Geological Survey of Denmark and Greenland Bulletin 1, 75-114 75 The Upper Jurassic of Europe: its subdivision and correlation Arnold Zeiss In the last 40 years, the stratigraphy of the Upper Jurassic of Europe has received much atten- tion and considerable revision; much of the impetus behind this endeavour has stemmed from the work of the International Subcommission on Jurassic Stratigraphy. The Upper Jurassic Series consists of three stages, the Oxfordian, Kimmeridgian and Tithonian which are further subdivided into substages, zones and subzones, primarily on the basis of ammonites. Regional variations between the Mediterranean, Submediterranean and Subboreal provinces are discussed and correlation possibilities indicated. The durations of the Oxfordian, Kimmeridgian and Tithonian Stages are reported to have been 5.3, 3.4 and 6.5 Ma, respectively. This review of the present status of Upper Jurassic stratigraphy aids identification of a num- ber of problems of subdivision and definition of Upper Jurassic stages; in particular these include correlation of the base of the Kimmeridgian and the top of the Tithonian between Submediterranean and Subboreal Europe. Although still primarily based on ammonite stratigraphy, subdivision of the Upper Jurassic is increasingly being refined by the incorporation of other fossil groups; these include both megafossils, such as aptychi, belemnites, bivalves, gastropods, brachiopods, echino- derms, corals, sponges and vertebrates, and microfossils such as foraminifera, radiolaria, ciliata, ostracodes, dinoflagellates, calcareous nannofossils, charophyaceae, dasycladaceae, spores and pollen. Important future developments will depend on the detailed integration of these disparate biostratigraphic data and their precise combination with the abundant new data from sequence stratigraphy, utilising the high degree of stratigraphic resolution offered by certain groups of fos- sils. This article also contains some notes on the recent results of magnetostratigraphy and sequence chronostratigraphy. Keywords: Europe, Upper Jurassic, Oxfordian, Kimmeridgian, Tithonian, Volgian, ammonite zonal and subzonal biostratigraphy and correlations, subdivision by non-ammonite fossil groups, chronometric data, magnetostrati- graphy, sequence stratigraphy Institut für Paläontologie der Universität Erlangen-Nürnberg, Loewenichstr. 28, D-91054 Erlangen, Germany. Present address: Albert Schweitzer Strasse 19, D-91080 Uttenreuth, Germany. E-mail: arnold.zeiss@t-online.de Geological Survey of Denmark and Greenland Bulletin 1, 75–114 (2003) © GEUS, 2003 76 Contents Subdivision and definition of stages: status and unsolved problems 78 The Upper Jurassic (Malm) Series 79 Boundaries of the Upper Jurassic Series 79 Lower boundary (Middle–Upper Jurassic Series boundary) 79 Upper boundary (Jurassic–Cretaceous System boundary) 80 Upper Jurassic stages – subdivision and correlation 81 Oxfordian 81 Lower boundary 82 Substages 82 Zones 84 Correlation 85 Chronometric data 85 Kimmeridgian 85 Lower boundary 86 Additional remarks on Lower Kimmeridgian correlation 89 Substages 89 Zones 90 Correlation 91 Chronometric data 91 Tithonian and Volgian 91 Lower boundary 92 Substages 92 Zones and Subzones 93 Correlation 95 Chronometric data 96 Biochronological importance of non-ammonite fossil groups: a review 96 Invertebrate megafossil groups 96 Cephalopods – other than ammonite conchs 96 Aptychi 96 Belemnites 96 Bivalves 97 Gastropods 97 Brachiopods 97 Echinoderms 97 Corals (scleractinians) 97 Sponges 97 Vertebrate megafossils 97 Invertebrate microfossils 98 Foraminifera 98 Radiolaria 98 Ciliata 98 Ostracodes 99 Plant microfossils 99 Dinoflagellata 99 Calcareous nannofossils (coccoliths, nannolith groups) 99 Charophyaceae 100 Dasycladaceae 100 Spores and pollen 100 Magnetostratigraphy 100 Sequence chronostratigraphy 