample PAT 109-ox, E.F.C. O 30/1 Lithodinia stoveri ×500, sample PAT 115-ox, E.F.C. D28 Lithodinia stoveri ×500, sample PAT 130-ox, E.F.C. L 66/2 Lithodinia amlasis ×550, sample PAT 112-ox, E.F.C. Y 33/1 cf. Desmocysta plekta ×600, sample PAT BIS 6-ox, E.F.C. U 18/2 Atopodinium haromense ×550, sample PAT 116-ox, E.F.C. R 22/3 cf. Tenua hystrix ×550, sample PAT 86, E.F.C. K 55 cf. Tenua hystrix ×550, sample PAT 128-ox, E.F.C. E 54/1 cf. Tenua hystrix, operculum, ×750, sample PAT 87-ox, E.F.C. G 27/1 Circulodinium distinctum ×500, sample PAT 95-ox, E.F.C. Y 28/1 Circulodinium distinctum longispinatum ×600, sample PAT 4-ox, E.F.C. Q 60 Cyclonephelium intonsum ×500, sample PAT 15-ox, E.F.C. P 36

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PLATE IV

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PLATE V

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PLATE VI

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by low sutural crests and occasionally few of the crests may be high and perforate. One of the encountered specimens exhibits faintly fibrous processes and a prominent crest in apical position (Plate XIV, 5). Stratigraphic occurrence: Rare from the uppermost Hauterivian to the Lower Barremian of the Cismon core. Batiacasphaera angularis Stevens and Helby, 1987 (139, Plate I, 2).

Batiacasphaera macrogranulata Morgan, 1975 (12, Plate I, 4 and 5). Batiacasphaera saidensis Below, 1981 (103, Plate I, 6). Remarks: The diameters of the specimens of B. saidensis recovered from the Cismon core (50– 60 mm) are generally larger than those of the type material (45–48 mm) documented by Below (1981) from the Late Hauterivian of Morocco, except for the holotype which is 56 mm wide.

PLATE IV 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

Prolixosphaeridium conulum ×750, sample PAT 4-ox, E.F.C. Q 53 Prolixosphaeridium parvispinum ×550, sample PAT 83-ox, E.F.C. E 72 Prolixosphaeridium parvispinum ×600, sample PAT 13-ox, E.F.C. D 48/2 Tanyosphaeridium spp. ×600, sample PAT 63-ox, E.F.C. M 30/3 Chlamydophorella nyei ×600, sample PAT 116-ox, E.F.C. K 31/1 Gochteodinia villosa multifurcata ×600, sample PAT 130-ox, E.F.C. S 54/2 Dapsilidinium warrenii ×550, sample PAT 113-ox, E.F.C. Q 66/2 Dapsilidinium warrenii ×600, sample PAT 115-ox, E.F.C. S 51/1 Kallosphaeridium dolomiticum sp. nov. ×600, sample PAT 128-ox, E.F.C. K 31/4. Holotype Dapsilidinium multispinosum ×600, sample PAT 5-ox, E.F.C. U 31/2 Chlamydophorella sp. A ×750, sample PAT 87, E.F.C. G 54/4 Kallosphaeridium dolomiticum sp. nov. ×750, sample PAT 132-ox, E.F.C. M 23/3

PLATE V 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11.

Bourkidinium granulatum ×600, sample PAT 133-ox, E.F.C. L 24. Specimen with 12 precingular processes Bourkidinium granulatum ×600, sample PAT 128-ox, E.F.C. V 19/2 Bourkidinium granulatum ×600, sample PAT 112-ox, E.F.C. P 61/1 Bourkidinium granulatum ×600, sample PAT 128-ox, E.F.C. Y 28 Bourkidinium granulatum ×600, sample PAT 107-ox, E.F.C. Q 56/3 Bourkidinium granulatum ×550, sample PAT 111-ox, E.F.C. P 36 Bourkidinium granulatum ×550, sample PAT 116-ox, E.F.C. N 58/2. Specimen with 10 precingular processes Bourkidinium granulatum ×750, sample PAT 115, E.F.C. L 63 Bourkidinium granulatum ×600, sample PAT 120-ox, E.F.C. T 57/3 Bourkidinium cf. B. ?cylindricum ×600, sample PAT 109-ox, E.F.C. K 54 Tanyosphaeridium spp. ×750, sample PAT 106-ox, E.F.C. N 52/4

PLATE VI 1. 2. 3. 4. 5. 6.

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Oligosphaeridium ?asterigerum ×550, sample PAT 85-ox, E.F.C. W 33 Oligosphaeridium cf. O. complex ×600, sample PAT 125-ox, E.F.C. Y 33/1 Oligosphaeridium ?asterigerum ×600, sample PAT 16-ox, E.F.C. G 23/2 Cymososphaeridium validum ×550, sample PAT 133-ox, E.F.C. V 55 Cymososphaeridium validum ×600, sample PAT 120-ox, E.F.C. W 21/3 Oligosphaeridium cf. O. complex ×600, sample PAT 109-ox, E.F.C. E 28

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PLATE VII

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PLATE VIII

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PLATE IX

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Stratigraphic occurrence: Sporadic in the Upper Barremian and continuous in the Aptian of the Cismon core. Batiacasphaera subtilis Stover and Helby, 1987 (81, Plate I, 3). Batiacasphaera cf. B. ovata Backhouse, 1987 (140a) Remarks: This form resembles B. ovata in the ovoidal outline, similar dimensions and archeopyle margin characterised by short accessory sutures, with a marked sulcal notch and free operculum. However, it displays a coarser granular ornamentation with rows of interconnected

PLATE VII 1. Oligosphaeridium porosum ×550, sample PAT 96-ox, E.F.C. O 55 2. Oligosphaeridium totum ssp. minus ×550, sample PAT 85-ox, E.F.C. T 22 3. Oligosphaeridium complex ×600, sample PAT 53-ox, E.F.C. F 52 4. Oligosphaeridium totum ssp. totum ×550, sample PAT 129-ox, E.F.C. U 53/3 5. Oligosphaeridium albertense ×550, sample PAT 88-ox, E.F.C. O 50/3 6. Oligosphaeridium porosum ×750, sample PAT 15-ox, E.F.C. C 56/2 7. Oligosphaeridium poculum ×750, sample PAT 51-ox, E.F.C. Q 27/3

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gemmae particularly evident in the equatorial region. Furthermore, the thickening of the autophragm from the apical regions towards the cingular area is not so noticeable as in B. ovata. Stratigraphic occurrence: Sporadic in the Albian of the Cismon core. Batiacasphaera? sp. (2, Plate I, 7 and 8). Batioladinium micropodum ( Eisenack and Cookson, 1960) Brideaux, 1975 (148). Batioladinium varigranosum (Duxbury, 1977) Davey, 1982 (89, Plate I, 1). *B. granulatum Morgan, 1975; emend. (13, Plate V, 1–9). Bourkidinium cf. B. ?cylindricum Dolding, 1992 (90, Plate V, 10). Comments: Specimens recovered from the Cismon core and assigned to B. cf. ?cylindricum display a similar central body but longer processes (11–14 mm) in comparison to specimens of B. ?cylindricum documented from the type material (7–11 mm). Stratigraphic occurrence: Rare in the uppermost Hauterivian of the Cismon core. Dolding (1992) described the type material (five specimens) from the late Campanian sediments of Humps Island,