100 Acknowledgements 101 References 101 The term ‘Upper Jurassic’ (‘Oberer Jura’) was intro- duced by von Buch (1839). Arkell (1956) revived this name with only minor changes in its chronostratigraphic content. The term ‘Upper Jurassic’ in the sense of Arkell (1956) was accepted by the First and Second ‘Colloque du Jurassique’ at Luxembourg in 1962 and 1967; only the stage name ‘Purbeckian’ was eliminated, as it was considered to characterise merely a distinct lithofacies. This usage was followed by the five subsequent International Symposia on Jurassic Stratigraphy at Erlangen in 1984, Lisbon in 1987, Poitiers in 1991, Mendoza in 1994 and Vancouver in 1998. Focus on the formal stratigraphic subdivision of the Jurassic, and the Upper Jurassic in particular, is reflected in the series of key meetings since the early 1960’s (Table 1). The term ‘Malm’ was included in the recommenda- tions of the First Luxembourg Colloquium in 1962 as an alternative term for the ‘Upper Jurassic’ (Maubeuge 1964). Although this term, like the term ‘Tithonian’ (see below), is not based on a geographical site, it has been widely used since its introduction by Oppel (1858, 1865). Referring to the Tithonian stage, Arkell (1956, p. 8) wrote: “it is too late to abolish it after a hundred years of continuous use”; this also applies to the term ‘Malm’. It is important to note that both ‘Upper Jurassic’ and ‘Malm’ are chronostratigraphic terms; the latter, in particular, has frequently been used in a lithostrati- graphic sense by some authors. At the First Colloquium in Luxembourg in 1962, a sub- division of the Jurassic System into stages was pro- posed, the basic framework of which has survived to the present day. The stages were defined by their lower and upper ammonite zones. The recommendations of the First Colloquium (Maubeuge 1964) were thus a landmark in the history of international agreements concerning the subdivision of the Jurassic System into series and stages. After a period of discussion following the publication of the resolutions of the Luxembourg Colloquia (Mau- beuge 1964, 1970), these proposals have been accepted 77 Table 1. Key events in Upper Jurassic stratigraphy since 1960 Date Place Event Reference 1962 Luxembourg Colloque du Jurassique à Luxembourg Maubeuge 1964 1965 Sofia VII Congress, Carpatho-Balkan Geological Association CBGA 1965 1967 Luxembourg Colloque du Jurassique à Luxembourg Maubeuge 1970; BRGM 1974 1967 Moscow International Symposium on Upper Jurassic stratigraphy ANSSR 1974 1969 London William Smith Symposium on Jurassic geology 1969 Budapest Colloque du Jurassique Méditerranéen Végh-Neubrandt 1971 1973 Neuchâtel Colloque sur la limite Jurassique–Cretacé, Lyon BRGM 1975 1975 Sofia International Symposium on the Jurassic–Cretaceous boundary in Bulgaria Nikolov & Sapunov 1977 1977 Stuttgart International Field Meeting on the Jurassic System of southern Germany – this meeting Zeiss 1977; Ziegler 1977 initiated the reorganisation of the International Subcommission on Jurassic Stratigraphy 1979 Novosibirsk International Colloquium on the Upper Jurassic and the Jurassic–Cretaceous boundary Saks 1979 1984 Erlangen International Symposium on Jurassic stratigraphy Michelsen & Zeiss 1984 1984 Sümeg Meeting of the Working Group for the Jurassic–Cretaceous boundary Fülöp 1986 1987 Lisbon 2nd International Symposium on Jurassic stratigraphy Rocha & Soares 1988 1987 International Field Meeting on Jurassic–Cretaceous boundary problems at Menner 1990 the Northern Caucasus 1988 Zaragoza 1st Oxfordian Working Group Meeting, Zaragoza – Iberian Chain Meléndez 1990 1990 Basel 2nd Oxfordian Working Group Meeting, Basel and Jura range of northern Switzerland Gygi 1990b 1991 Poitiers 3rd International Symposium of Jurassic stratigraphy Cariou & Hantzpergue 1994 1992 Warsaw Joint Meeting of the Oxfordian and