PLATE IX

PLATE VIII 1. Hystrichosphaeridium atlasiense ×750, sample PAT 4-ox, E.F.C. Q 52 2. Hystrichosphaeridium atlasiense ×600, sample PAT 4-ox, E.F.C. M 38/1 3. Hystrichosphaeridium atlasiense ×600, sample PAT BIS 6-ox, E.F.C. P 37 4. Litosphaeridium arundum ×750, sample PAT BIS 1, E.F.C. J 26/1 5. Litosphaeridium arundum ×750, sample PAT BIS 1, E.F.C. K 26/4 6. Florentinia mantellii ×750, sample PAT BIS 5-ox, E.F.C. V 36 7. Florentinia mantellii ×550, sample PAT BIS 6-ox, E.F.C. O 20 8. Florentinia cf. F. abjuncta ×550, sample PAT 20-ox, E.F.C. M 53/3 9. Hystrichosphaeridium atlasiense ×600, sample PAT 4-ox, E.F.C. T 57/1 10. Dapsilidinium sp. ×550, sample PAT 8-ox, E.F.C. S 57/3

1. Systematophora spp. ×550, sample PAT 124-ox, E.F.C. L 26 2. Systematophora spp. ×550, sample PAT 110-ox, E.F.C. T 30 3. Systematophora spp. ×600, sample PAT 117-ox, E.F.C. U 36/1 4. Cometodinium spp. ×600, sample PAT 120, E.F.C. N 57/1 5. Cometodinium spp. ×750, sample PAT 108-ox, E.F.C. P 48 6. Systematophora ?daveyi ×550, sample PAT 116-ox, E.F.C. M 54/1 7. Downiesphaeridium aciculare ×600, sample PAT 134-ox, E.F.C. R 26 8. Downiesphaeridium sp. ×550, sample PAT 113-ox, E.F.C. M 39/2 9. Heterosphaeridium araneosum ×550, sample PAT 116-ox, E.F.C. L 39 10. Aptea polymorpha ×550, sample PAT 4-ox, E.F.C. O 59 11. Pseudoceratium eisenackii ×550, sample PAT 4-ox, E.F.C. K 54. 12. Pseudoceratium eisenackii ×550, sample PAT 12-ox, E.F.C. T 52/4

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PLATE X

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Antarctic Peninsula, pointing out the presence of abundant Permian and Lower Cretaceous recycled palynomorphs in his original assemblages. Callaiosphaeridium asymmetricum (Deflandre and Couteville, 1939) Davey and Williams, 1966 (34, Plate XIV, 1). Cassiculosphaeridia magna Davey, 1974 (10, Plate II, 3). Cassiculosphaeridia reticulata Davey, 1969 (43, Plate II, 4–9). Remarks: The specimens herein assigned to this taxon, which were abundantly recovered from several samples, are characterized by a subspherical outline with a variable development of reticulation, also within the same assemblage, ranging from forms with a complete reticulum outlining polygonal lumina (Plate II, 4 and 7), to forms ornamented with low, more or less continuous, smooth ridges forming an incomplete and relatively large mesh reticulum (Plate II, 8 and 9). Stratigraphic occurrence: From the Upper Hauterivian to the Upper Aptian unconformity, particularly abundant in the Barremian–Aptian interval of the Cismon core. Cerebrocysta sp. (72, Plate XV, 3). Chlamydophorella nyei Cookson and Eisenack, 1958 (74, Plate IV, 5). Chlamydophorella sp. A (97, Plate IV, 11). Remarks: Holocavate cysts with a spheroidal central body densely covered by broad, tubular and distally flared processes supporting a thin, discontinuous ectophragm. Archeopyle apical,

PLATE X 1. Pseudoceratium retusum ×600, operculum, sample PAT 13-ox, E.F.C. R 27/1 2. Pseudoceratium retusum ×600, sample PAT 13-ox, E.F.C. O 36/2 3. Pseudoceratium retusum ×600, sample PAT 13-ox, E.F.C. K 53/2 4. Pseudoceratium retusum ×600, sample PAT 13-ox, E.F.C. H 49 5. Pseudoceratium retusum ×600, sample PAT 13-ox, E.F.C. Y 30 6. Pseudoceratium pelliferum ×600, sample PAT 128-ox, E.F.C. Y 27/4

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type tA. Apical protuberance absent. Overall diameter: 40–50 mm. The processes of Chlamydophorella sp. A are considerably broader than those of C. nyei. Stratigraphic occurrence: This form is common in a narrow interval within the uppermost Lower Barremian of the Cismon core. Forms similar to Chlamydophorella sp. A were reported by Srivastava (1984, Plate 6, 3–5) as Cleistosphaeridium ancoriferum from the Upper Barremian of SE France. Chytroeisphaeridia ?scabrata Pocock, 1972 (14, Plate XXI, 9). Circulodinium brevispinosum (Pocock, 1962) Jansonius, 1986/Circulodinium distinctum (Deflandre and Cookson, 1955) Jansonius, 1986 (3, Plate III, 10). Circulodinium distinctum subsp. longispinatum (Davey, 1978) Lentin and Williams, 1989 (132, Plate III, 11). Remarks: Due to the long stratigraphical range and to the intergrading nature of their morphological features, no attempt has been made in the present study to distinguish forms with short spines (C. brevispinosum) from forms with longer spines (C. distinctum), except for specimens characterised by long spines (>10 mm) which were assigned to C. distinctum subsp. longispinatum (Plate III, 11). Stratigraphic occurrence: The C. brevispinosum/distinctum group was recorded throughout the Cismon core, particularly abundant in the Upper Hauterivian–Lower Barremian interval. By contrast, C. distinctum subsp. longispinatum was found in the Albian only. Cithariplana caperata Srivastava, 1984 (119, Plate XXI, 10). Cometodinium spp. (24, Plate IX, 4 and 5). Coronifera oceanica Cookson and Eisenack, 1958 (93, Plate XV, 7). Cribroperidinium spp. (5, Plate XVII, 2–4). Remarks: Apart from the identification of Cribroperidinium boreas (Plate XVII, 1) and Cribroperidinium magnum (Plate XV, 1), no attempt has been made to distinguish other morphologically closely related and highly variable species of Cribroperidinium. Stratigraphic occurrence: This genus is regularly

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PLATE XI

1. 2. 3. 4. 5. 6. 7. 8.

Muderongia Muderongia Muderongia Muderongia Muderongia Muderongia Muderongia Muderongia

staurota ×500, sample PAT 106-ox, E.F.C. Y 60 perforata ×600, sample PAT 109-ox, E.F.C. G 64 tabulata sensu Monteil ×550, sample PAT 86, E.F.C. T 27/2 tabulata sensu Monteil ×550, sample PAT 86, E.F.C. Q 25 tabulata sensu Monteil ×550, sample PAT 86, E.F.C. F 33 siciliana ×600, sample PAT 107-ox, E.F.C. T 65 aequicornis ×600, sample PAT 34, E.F.C. G 28/3 testudinaria ×500, sample PAT 115, E.F.C. F 54/3

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present, but never abundant, throughout the Cismon core. Cribroperidinium boreas (Davey, 1974) Helenes, 1984 (121, Plate XVII, 1). Cribroperidinium magnum (Jain, 1977) Williams et al., 1998 (64, Plate XV, 1). Ctenidodinium elegantulum Millioud, 1969 (48, Plate XIV, 3). Cyclonephelium intonsum Duxbury, 1983 (128, Plate III, 12). C. validum Davey, 1982 (6, Plate VI, 4 and 5). Damassadinium chibane (Below, 1981) Fensome et al., 1993 (133, Plate XX, 7). Remarks: Forms herein retained as Damassadinium chibane are slightly smaller than those of the type material described and illustrated from the Aptian (Gargasian) of Morocco by Below (1981). Stratigraphic occurrence: Five specimens were recorded at 7.50 m from the Middle Albian (T. primula planktonic foraminifer Zone). Dapsilidinium laminaspinosum (Davey and Williams, 1966) Lentin and Williams, 1981 (142). Dapsilidinium multispinosum (Davey, 1974) Bujak et al., 1980 (29, Plate IV, 10). Dapsilidinium warrenii (Habib, 1976) Lentin and Williams, 1981 (7, Plate IV, 7 and 8). cf. Desmocysta plekta Duxbury, 1983 (141, Plate III, 5). Remarks: The specimens of D. plekta described from type material by Duxbury (1983) from the Aptian of the Isle of Wight, southern England, exhibit a two-plate precingular archeopyle. Two specimens which were recorded in the present study are very similar in size and appearence to D. plekta, but differ from the holotype by having an apical archeopyle. Stratigraphic occurrence: Rare in the Upper Albian of the Cismon core. Cysts with an apical archeopyle were recorded by Nøhr-Hansen (1993) from the Upper Albian of NE Greenland as D. plekta although, according to this author (p. 65), ‘‘the different archeopyle type may advocate the erection of a new species and maybe also a new genus’’. Dingodinium cerviculum Cookson and Eisenack, 1958 (67, Plates XII, 7 and XV, 10). Remarks: Stover and Helby (1987a, p. 282)

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retained D. cerviculum and Dingodinium albertii Sarjeant 1966 as two separate species, stating that, in addition to the difference in size, the endocysts of D. albertii tend to be subcircular in outline rather than elliptical, whereas the epipericoels are much reduced in comparison to those of D. cerviculum. I accept the taxonomic choice of Williams et al. (1998) who considered D. cerviculum a senjor synonym of D. albertii due to the intergrading nature of their morphological features and to their very similar stratigraphical distributions. Stratigraphic occurrence: This taxon exhibits a sporadic record from the Upper Hauterivian to the Upper Barremian of the Cismon core. Dissiliodinium globulus Drugg, 1978 (35, Plate II, 2). Downiesphaeridium spp. (4, Plate IX, 8). Downiesphaeridium aciculare (Davey, 1969) Islam, 1993 (15, Plate IX, 7). Endoscrinium campanula (Gocht, 1959) Vozzhennikova, 1967 (25, Plate XXI, 7 and 8). Exochosphaeridium muelleri Yun, 1981/Exochosphaeridium phragmites Davey et al., 1966 (17, Plate XXI, 3 and 4). Remarks: Due to the intergrading nature of the characteristic features, no attempt has been made to distinguish between E. phragmites and E. muelleri. Stratigraphic occurrence: The E. phragmites/ muelleri group occurs from the bottom ( Upper Hauterivian) to the top ( Upper Albian) of the Cismon core, particularly consistent in the Hauterivian. Florentinia mantellii (Davey and Williams, 1966) Davey and Verdier, 1973 (122, Plate VIII, 6 and 7). Florentinia radiculata (Davey and Williams, 1966) Davey and Verdier, 1973 (129). Florentinia cf. F. abjuncta Duxbury, 1983 (124, Plate VIII, 8). Remarks: One specimen encountered at 19.95 m displays morphological features consistent with the original description of F. abjuncta (Duxbury, 1983), except for the large antapical process which exhibits the lateral bifurcation of a slender element very similar in size and shape to the other processes.

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PLATE XII

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PLATE XIII

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PLATE XIV

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PLATE XII 1. Odontochitina operculata ×500, sample PAT 4-ox, E.F.C. E 64 2. Phoberocysta neocomica ×600, sample PAT 112-ox, E.F.C. G 28 3. Odontochitina ancala ×550, sample PAT 73, E.F.C. J 57/3 4. Odontochitina spp., operculum ×500, sample PAT 23-ox, E.F.C. R 54/4 5. Phoberocysta neocomica ×550, sample PAT 111-ox, E.F.C. J 25/1 6. Muderongia pariata ×600, sample PAT 25-ox, E.F.C. N 23 7. Dingodinium cerviculum ×500, sample PAT 114-ox, E.F.C. ×55/1 8. Muderongia pariata ×550, sample PAT 71-ox, E.F.C. H 55/4

PLATE XIII Pinocchiodinium erbae Torricelli, gen. et sp. nov. 1. ×550, sample PAT 10-ox, E.F.C. H 61/3 2. ×600, sample PAT 16-ox, E.F.C. V 46/3. Circular operculum inside the cyst 3. ×600, sample PAT 14-ox, E.F.C. V 56. Circular operculum inside the cyst. Paratype 4. ×600, sample PAT 13, E.F.C. F 22/4 5. ×600, sample PAT 12-ox, E.F.C. K 54/3. Paratype 6. ×600, sample PAT 13-ox, E.F.C. R 23/1 7. ×600, sample PAT 14-ox, E.F.C. ×31/2. Holotype 8. ×550, sample PAT 12-ox, E.F.C. U 24/3 9. ×550, sample PAT 12-ox, E.F.C. U 49/2 10. ×550, sample PAT 11-ox, E.F.C. S 49/2. 11. ×550, sample PAT 8-ox, E.F.C. H 30/3. Circular operculum inside the cyst 12. ×550, sample PAT 7-ox, E.F.C. G 39/4

Stratigraphic occurrence: Within the Selli Level, Lower Aptian of the Cismon core. Fromea amphora Cookson and Eisenack, 1958 (120, Plate XXIII, 8). Gardodinium trabeculosum (Gocht, 1959) Alberti, 1961 (56, Plate XV, 8 and 9). Remarks: The specimens of G. trabeculosum recorded here consist only of the internal body of the original cysts, probably due to the breaking of the thin periphragm which characterises this holocavate taxon. Stratigraphic occurrence: Upper Hauterivian to Upper Barremian of the Cismon core. Gochteodinia villosa subsp. multifurcata Davey, 1982 (39, Plate IV, 6). Gonyaulacysta spp. (28). Gonyaulacysta teichos Davey, 1974 (100, Plate XVIII, 9). cf. Gonyaulacysta polythyris Davey, 1979 (125, Plate XV, 6). Remarks: The few specimens herein recorded as cf. G. polythyris differ from the type material of G. polythyris described by Davey (1979) from the Bay of Biscay in lacking strongly developed parasutural crests. By contrast, the other cyst features, such as dimensions, the presence of a subconical apical horn exhibiting elongate perforations and the smooth cyst wall bearing stout intratabular spines, are very similar to those of the holotype. Stratigraphic occurrence: This form was recorded only within the Lower Aptian Selli Level of the Cismon core.

PLATE XIV 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11.

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Callaiosphaeridium asymmetricum ×750, sample PAT BIS 1, E.F.C. D 52/4 Scriniodinium attadalense ×600, sample PAT 85-ox, E.F.C. V 34/4 Ctenidodinium elegantulum ×550, sample PAT 23-ox, E.F.C. D 55 Avellodinium cf. A. falsificum ×600, sample PAT 109-ox, E.F.C. T 26 Avellodinium cf. A. falsificum ×500, sample PAT 95-ox, E.F.C. O 22 Avellodinium falsificum ×550, sample PAT 117-ox, E.F.C. K 29/1 Heslertonia heslertonensis ×750, sample PAT 118-ox, E.F.C. C 59/2 Hapsocysta peridictya ×600, sample PAT 115-ox, E.F.C. R 65/1 Systematophora silybum ×750, sample PAT 40-ox, E.F.C. F 51/1 Taleisphaera hydra ×750, sample PAT 118-ox, E.F.C. O 51/3 Hapsocysta peridictya ×550, sample PAT 121-ox, E.F.C. P 51/3

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PLATE XV

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PLATE XVI

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Hapsocysta peridictya ( Eisenack and Cookson, 1960) Davey, 1979 (30, Plate XIV, 8 and 11). Heslertonia heslertonensis (Neale and Sarjeant, 1962) Sarjeant, 1966 (36, Plate XIV, 7). Heterosphaeridium spp. (79). Heterosphaeridium araneosum (Brideaux, 1977) Islam, 1993 (75, Plate IX, 9). Hystrichodinium furcatum Alberti, 1961 (86). Hystrichodinium pulchrum Deflandre, 1935 (27, Plate XXII, 4). Hystrichodinium ramoides Alberti, 1961 (40, Plate XXII, 5 and 6). Hystrichosphaeridium atlasiense Below, 1982 (134, Plate VIII, 1–3, 9). Hystrichostrogylon stolidotum (Duxbury, 1980) Stover and Williams, 1987 (87, Plate XX, 11 and 12). Remarks: Specimens of H. stolidotum recovered from the Cismon core display a greatly reduced pericoel in comparison to the type material documented by Duxbury (1980) from Speeton, England.