Kimmeridgian Working Groups Atrops et al. 1993a 1993 London W. J. Arkell Symposium of Jurassic geology Taylor 1996 1994 Mendoza 4th International Symposium on Jurassic stratigraphy Riccardi 1996 1994 Lyon 4th Oxfordian and Kimmeridgian Working Groups Meeting, Lyon and Atrops & Meléndez 1994a South-Eastern France Basin 1997 Warsaw Oxfordian (Jurassic) Meeting in Poland Glowniak et al. 1997 1998 Vancouver 5th International Symposium on the Jurassic System Pálfy 1998, Hall & Smith 2000 worldwide, the only exception being that in the former Soviet Union the Callovian has been considered to belong to the Upper Jurassic (see Krymholts et al. 1988), while in the rest of the world the Callovian is included in the Middle Jurassic. However, following a decision by the Interdepartmental Stratigraphic Commitee in 1989, the Callovian is also now considered in Russia to belong to the Middle Jurassic (Zhamojda 1991). According to the most recent estimates, the Late Jurassic had a duration of a little more than 15 million years according to Gradstein et al. (1994, 1995; Ogg 1995), or 12 million years (+5.6/-7.3) according to Pálfy et al. (1998). The data of Gradstein et al. (1995) have been used in Figures 2, 4 and 5 of this paper; on this basis each of the three Upper Jurassic stages has an aver- age length of 5 million years, while zones and subzones have approximate durations of 700 000 and 300 000 years, respectively. Each subzone comprises at least three horizons, each of which has an approximate dura- tion of 100 000 years. Subdivision and definition of stages: status and unsolved problems On the basis of the recommendations of the First Luxembourg Colloquium (Maubeuge 1964), the Upper Jurassic Series was subdivided into four stages for the Boreal and Subboreal regions: Oxfordian, Kimmeridgian (sensu anglico), Portlandian (sensu anglico) and Volgian, and three for the Submediterranean and Mediterranean regions: Oxfordian, Kimmeridgian (sensu gallico equiv- alent to ‘Crussolian’) and Tithonian (equivalent to ‘Danubian’ and ‘Ardescian’). In the following decades, there has been much confusion as the Kimmeridgian and Portlandian Stages have often been used differ- ently in different parts of Europe. In 1990, a formal vote of the International Subcom- mission on Jurassic Stratigraphy (ISJS) led to the deci- sion to use stages with approximately the same vertical age ranges and uniform names in both regions: Kimmeridgian (sensu gallico) and Tithonian (Zeiss 1991a). With regard to the still unresolved correlation problems between the Boreal and Mediterranean provinces, it was agreed that the Volgian can be used as an alternative stage for the Tithonian in Subboreal and Boreal regions (Fig. 1). The main problem which remained to be solved was the definition of the lower boundary of each stage. To date, no stage has a type locality and a defined lower boundary (global stratotype section and point, or GSSP) formally accepted by the International Commission on Stratigraphy (ICS). There are of course a lot of pro- posals, but they have not been validated according to the guidelines and rules of the ICS (Cowie et al. 1986; Remane et al. 1996). The most intractable problems are to find isochro- nous levels in Submediterranean (and Mediterranean) and Subboreal (and Boreal) Europe for the lower bound- ary of the Kimmeridgian Stage and for the upper bound- ary of the Tithonian (Volgian) Stage. As the former is of particular significance for Upper Jurassic subdivision and correlation, it will be treated here in some detail (see below). Other problems are the unification of the differing subdivisions of stages into substages, correlation of the zones of each stage between the different areas of Europe and the development of better correlation charts from the Boreal regions to the Mediterranean areas. Provisional correlation charts on a zonal and subzonal level for each stage of the European Upper Jurassic are presented here (see Figs 2–5). Zones and subzones are used here as chronozones following the International Stratigraphic Guide (Salvador 1994); originally, many of them were defined as biozones whereas others were used as standard zones, standard chronozones or biochronological standard zones, i.e. only the base is defined while the top is defined by the base of the next overlying unit (Callomon 1965, 1984a, 1994). Problems arise, however, due to inconsistent usage of the term ‘standard zone’. In Northwest Europe, standard zones are mostly used following the concept of Callomon (1994), whereas in central and southern Europe, stan- dard zones are often synonymous with biostratigraphic zones for use in biochronology (Cariou & Hantzperque 1997). It is often difficult, therefore, to determine in which meaning ‘standard zones’ are used. The prob- lems encountered in moving from biostratigraphic field data to biochronological interpretations have been dis- cussed recently by Remane (1991). The term biochronological zone is now used by many authors instead of chronostratigraphic zone, if the zone is based on fossil data. As the ultimate sub- division of biochronology, French authors use the term ‘biohorizon’ (e.g. Enay 1997); their concept is therefore ‘sensiblement different’ from the pure biostratigraphic horizon concept of J.H. Callomon (Dommergues 1997). No attempt has been made here to correlate ammonite faunal horizons due to the variable nature of the pub- lished research on the Upper Jurassic in the various sedimentary basins of Europe. Where necessary, how- ever, correlations of horizons are discussed in the text. 78 For an example of such horizon correlations, the reader is referred to the work of Callomon (1984c) on the Upper Jurassic of North America (this study also cov- ers the European Amoeboceras subdivision). Although attempts have been made to generalise horizons for the whole ‘domaine tethysien’ and ‘domaine boréal’ (Cariou et al. 1997; Hantzperque et al. 1997), these appear premature and of little practical use, given the present state of knowledge. In Europe, subdivisions down to the level of ammonite faunal horizons have been proposed for sev- eral sedimentary basins. Such studies include that of the Upper Jurassic of East Greenland by Callomon & Birkelund (1980, 1982; Birkelund & Callomon 1985), the Lower Kimmeridgian of southern England by Birkelund et al. (1983), the Kimmeridgian of Spitsbergen by Wierzbowski (1989), the Kimmeridgian of the Barents Sea by Wierzbowski & Smelror (1993), the lowermost Oxfordian of northern France by Vidier et al. (1993), the Upper Oxfordian, Kimmeridgian and Lower Tithonian of western France by Hantzperque (1989), the lowermost Oxfordian of south-east France by Fort- wengler & Marchand (1994b), the Lower Kimmeridgian of south-east France by Atrops (1982), the Upper Oxfordian, Upper Kimmeridgian and Lower Tithonian of south-west Germany by Schweigert (1994, 1995a, b, 1996a, b; Schweigert & Callomon 1997), the Oxfordian– Kimmeridgian of Poland (Matyja & Wierzbowski 1997), the Middle Volgian of central Poland (Kutek 1994) and the Oxfordian of north-east Spain by Cariou et al. (1991a) and Meléndez & Fontana (1993). The ‘faunal horizon’ approach clearly represents a method for increasing precision in correlation and dat- ing in the future, when the data from the various sed- imentary basins reach the necessary standard. It is already proving useful in deciphering the history of basin deposits at a resolution that was hitherto impos- sible; such data, in particular, allow us to date events more precisely and to determine the ‘completeness’ of the sedimentary record i.e. to identify accurately the posi- tion and duration of hiatuses. A prerequisite is, how- ever, that