Stratigraphic occurrence: From the uppermost Hauterivian to the Upper Barremian interval of the Cismon core; particularly consistent within the Lower Barremian. 1Kallosphaeridium dolomiticum sp. nov. (16, Plate IV, 9 and 12). Kallosphaeridium ?helbyi subsp. psilatum (Burger, 1980) Lentin and Williams, 1989 (135, Plate I, 9). Kiokansium unituberculatum ( Tasch, 1964) Stover and Evitt, 1978 (104). Kleithriasphaeridium corrugatum Davey, 1974 (37, Plate XXI, 1 and 6). Kleithriasphaeridium eoinodes ( Eisenack, 1958) Davey, 1974 (55, Plate XXI, 5). Kleithriasphaeridium fasciatum (Davey and Williams, 1966) Davey, 1974 (41, Plate XXI, 2). Kleithriasphaeridium cf. K. fasciatum (Davey and Williams, 1966) Davey, 1974 (98, Plate XXII, 1–3). Remarks: This form differs from Kleithriasphaeridium fasciatum in possessing tubiform processes connected by thin, membranous crests. These crests may be low or high, in the latter

PLATE XV 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

Cribroperidinium magnum ×550, sample PAT 63-ox, E.F.C. R 54 Tehamadinium coummia ×550, sample PAT 113-ox, E.F.C. G 64 Cerebrocysta sp. ×500, sample PAT 87-ox, E.F.C. E 50/3 Pyxidinopsis sp. A ×600, sample PAT 97-ox, E.F.C. M 50 Aprobolocysta neista ×550, sample PAT 116-ox, E.F.C. K 28/2 cf. Gonyaulacysta polythyris ×550, sample PAT 19-ox, E.F.C. E 30 Coronifera oceanica ×600, sample PAT BIS 5-ox, E.F.C. B 39 Gardodinium trabeculosum, internal body, ×750, sample PAT 115-ox, E.F.C. L 62 Gardodinium trabeculosum, internal body, ×600, sample PAT 112-ox, E.F.C. P 57/1 Dingodinium cerviculum ×750, sample PAT 106, E.F.C. K 26 Pareodinia sp I Davey 1982 ×750, sample PAT 115-ox, E.F.C. S 41 Pareodinia sp I Davey, 1982 ×550, sample PAT 116-ox, E.F.C. H 35/3

PLATE XVI 1. 2. 3. 4. 5. 6. 7. 8.

Nexosispinum vetusculum ×550, sample PAT 75-ox, E.F.C. O 31 Nexosispinum hesperus subsp. brevispinosum ×550, sample PAT 124-ox, E.F.C. J 26 Nexosispinum hesperus subsp. hesperus ×550, sample PAT 116-ox, E.F.C. R 30 Nexosispinum hesperus subsp. brevispinosum ×1500, sample PAT 112-ox, E.F.C. W 54. Holotype Nexosispinum hesperus subsp. brevispinosum ×600, sample PAT 112-ox, E.F.C. W 54. Holotype Nexosispinum hesperus subsp. brevispinosum ×600, sample PAT 116-ox, E.F.C. K 29/2 Occisucysta ?echinata ×750, sample PAT BIS 1, E.F.C. Q 32/3 Tehamadinium sousense ×550, sample PAT 19-ox, E.F.C. E 28

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case resulting in an entire encircling flange. Some specimens may superficially resemble the dinoflagellate cyst genus Pterodinium Eisenck 1958, although the striate central body and tubiform processes are distinctive of K. cf. fasciatum. Kleithriasphaeridium corrugatum never develops interconnecting crests between processes. Stratigraphic occurrence: This taxon was abundantly recovered from a narrow interval of the Cismon core straddling the Lower–Upper Barremian boundary, from 59.50 to 60.90 m. Forms similar to K. cf. fasciatum as described here were found in the Upper Hauterivian of SE Spain (Leereveld, 1995, 1997). Leptodinium millioudii (Sarjeant, 1963) Sarjeant, 1969 (8, Plate XVIII, 4–6). Lithodinia amlasis (Below, 1981) Williams et al., 1993 (66, Plate III, 4). Lithodinia stoveri (Millioud, 1969) Gocht, 1976 (38, Plate III, 1–3). Litosphaeridium arundum ( Eisenack and Cookson, 1960) Davey, 1979 (145, Plate VIII, 4 and 5). Remarks: Two specimens of L. arundum were recorded in the present study, one with truncated processes and the other with processes displaying slightly flared tips as specimens from the type material described by Eisenack and Cookson (1960) from the Albian of south Australia. Both forms are consistent with the emended description of L. arundum provided by Lucas-Clark (1984, p.188), who states that the periphragm ornamentation is granular and ‘‘the processes are cylindrical to gently tapering, distally truncated or slightly flared…’’. Stratigraphic occurrence: Rare at 2.10 m in the Upper Albian of the Cismon core. Litosphaeridium conispinum Davey and Verdier, 1973 (146). Mendicodinium sp. (127, Plate II, 1). Muderongia spp. (26). ˚ rhus et al., 1990 (52, Muderongia aequicornis A Plate XI, 7). Muderongia pariata Duxbury, 1983 (62, Plate XII, 6 and 8). Muderongia perforata Alberti, 1961 (91, Plate XI, 2).

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Muderongia siciliana Torricelli, 1997 (88, Plate XI, 6). Muderongia simplex Alberti, 1961 (68). Muderongia staurota Sarjeant, 1966 (75a, Plate XI, 1). Muderongia tabulata sensu Monteil (Raynaud, 1978) Monteil, 1991 (18, Plate XI, 3–5). Muderongia testudinaria Burger, 1980 (80, Plate XI, 8). Nannoceratopsis spp. (19, Plate XXIII, 4, 7 and 10–12). *Nexosispinum hesperus Davey, 1979 subsp. hesperus (Autonym) (77, Plate XVI, 3). *Nexosispinum hesperus Davey, 1979 subsp. brevispinosum subsp. nov. (49, Plate XVI, 2 and 4–6). Nexosipinum vetusculum (Davey, 1974) Davey, 1979 (109, Plate XVI, 1). Occisucysta duxburyi Jan du Cheˆne et al., 1986 (131, Plate XIX, 6). Occisucysta ?echinata Duxbury, 1983 (147, Plate XVI, 7). Odontochitina operculata ( Wetzel, 1933) Deflandre and Cookson, 1955 (108, Plate XII, 1). Odontochitina ancala Bint, 1986 (110, Plate XII, 3). Oligosphaeridium albertense (Pocock, 1962) Davey and Williams, 1969 (92, Plate VII, 5). Oligosphaeridium ?asterigerum (Gocht, 1959) Davey and Williams, 1969 (59, Plate VI, 1 and 3). Oligosphaeridium complex ( White, 1842) Davey and Williams, 1966 (46, Plate VII, 3). Oligosphaeridium cf. O. complex (20, Plate VI, 2 and 6). Oligosphaeridium dividuum Williams, 1978 (63). Oligosphaeridium poculum Jain, 1977 (50, Plate VII, 7). Oligosphaeridium porosum Lentin and Williams, 1981 (95, Plate VII, 1 and 6). Oligosphaeridium pulcherrimum (Deflandre and Cookson, 1955) Davey and Williams, 1966 (31). O. totum subsp. totum (Brideaux, 1971) Lentin and Williams, 1973 (9, Plate VII, 4). O. totum subsp. minus (Brideaux, 1971) Lentin and Williams, 1973 (105, Plate VII, 2). Ovoidinium diversum Davey, 1979 (136, Plate I, 10 and 11). Ovoidinium incomptum Duxbury, 1983 (140, Plate I, 12).