it is possible to reconstruct the complete succession of faunal horizons by correlating individual local successions. In this context, it is worth mentioning that the meth- ods of Jurassic stratigraphy and high-resolution geo- chronology have been discussed in detail by Callomon (1984a, b, 1994, 1995), Page (1995), Corna et al. (1997), Blau (1998) and Blau & Meister (2000); formal aspects were covered by Remane (1996). The Upper Jurassic (Malm) Series (Fig. 1) In this paper, subdivision and correlation of the Upper Jurassic Series have been carried out mainly using ammonites. Other fossil groups are reviewed briefly, however, with request to their biochronologic resolu- tion and correlation potential. Many papers have been published on Upper Jurassic ammonites and their chronostratigraphic resolution (see detailed discussion below). For a broad overview, the reader is referred to the papers of Cariou et al. (1997), Geyssant (1997) and Hantzpergue et al. (1997) for western Europe and the Mediterranean. Other important, partly regional com- pilations and revisions have been published by Sapunov (1979), Donovan et al. (1981), Krymholts et al. (1988), Malinowska et al. (1988), Enay et al. (1994) and Schle- gelmilch (1994). Boundaries of the Upper Jurassic Series Lower boundary (Middle–Upper Jurassic Series boundary) The lower boundary of the Oxfordian Stage is rather well-defined by ammonite zones and subzones and only requires more precise definition with respect to the lowermost faunal horizon, which then would char- acterise the beginning of the lowermost subzone (and zone) of the stage. Furthermore, it appears that the lower boundary is approximately (on a subzonal level) the same in Boreal and Mediterranean areas. Once the type faunal horizon has been chosen, then the prob- lem of the type locality for the boundary will also have been solved. At present, this boundary lies in France between the uppermost horizon of the Quenstedtoceras lamberti Zone of the Upper Callovian Substage (the Cardioceras paucicostatum horizon) and the lower- most horizon of the Q. mariae Zone; this was first named in France after Peltoceratoides elisabethae (Fortwengler & Marchand 1994a), but afterwards was changed to Hecticoceras (Brightia) thuouxense (Fort- wengler & Marchand 1994b, c), a species described only recently (Fortwengler et al. 1997). In Dorset, how- ever, Cardioceras cf. woodhamense and C. woodhamense are found in the lowermost levels of the Q. mariae Zone (Callomon & Cope 1996), whereas in north-west France, C. woodhamense has been collected only in the third horizon of the Q. mariae Zone (Vidier et al. 1993). In south-east France, this horizon is only recog- nised tentatively. These different faunal horizons all lie 79 in the Cardioceras scarburgense Subzone, the lower subzone of the Q. mariae Zone, so that the age differ- ence of these horizons (if any) should not be too large. A vote by the Callovian/Oxfordian Boundary Working Group in 1995 resulted in a preference for a type local- ity in south-east France, with the consequence that the Oxfordian would begin with the H.(B.) thuouxense horizon (see above), but a final decision was not taken (Meléndez 1995; Meléndez et al. 1998). Upper boundary (Jurassic–Cretaceous System boundary) In accordance with the decision of the ISJS (see above), there are two alternative stages for the uppermost part of the Jurassic System: Tithonian and Volgian. As they differ in duration, the boundary may be drawn at two different levels, i.e. there are two variants of the Jurassic–Cretaceous boundary. Accordingly, the mem- bers of the former Jurassic–Cretaceous Boundary Working Group agreed to work provisionally with two bound- aries (Remane 1986; Remane et al. 1986; Zeiss 1986). 