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PLATE XVII

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PLATE XVIII

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PLATE XIX

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Palaeoperidinium cretaceum Pocock, 1962 (118, Plate XXIII, 1). Palaeoperidinium? sp. A (143, Plate XXIII, 2). Remarks: Cyst proximate, composed of two layers, delphicavate to circumcavate. The endocyst has more or less the same outline as that of the pericyst. The pericyst is slightly longitudinally elongate, with a subpentagonal outline resulting from well developed apical (one axial ) and antapical (two non-axial ) horns.

PLATE XVII 1. Cribroperidinum boreas ×600, sample PAT 23-ox, E.F.C. M 35/3 2. Cribroperidinum spp. ×500, sample PAT BIS 1, E.F.C. D 52 3. Cribroperidinum spp. ×550, sample PAT 12-ox, E.F.C. P 27 4. Cribroperidinum spp. ×600, sample PAT 112-ox, E.F.C. R 23/2 5. Apteodinium sp. A ×550, sample PAT 127-ox, E.F.C. G 32/4 6. Apteodinium granulatum ×550, sample PAT 116-ox, E.F.C. L 24/2 7. Apteodinium reticulatum ×600, sample PAT 112-ox, E.F.C. R 49/1

PLATE XVIII 1. Pterodinium premnon ×600, sample PAT 72-ox, E.F.C. V 53 2. Pterodinium premnon ×550, sample PAT 73-ox, E.F.C. J 54/4 3. Pterodinium spp. ×750, sample PAT BIS 5-ox, E.F.C. O 55 4. Leptodinium millioudii ×750, sample PAT 87-ox, E.F.C. V 59/4 5. Leptodinium millioudii ×600, sample PAT 117-ox, E.F.C. K 56/2 6. Leptodinium millioudii ×550, sample PAT 129-ox, E.F.C. E 34 7. Wrevittia helicoidea ×600, sample PAT 72-ox, E.F.C. M 57 8. Wrevittia ?diutina ×500, sample PAT 95-ox, E.F.C. T 37/1 9. Gonyaulacysta teichos ×550, sample PAT 11-ox, E.F.C. M 50/4 10. Stanfordella ?cretacea ×550, sample PAT 124-ox, E.F.C. U 34/1 11. Stanfordella fastigiata ×550, sample PAT 88-ox, E.F.C. M 28 12. Wrevittia cassidata ×600, sample PAT 121-ox, E.F.C. E 29/4

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Periphragm finely ornamented with granules and short spines (1–3 mm). Broad cingulum indicated by transverse, parallel parasutural ridges. Pericyst length=60–65 mm; pericyst width=40– 42 mm. Archeopyle not perfectly discernible, possibly involving intercalary paraplates. Operculum adherent. This species is questionably palced in the genus Palaeoperidinium due to the undefined nature of the archeopyle. Stratigraphic occurrence: Upper Albian of the Cismon core. Pareodinia sp I Davey, 1982 (76, Plate XV, 11 and 12). Pervosphaeridium pseudhystrichodinium (Deflandre, 1937) Yun, 1981 (115). Phoberocysta neocomica (Gocht, 1957) Millioud, 1969 (45, Plate XII, 2 and 5). Remarks: When no clearly developed processes occur on the periphragm, the present study follows the Monteil’s (1991) key for determination of species of Muderongia. By contrast, the specimens displaying a thin periphragm ornamented with processes were assigned to P. neocomica, although in many cases it was extremely difficult to evaluate the nature of lateral horns and of the second antapical horn, if any are developed. *Pinocchiodinium erbae gen. et sp. nov. (117, Plate XIII, 1–12). Prolixosphaeridium conulum Davey, 1969 (137, Plate IV, 1). Prolixosphaeridium parvispinum (Deflandre, 1937) Davey et al., 1966 (107, Plate IV, 2 and 3).

PLATE XIX 1. Rhynchodiniopsis aptiana ×550, sample PAT 71-ox, E.F.C. M 26 2. Rhynchodiniopsis aptiana ×550, sample PAT 71-ox, E.F.C. E 59 3. Rhynchodiniopsis aptiana ×550, polar veiw, sample PAT 97-ox, E.F.C. O 56 4. Rhynchodiniopsis cf. R. cladophora ×600, sample PAT 112-ox, E.F.C. H 44 5. Rhynchodiniopsis aptiana ×600, sample PAT 36-ox, E.F.C. S 52/1 6. Occisucysta duxburyi ×550, sample PAT 6-ox, E.F.C. F 49/3

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PLATE XX

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PLATE XXI

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PLATE XXII

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Protoellipsodinium seghire subsp. medaaure Below, 1981 (69, Plate XXI, 12). Protoellipsodinium touile subsp. mugatae¨ Below, 1981 (70, Plate XXI, 11 and 13). Pseudoceratium eisenackii (Davey, 1969) Bint, 1986 (123, Plate IX, 11 and 12). Pseudoceratium pelliferum Gocht, 1957 (51, Plate X, 6).

Pseudoceratium retusum Brideaux, 1977 (130, Plate X, 1–5). Remarks: The ornamentation of the present material is composed of a network of low crests and relatively short processes covering the entire surface of the cyst, sometimes slightly reduced on the mid-dorsal and mid-ventral areas. The processes are often interconnected basally

PLATE XX 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

Spiniferites spp. ×550, sample PAT 19-ox, E.F.C. G 63/4 Spiniferites spp. ×600, sample PAT 37-ox, E.F.C. E 45/4 Spiniferites spp. ×600, sample PAT 16-ox, E.F.C. R 19/4 Spiniferites spp. ×550, sample PAT 68-ox, E.F.C. F 72/4 Spiniferites spp. ×600, sample PAT BIS 1, E.F.C. P 30 Achomosphaera neptunii ×600, sample PAT 61-ox, E.F.C. M 59 Damassadinium chibane ×550, sample PAT 4-ox, E.F.C. H 64 Rhynchodiniopsis cf. R. cladophora ×750, sample PAT 106-ox, E.F.C. Y 61 Achomosphaera neptunii ×550, sample PAT 116-ox, E.F.C. H 51/3 Rhynchodiniopsis cf. R. cladophora ×750, sample PAT 106-ox, E.F.C. P 26 Hystrichostrogylon stolidotum ×750, sample PAT 97-ox, E.F.C. G 63 Hystrichostrogylon stolidotum ×550, sample PAT 95-ox, E.F.C. E 66

PLATE XXI 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13.