1. In Mediterranean and Submediterranean Europe, the boundary is placed between the top of the Tithonian Stage (top of Durangites vulgaris Zone and/or of Calpionellid Zone A) and the base of the Berriasian Stage (base of Berriasella jacobi Zone s.l. (= Ber- riasella jacobi and Pseudosubplanites grandis Subzones or Pseudosubplanites euxinus Zone) and/or base of Calpionellid Zone B). 2. In Subboreal and Boreal Europe, the boundary lies between the Upper Volgian (top of Craspedites nodi- ger or Chetaites chetae Zone) and the Ryazanian or ‘Boreal Berriasian’ (base of Chetaites sibericus, Rjasanites rjasanensis or Runctonia runctoni Zone) (Rawson et al. 1978; Kejsi et al. 1988; Sey & Kalacheva 1993a). 80 Upper Upper Upper Upper Upper UpperUpper Upper Middle Middle Middle Middle Middle Middle Middle Lower Lower Lower Lower Lower Lower LowerLowerLower Abnormis Scythicus Klimovi Autissiodorensis Mutabilis Baylei Glosense Plicatilis Mariae Semiforme Hybonotum Beckeri Acanthicum Platynota Bimammatum Bifurcatus Transversarium Mariae Tithonian Kimmeridgian Oxfordian Tithonian (Volgian) StagesStages SubstagesSubstages Germany France Submediterranean Province Subboreal Province Basal zones of substages Basal zones of substages Kimmeridgian Oxfordian Fig. 1. Subdivision of the Upper Jurassic Series of Europe into stages, substages and zones. Substage usage varies in the literature, dependent on author; those indicated are only examples. In the first case, the type locality should be best selected in south-eastern France, where the Ardescian Substage (Upper Tithonian) and the Berriasian Stage were orig- inally described. Subsequent studies have revealed that the sequences are not complete at the base, however, so that it has been suggested that the best sections illustrating the Jurassic–Cretaceous boundary beds and their fauna are situated in southern Spain (Enay & Geyssant 1975; Tavera 1985; Tavera et al. 1994; Enay et al. 1998a, b). In the second case, the boundary should correspond to the base of the Berriasella boissieri Zone in the Mediterranean area. Thus, the Upper Volgian Substage corresponds to the Lower Berriasian (Zeiss 1974, 1979, 1983, 1986; Rawson et al. 1978; Hoedemaker 1990; Sey & Kalacheva 1993a; W.A. Wimbledon in: Callomon & Cope 1996), and is not equivalent to the Upper Tithonian as Mesezhnikov (1988) and other authors have assumed. In a recent review of the Berriasian Stage, Hoede- maker (1994) stated that the Jurassic–Cretaceous bound- ary is typically placed at one of two different levels, either at the base or at the top of the Jacobi Chronozone: “Investigators of Jurassic stratigraphy prefer the lower of these two boundaries, investigators of the Cretaceous stratigraphy the upper” (Hoedemaker 1994, p. 12). At the same time, there has also been an attempt to trace the Jurassic–Cretaceous boundary based on geo- magnetic anomalies from the Tethys to southern England (Ogg et al. 1994). In the Tethyan–Atlantic faunal realm, the top of magnetic polarity reversal M19r approximately coincides with the Tithonian–Berriasian boundary in the Mediterranean area. This reversal is difficult to place pre- cisely in England, but it seems to be situated in the low- ermost Purbeck beds. If so, it would demonstrate once again that the ‘Upper Volgian’ (Casey 1973) or ‘Upper Portlandian’ of England (Wimbledon 1980), i.e. the zonal sequence Subcraspedites primitivus – Subcraspedites lam- plughi, overlaps with the Lower Berriasian. Wimbledon (1980) also included the ‘Upper Volgian’ zones of Casey (1973) in the ‘Portlandian’ of Britain, thus extending the stage upwards by three further zones (termed here ‘Upper Portlandian’). In a recent compi- lation chart, W.A. Wimbledon (in: Callomon & Cope 1996) correlated these ‘Upper Portlandian’ zones and the Upper Volgian zones of the Russian platform with parts of the Lower Berriasian. In Poland, the Jurassic–Cretaceous boundary has been traced by joint studies