Kleithriasphaeridium corrugatum ×600, sample PAT 128-ox, E.F.C. T 29 Kleithriasphaeridium fasciatum ×550, sample PAT 130-ox, E.F.C. V 65/1 Exochosphaeridium phragmites/muelleri ×600, sample PAT 125-ox, E.F.C. M 18 Exochosphaeridium phragmites/muelleri ×600, sample PAT 117-ox, E.F.C. F 52 Kleithriasphaeridium eoinodes ×550, sample PAT 88-ox, E.F.C. T 26 Kleithriasphaeridium corrugatum ×550, sample PAT 127-ox, E.F.C. V 56/3 Endoscrinium campanula ×600, sample PAT 115-ox, E.F.C. L 60/2 Endoscrinium campanula ×550, sample PAT 116-ox, E.F.C. C 24/3 Chytroeisphaeridia ?scabrata ×550, sample PAT 133-ox, E.F.C. E 28/1 Cithariplana caperata ×550, sample PAT 7-ox, E.F.C. D 22/1 Protoellipsodinium touile ssp. mugatae¨ ×550, sample PAT 116-ox, E.F.C. G 61 Protoellipsodinium seghire ssp. medaaure ×750, sample PAT 118-ox, E.F.C. C 28 Protoellipsodinium touile ssp. mugatae¨ ×750, sample PAT 118-ox, E.F.C. N 58/1

PLATE XXII 1. 2. 3. 4. 5. 6.

257

Kleithriasphaeridium cf. K. fasciatum ×750, sample PAT 87-ox, E.F.C. U 28 Kleithriasphaeridium cf. K. fasciatum ×750, sample PAT 87-ox, E.F.C. T 51/2 Kleithriasphaeridium cf. K. fasciatum ×750, sample PAT 87-ox, E.F.C. K 27 Hystrichodinium pulchrum ×750, sample PAT 7-ox, E.F.C. V 45 Hystrichodinium ramoides ×600, sample PAT 112-ox, E.F.C. M 47/3 Hystrichodinium ramoides ×750, sample PAT 106-ox, E.F.C. M 23/3

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PLATE XXIII

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and/or distally joined by trabeculae. An incomplete ectophragm is present, particularly involving the distal linkage of elements at the apices of the horns. Although the ornamentation pattern recorded in the present specimens is constant, a large variability of the cyst shape was observed even within the same assemblage: individuals with reduced antapical and right lateral horns (Plate X, 4 and 5) coexist with individuals characterized by long horns and subtriangular outline (Plate X, 2 and 3). A prominent apical horn is always present (Plate X, 1 and 5). Some of the specimens herein assigned to P. retusum are slightly larger and possess longer lateral and antapical horns than specimens from the type material documented by Brideaux (1977) from Arctic Canada; however, the ornamentation pattern is very similar. Stratigraphic occurrence: This species was recorded from a narrow interval extending from 14.90 to 15.30 m, in the Upper Aptian of the Cismon core. Pterodinium spp. (102, Plate XVIII, 3). Pterodinium premnon Duxbury, 1980 (101, Plate XVIII, 1 and 2). Pyxidinopsis sp. A (94, Plate XV, 4). Remarks: cyst proximate, with a subspherical outline, characterised by a relatively thin autophragm which surface is densely covered with small granules, causing a microreticulate appearance. Evidence of cingulum or sulcus lacking. Archeopyle large and precingular, formed by the loss of a single plate (type P).

Operculum simple, free or adherent. Overall diameter=40–48 mm. Stratigraphic occurrence: Lower Barremian of the Cismon core. Rhynchodiniopsis aptiana Deflandre 1935 (78, Plate XIX, 1–3 and 5). Remarks: Specimens of Rhynchodiniopsis were abundantly recovered from several samples. Most of them are characterised by elevated parasutural crests densely ornamented with spines (Plate XIX, 1–3) and must be assigned to R. aptiana. However, for some specimens occurring in the same assemblages and herein retained as R. aptiana (Plate XIX, 5), the distinction between ‘‘elevated parasutural crests densely ornamented with long spines’’ and ‘‘low crests with well spaced spines’’ which are the diagnostic features of respectively R. aptiana and Rhynchodiniopsis fimbriata according to Jan du Cheˆne et al. (1986), could not be unambigously made due to the intergrading nature of these diagnostic features and/or to the poor preservation of some specimens. Stratigraphic occurrence: In this study R. aptiana was found from the Upper Hauterivian to the lowermost Aptian. Its occurrence is particularly consistent in the Barremian. Rhynchodiniopsis cladophora (Deflandre, 1938) Below, 1981 (82). Rhynchodiniopsis cf. R. cladophora (Deflandre, 1938) Below, 1981 (53, Plates XIX, 4 and XX, 8 and 10).

PLATE XXIII 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

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Palaeoperidinium cretaceum ×550, sample PAT 8-ox, E.F.C. H 65 Palaeoperidinium? sp A ×750, sample PAT BIS 5-ox, E.F.C. T 63/3 Subtilisphaera sp A ×750, sample PAT BIS 5-ox, E.F.C. P 56/3 Jurassic reworking: Nannoceratopsis cf. N. plegas ×600, sample PAT 131-ox, E.F.C. V 20/1 Subtilisphaera perlucida ×750, sample PAT 87-ox, E.F.C. M 51/4 Subtilisphaera sp A ×750, sample PAT BIS 5-ox, E.F.C. M 61 Jurassic reworking: Nannoceratopsis gracilis ×750, sample PAT 132-ox, E.F.C. R 35/4 Fromea amphora ×550, sample PAT 8-ox, E.F.C. M 30/4 Wallodinium luna ×600, sample PAT 63, E.F.C. P 57 Jurassic reworking: Nannoceratopsis deflandrei ssp. deflandrei ×600, sample PAT 65-ox, E.F.C. T 34/2 Jurassic reworking: Nannoceratopsis deflandrei ssp. senex ×550, sample PAT 110-ox, E.F.C. T 24/3 Jurassic reworking: Nannoceratopsis deflandrei ssp. deflandrei ×550, sample PAT 113-ox, E.F.C. J 25

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Scriniodinium attadalense (Cookson and Eisenack, 1958) Eisenack, 1967 (106, Plate XIV, 2). Sentusidinium spp. (42, Plate II, 11–13). Spiniferites spp. (21, Plate XX, 1–5). Remarks: In this study no attempt has been made to distinguish between the species of Spiniferites. Stanfordella ?cretacea (Neale and Sarjeant, 1962) Helenes and Lucas-Clark, 1997 (58, Plate XVIII, 10). Stanfordella fastigiata (Duxbury, 1977) Helenes and Lucas-Clark, 1997 (57, Plate XVIII, 11). Subtilisphaera perlucida (Alberti, 1961) Jain and Millepied, 1973 (99, Plate XXIII, 5). Subtilisphaera sp. A (144, Plate XXIII, 3 and 6). Remarks: Cyst camocavate to delphicavate, with an outline of the periphragm longitudinally elongate and ranging from ovoidal to subpentagonal. An apical horn and a non-axial antapical horn are present, defining rounded subconical projections. Periphragm thin and finely scabrate, smooth in the cingular region, with no evidence of cingular folds/ridges. Archeopyle not discernible. Pericyst length=68–71 mm; pericyst width= 40–43 mm. Subtilisphaera sp. A resembles in many aspects some species belonging to the dinoflagellate cyst genera Isabelidinium Lentin and Williams 1977 and Alterbidinium Lentin and Williams 1985 which are otherwise characterised by the presence of an intercalary archeopyle. Futhermore, species of Alterbidinium are always circumcavate, exhibit a paracingulum and an endocyst with the same outline as that of the pericyst. Stratigraphic occurrence: Upper Albian of the Cismon core. Surculosphaeridium spp. (47). Systematophora spp. (61, Plate IX, 1–3). Systematophora ?daveyi Riding and Thomas, 1988 (60, Plate IX, 6). Systematophora silybum Davey, 1979 (116, Plate XIV, 9). Taleisphaera hydra Duxbury, 1979 (71, Plate XIV, 10). Tanyosphaeridium spp. (22, Plates IV, 4 and V, 11). *Tanyosphaeridium magneticum Davies, 1983; emend. (32).

Tehamadinium coummia (Below, 1981) Jan du Cheˆne et al., 1986 (83, Plate XV, 2). Tehamadinium sousense (Below, 1981) Jan du Cheˆne et al., 1986 (126, Plate XVI, 8). cf. Tenua hystrix Eisenack, 1958, (65, Plate III, 7–9). Remarks: Forms referred to as cf. Tenua hystrix combine morphological features of both the Circulodinium brevispinosum/distinctum group and Tenua hystrix s.s. (Sarjeant, 1985), showing a reduction of processes on mid-dorsal and midventral areas and a more or less vague penitabular arrangement of processes which only locally is clearly discernable, particularly along the archeopyle margin. No unequivocal specimens of Tenua hystrix were recorded, although subjective criteria could have influenced this result. Stratigraphic occurrence: Recorded from the Upper Hauterivian to the lowermost Upper Aptian of the Cismon core with a gap in the Upper Barremian. Leereveld (1995) observed that the stratigraphical evolution from the C. brevispinosum/distinctum type of ornamentation towards the penitabular alignment of processes in Cerbia tabulata Below 1981 [taxonomic junior synonym of T. hystrix following Williams et al. (1998)] seems to be gradual via intermediate forms. After an examination of Hauterivian– Barremian material from England, Germany, Portugal, Spain and France, Leereveld (1995) concluded that the pre-mid Barremian records of C. tabulata in the literature probably are cf. T. hystrix as described here. In this study no significant differences were recorded between the pre-Upper Barremian and the post-Upper Barremian specimens. Valensiella tazadensis (Below, 1981) Lentin and Williams, 1993 (112, Plate II, 10). Wallodinium luna (Cookson and Eisenack, 1960) Lentin and Williams, 1973 (114, Plate XXIII, 9). Wallodinium krutzschii (Alberti, 1961) Habib, 1972 (54). Wrevittia cassidata (Eisenack and Cookson, 1960) Helenes and Lucas-Clark, 1997 (28, Plate XVIII, 12). Wrevittia ?diutina (Duxbury, 1977) Helenes and Lucas-Clark, 1997 (96, Plate XVIII, 8). Wrevittia helicoidea ( Eisenack and Cookson, 1960)

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Helenes and Plate XVIII, 7).

Lucas-Clark,

1997

(111,

Appendix B: Systematic palynology The systematics follow Fensome et al. (1993). The generic allocation of taxa described prior to 1998 follows Williams et al. (1998). Division DINOFLAGELLATA (Bu¨tschli, 1885) Fensome et al. (1993). Subdivision DINOKARYOTA Fensome et al. (1993). Class DINOPHYCEAE Pascher, 1914 Subclass PERIDINIPHYCIDAE Fensome et al. (1993). Order GONYAULACALES Taylor, 1980 Genus Bourkidinium Morgan, 1975; emend. NøhrHansen, 1993

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processes at the apex are generally concentrated on the free, composite operculum, but up to 6 processes may occur on the uppermost part of the precingular plate series leaving a wide equatorial zone devoided of processes’’. Nevertheless, in some specimens encountered in this study, whose main body features and process morphology are identical to the other specimens of B. granulatum, more than six precingular processes (up to 12) were observed (Plate V, 1, 2, 5 and 7). Therefore, NøhrHansen’s (1993) emendation is here slightly modificated to include also these forms in B. granulatum. Comparisons: B. granulatum is very similar to Tanyosphaeridium magneticum in size and shape but completely lacks cingular processes. Stratigraphic occurrence: This species is common and regularly recorded throughout the Upper Hauterivian portion of the Cismon core. Its extinction approximates the Hauterivian–Barremian boundary. Genus Kallosphaeridium de Coninck, 1969.

Bourkidinium granulatum Morgan, 1975; emend. (Plate V, 1–9) Emended diagnosis: Chorate cysts with an elongate ellipsoidal body which bears long, hollow, distally expanded processes arranged in two groups, one in the apical region and the other in the antapical region. The number of processes is variable both at the apex and at the antapex. The processes are cylindrical, slightly tapering toward the tip which is sometimes strongly flared, with serrate or simple margins. The central body is densely granulate and lacks indication of a cingulum or paratabulation. The archeopyle is apical, type tA. The processes at the apex are both concentrated on the free, compound operculum and on the upper part of the precingular plate series, leaving a wide equatorial zone devoided of processes. Discussion: Cysts belonging to the genus Bourkidinium Morgan 1975 were abundantly recovered from the lower portion of the Cismon core. The overall morphology of most of these specimens perfectly corresponds to the emendation of B. granulatum proposed by Nøhr-Hansen (1993, pp. 47–50) which extends Morgan’s original description (1975, pp. 160–161) stating that ‘‘the

Kallosphaeridium dolomiticum Torricelli sp. nov. (Plate IV, 9 and 12) Holotype: slide PAT 128-ox, England Finder Coordinate K 31/4 (Plate IV, 9). Type locality: Cismon borehole, Belluno Province, Veneto, Italy. Type stratum: Biancone Formation. Repository: Palynological slide collection, Stratigraphic Department (STIG) of ENI S.p.A., Agip Division, S. Donato Milanese, Italy. Etymology: Named after the Dolomites, a mountain region in the immediate vicinity of the type locality. Diagnosis: A species of Kallosphaeridium possessing hollow spines distally expanded and closed. Description: Cyst subspherical, proximate, with a smooth, thin autophragm which bears numerous nontabular spines. Spines are smooth and hollow, 3–5 mm in length, gently tapering and distally capitate. Their basal diameter is ca. 2 mm. The archeopyle is apical, type tA. The operculum is attached. Accessory splits between precingular paraplates are present and usually long. Other evidences of paratabulation are absent.

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Dimensions: Holotype: central body length=38 mm (without operculum); width=50 mm; processes length=3–4 mm. Range: central body length=35– 46 mm (without operculum); width=36–52 mm; processes length=3–5 mm. A total of 11 specimens measured. Comparisons: The new species differs from all other species of Kallosphaeridium by the morphology of spines. In particular, Kallosphaeridium nigeriaense Jan du Cheˆne et al. 1984 displays slender acuminate spines. Stratigraphic occurrence: Upper Hauterivian, Lower Cretaceous of the southern Alps.

Emended diagnosis: A species of Nexosispinum possessing a thin, lightly punctate or intraperforate wall. Simple processes are common; others anastomoze in an uncomplicated manner with one or two neightbuoring processes. The anastomozing is generally proximal or medial. The processes are either irregularly arranged over the cyst surface or occur in small clumps or alignments. Distally, they are generally capitate or slightly bifurcate. The archeopyle is precingular, formed by the loss of two plates. Dimensions: Cyst diameter (excluding processes)= 38 (47) 69 mm. Processes length=2–9 mm. Discussion: According to the original description provided by Davey (1979, p. 558), the length of processes in Nexosispinum hesperus ranges from 6 to 9 mm. This species is here emended primarily to include specimens with processes <6 mm (ranging in length between 2 and 4 mm), that are assigned to Nexosispinum hesperus subsp. brevispinosum subsp. nov. Two infra-specific taxa within Nexosispinum hesperus were recognized because, except for the length of processes, all other morphological fatures (wall, shape of processes, dimensions) are identical.