of ostracodes and ammonites (Marek et al. 1989) whereby the Upper Tithonian and the lower part of the Lower Berriasian could be recog- nised as well as the Upper Berriasian (= ‘Ryazanian’). In a recent paper (Marek & Shulgina 1996), the am- monites of the Berriasian (Ryazanian) were considered to belong to the interval upper occitanica – lower boissieri Zones. In a recent development, the Interdepartmental Stratigraphic Committee of Russia (ISC) approved the following resolutions of its commissions on the Jurassic and Cretaceous Systems (Rostovtsev & Prozorovskiy 1997, p. 48). “1. To draw the Jurassic–Cretaceous boundary in the Boreal Realm between the middle and upper sub- stages of the Volgian, and not …… as …… earlier adopted in Russia (1978). This boundary mainly cor- responds to the Tithonian/Berriasian boundary in Tethyan realm (Colloque Lyon–Neuchâtel, 1975). Correspondingly, the Lower Volgian in the whole cor- related with the Lower and Middle Tithonian; the Middle Volgian, with the Upper Tithonian; the Upper Volgian, with two lower zones of the Berriasian (Jacobi/Grandis and Occitanica). 2. To transfer the Volgian Stage in its former range to the category of regional stratigraphic units (regional stage). To distinguish as chronostratigraphic units in the boundary part of the Jurassic and Cretaceous scale of Russia only Tithonian and Berriasian.” These resolutions, which were precipitated by the work of Sey & Kalacheva (1993a), confirmed the earlier opin- ions of many authors concerning Upper Jurassic/Lower Cretaceous correlations. It is clear that general consensus has not yet been reached; it is assumed, however, that the present Tithonian–Berriasian boundary is not suitable for global correlation. It may be preferable, therefore, to return to an old proposal: to define the Jurassic–Cretaceous boundary at the base of the B. boissieri Zone, where many guide fossils of different groups are available for correlation. Recent studies in the Caucasus area by Remane (1997) are supportive of this proposal. Upper Jurassic stages – subdivision and correlation Oxfordian (Figs 2, 3) The Colloquium at Luxembourg in 1962 (Maubeuge 1964, p. 85) came to the resolution “...... that it seemed necessary to return to the original sense of this stage 81 [the Oxfordian] as defined by A. d’Orbigny and given precision by W.J. Arkell (1956)”. The ‘base’ was indi- cated to be the ‘Zone of Quenstedtoceras mariae’ and the ‘top’ the ‘Zone of Ringsteadia pseudocordata (= Zone of Idoceras planula), (= Zone of Epipeltoceras bimammatum)’. It was recommended that other stage and substage names then still in use, e.g. the Argovian (Marcou 1848), Rauracien (Greppin 1867), Sequanian (Marcou 1848) and the Lusitanien (Choffat 1885; Haug 1910) should be abandoned. These stages had been interpreted in different ways so that continued usage would have created only more confusion. Subsequent studies (e.g. Enay 1980a; Gygi & Persoz 1986; Enay et al. 1988) demonstrated the validity of this resolution. Lower boundary See discussion above. Substages Although the Oxfordian has been subdivided into three substages, Lower, Middle and Upper Oxfordian, full agreement has not been reached on the zonal content of these substages and the position of their boundaries (Callomon 1988, 1990, fig. 10; Meléndez & Fontana 1993, fig. 5; Wright 1996a). The subdivision is thus essentially informal but the substages are capitalised in 82 SwitzerlandFrance, Spain S. Germany England Greenland, Scotland, Svalbard Galar (Gigantoplex, Grandiplex) Galar Planula Galar Planula (Grandiplex) Planula Proteron Pl an ul a Pl an ul a Luciaeformis (Wartae) Parandieri Parandieri Nunning- tonense Cautisnigrae Serratum Serratum Koldewayense Regulare Rosen- krantzi Variocostatum Caledonica Pseudoyo Parandieri Schilli Schilli Schilli Rotoides Stenocycloides Stenocycloides