Type locality: Cismon borehole, Belluno Province, Veneto, Italy. Type stratum: Biancone Formation. Repository: Palynological slide collection, Stratigraphic Department (STIG) of ENI S.p.A., Agip Division, S. Donato Milanese, Italy. Etymology: Latin ‘brevis’, short, and ‘spinosum’, in reference to the reduced length of processes. Diagnosis: A subspecies of Nexosispinum hesperus possessing very short processes, ranging in length between 2 and 4 mm. Description: Cysts spherical to subspherical, with a thin wall (ca. 0.5 mm) punctate or intraperforate, bearing numerous, short, solid processes which are apparently randomly arranged. The processes are commonly simple, but sometimes anastomose, proximally, medially or distally, with one or two neighbouring processes. Processes tips are acuminate, capitate or briefly bifurcate. The archeopyle is precingular, formed by the loss of two plates (type 2P). Dimensions: Holotype: central body length= 50 mm; width=45 mm; processes length=2–3 mm. Range: central body length=45–50 mm; width= 45–55 mm; processes length=2–4 mm. Five specimens measured. Comparisons: The new subspecies Nexosispinum hesperus brevispinosum differs from N. hesperus subsp. hesperus (Autonym) in possessing very short processes, ranging in length between 2 and 4 mm. Stratigraphic occurrence: Upper Hauterivian, Lower Cretaceous of southern Alps. This taxon was also found in the sediments of the Monte Soro Flysch, Lower Hauterivian of Sicily, Italy ( Torricelli, personal observation). A specimen illustrated by Leereveld (1995, plate VIII, d; 1997 fig. 9d) from the uppermost Lower Hauterivian of SE Spain and indicated as N. vetusculum is very similar, judged from the illustration, to N. hesperus brevispinosum subsp. nov. because of the very short processes. By contrast, N. vetusculum differs from N. hesperus by having longer and more complexly linked processes (Davey, 1979).

Nexosispinum hesperus subsp. brevispinosum Torricelli, subsp. nov. (Plate XVI, 2, 4–6)

Nexosispinum hesperus subsp. hesperus 1979 (Autonym) (Plate XVI, 3)

Holotype: Slide PAT 112-ox, England Finder Coordinate W 54 (Plate XVI, 4 and 5).

Remarks: this subspecies is automatically established by the erection of Nexosispinum hesperus

Genus Nexosispinum Davey, 1979 Nexosispinum hesperus Davey, 1979; emend.

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brevispinosum subsp. nov. It is described and illustrated by Davey (1979, plate 6, 1–5). Genus Tanyosphaeridium Davey and Williams, 1966.

Tanyosphaeridium magneticum Davies, 1983; emend. Emended diagnosis: Chorate cyst with an elongate ellipsoidal body with granulate ornamentation, bearing long, hollow, distally flared processes arranged mainly at the apex and antapex of the cyst. One to three cingular processes are always present. The shape of process tips varies from truncated to trumpet-shaped, and the distal margin of processes from smooth to serrate. The archeopyle is apical, type tA. Discussion: The holotype of Tanyosphaeridium magneticum (Davies, 1983; plate 8, 7) is reported as having two cingular processes, but the original description (p. 25) states that ‘‘the processes are located at the apex and antapex of the cyst, and 0–3 cingular processes are also present’’. Following the emendation of B. granulatum proposed by Nøhr-Hansen (1993) and modified herein, which includes in this taxon also forms with a precingular ‘corona’ of several processes, the specimen of T. magneticum illustrated by Davies (1983, plate 8, 5), which displays a clearly granulate body but lacks cingular processes, must be assigned to B. granulatum. This discussion advocates a restriction of Davies’ (1983) original description to include in T. magneticum only forms with at least one cingular process. Comparisons: B. granulatum is very similar to T. magneticum in size and shape but entirely lacks cingular processes. Remarks: The emendation of both T. magneticum and B. granulatum proposed in this study is an attempt to facilitate recognition. However, it is evident that T. magneticum possesses intermediate morphological features with respect to B. granulatum and other Tanyosphaeridium species, and all these forms possibly intergrade with each other. It is furthermore possible that, in badly preserved assemblages, the breaking of cingular processes

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could transform a cyst of T. magneticum in a B. granulatum. The most correct approach to the taxonomy of these forms still remains the ‘morphon concept’ proposed by Leereveld and Brinkhuis (1988). Stratigraphic occurrence: Rare in the Upper Hauterivian of the Cismon core. Group ACRITARCHA Evitt, 1963

Pinocchiodinium Torricelli, gen. nov. (Plate XIII, 1–12) Type: Pinocchiodinium erbae Torricelli, sp. nov. (Plate XIII, 7). Etymology: Named after the puppet Pinocchio, hero of the popular tale created by the Italian writer Carlo Collodi (1826–1890): the single lateral horn (or process) resembles Pinocchio’s famous nose. Diagnosis: Elongate ellipsoidal cysts with a singlelayered wall, rounded apex and antapex and a more or less circular opening at the apex whose margin does not exhibit the regular arrangement of parasutural splits which characterise dinoflagellate cyst apical archeopyles. The circular operculum is always detached. A single, hollow and distally open horn (or process) arises laterally from the equatorial region. There is not unequivocal evidence of paratabulation. Discussion: This genus differs from all the previously described dinoflagellate cyst and acritarch genera, and in particular from Fromea Cookson and Eisenack 1958, for the presence of the single tubular horn (or process) arising from the equatorial region of the cyst. Although some workers assigned the genus Fromea to the dinoflagellates (Lentin and Williams, 1993), I agree with Duxbury (1980), Fensome et al. (1990), Fensome et al. (1993) and Williams et al. (1998) who consider it to be an acritarch, since it lacks unequivocal morphological indication of dinoflagellate cyst affinity. This argument is valid also for Pinocchiodinium Torricelli gen. nov. However, amongst the dinoflagellate cysts, this genus displays vague morphological affinities with the Ceratioid group.

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Pinocchiodinium erbae Torricelli, sp. nov. (Plate XIII, 1–12) Holotype: Slide PAT 14-ox, England Finder Coordinate X 31/2 (Plate XIII, 7). Paratypes: Slide PAT 12-ox, England Finder Coordinate K 54/3 (Plate XIII, 5). Slide PAT 14-ox, England Finder Coordinate V 56 (Plate XIII, 3). Type locality: Cismon borehole, Belluno Province, Veneto, Italy. Type stratum: Scaglia Variegata. Repository: Palynological collection, Stratigraphic Department (STIG) of ENI S.p.A., Agip Division, S. Donato Milanese, Italy. Etymology: Named after Dr Elisabetta Erba ( University of Milan, Italy) who coordinated the multidisciplinary study of the Cismon core. Diagnosis: A species of Pinocchiodinium with smooth autophragm. Description: Elongate ellipsoidal cyst, with rounded apex and antapex. Sometimes the antapex exhibits an evident asymmetry due to the presence of a slightly prominent non-axial bulge. The singlelayered wall is thin and smooth. One long (about half of the main body width), tubular horn (or process) arises from the equatorial region: it is distally open, sometimes slightly, irregularly expanded (Plate XIII, 4 and 5). Usually the horn is perpendicular to the main cyst body, but in some cases it is turned downwards (Plate XIII, 10). A wide, more or less circular opening is always present at the apex. In well preserved specimens the opening margin shows neither accessory sutures nor the clearly polygonal outline which characterise dinoflagellate cyst apical archeopyles. The operculum, which has been observed inside some cysts (Plate XIII 2–4, 11), is perfectly round and always free. Dimensions: Holotype (without operculum): main body length=58 mm; width=45 mm; horn length= 20 mm; horn width=4 mm. Range (without operculum): main body length=53–84 mm; width=40– 62 mm; horn length=13–25 mm; 14 specimens measured. Stratigraphic occurrence: This taxon was continuously recorded from a layer 1.82 m below the base of the Selli Level (Lower Aptian) to the

Aptian/Albian unconformity. Certain samples yielded >20 specimens per slide. Rare and well preserved specimens of Pinocchiodinium erbae gen. et sp. nov. were observed in the Aptian interval of the Monte Soro Flysch, Sicily, southern Italy ( Torricelli, personal observation).

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