Grossouvrei Grossouvrei Antecedens Antecedens Antecedens Tenuiserratum Glosense (Alternoides) Glosense Ilovaiskyi Tenuiserratum Blakei Pu m ilu s C au ti sn ig ra e Antecedens Vertebrale Vertebrale Vertebrale Maltonense Vertebrale Cordatum Cordatum Cordatum Cordatum Cordatum Costicardia Costicardia Costicardia Costicardia Costicardia Bukowskii Bukowskii Bukowskii Bukowskii Bukowskii Praecordatum Praecordatum Praecordatum Praecordatum Praecordatum Scarburgense Scarburgense Scarburgense Scarburgense Scarburgense Plicatilis Pl ic at il. Pl ic at ili s C o rd at um C o rd at um C o rd at um CordatumCordatum M ar ia e M ar ia e M ar ia e MariaeMariae A lt er na ns O va le B au hi ni R in gs te ad ia Densiplicatum Densiplicatum Transversarium Transver- sarium Tr an sv er sa ri um Tr an sv er sa ri um Bifurcatus Bifurcatus Hypsel. Hauffianum Bimammatum Bimammatum Bimam-matum Bimammatum B im am m at um Ps eu do co rd at a B ay le i R av ni (S ub -) B o re al O x. /K i. bo un da ry Pseudocordata Evoluta Densicostata Baylei ? ? Bauhini (? Bayi) Bauhini Tonnerense Praecursor Praecursor Berrense Berrense Hauffian. Semimammatum B ifu rc at us Submediterranean Subboreal Boreal 15 9. 4 (± 3 .6 ) 15 4. 1 (± 3 .2 ) U pp er M id dl e Lo w er O xf o rd ia n Fig. 2. A tentative correlation chart for the Oxfordian Stage in Europe (thick lines indicate periods during which correlation is diffi- cult). Modified after Zeiss (1984), Mesezhnikov (1988), Cariou et al. (1991b), Wright (1996a, b), Matyja & Wierzbowski (1997, 1998), Schweigert & Callomon (1997) and Gygi (2000a, b). this paper, following common usage. An example of the ongoing debate is the inclusion of the D. bifurca- tus Zone in the Middle or Upper Oxfordian; this zone was introduced by Enay (1966) as the upper subzone of the G. transversarium Zone but was later considered as the lowermost zone of the Upper Oxfordian (Cariou et al. 1971). Preference is given here to a subdivision in which the D. bifurcatus Zone is included in the Middle Oxfordian (Fig. 1) as has also been proposed by Meléndez (1989), Cariou & Meléndez (1990), Cariou et al. (1991a) and Gygi (2000a) although not followed by Cariou et al. (1991b, 1997). While the lower and middle substages have the same lower boundaries in Submediterranean and Subboreal Europe, the position of the lower boundary of the upper substage differs. In Boreal Europe, it has been drawn at three different levels (Wright 1996a, fig. 6). The solu- tion to draw it at the base of the A. glosense Zone is well-known (Sykes & Callomon 1979; Wright 1980); it would correspond to the base of the P. luciaeformis Subzone in the G. transversarium Zone, i.e. the bound- ary would be drawn around one and a half zones deeper than in the Submediterranean subdivision. It seems preferable to draw the boundary at the lower bound- ary of the A. rosenkrantzi Zone, corresponding approx- imately to the lower boundary of the Upper Oxfordian both in Submediterranean Europe (base of E. bimam- matum Zone) and in Subboreal Europe (base of R. pseudocordata Zone), although the latter lies some- what deeper (Matyja & Wierzbowski 1997, fig. 4). Additional literature references pertinent to the sub- division of the Oxfordian Stage are Enay (1963, 1966), Zeiss (1966), Sequeiros (1974), Sapunov (1976), Gygi (1977, 1986, 1990a, 2000a, b, c), Wierzbowski (1978), Enay & Meléndez (1984), A. Zeiss (in: Enay & Meléndez 1984), Cariou & Meléndez (1990), Malinowska (1991), Meléndez & Fontana (1993), Schweigert (1995a, b), Fözy & Meléndez (1996), Matyja & Wierzbowski (1997, 1998), Groiss et al. (2000) and Schweigert & Callomon (1997) for the Submediterranean and Mediterranean provinces, and Sykes & Callomon (1979), Wright (1980, 1996a, b) and Mesezhnikov (1988) for the Subboreal and Boreal provinces. Mönning & Bertling (1995), Mönning (1998) and Gramann et al. (1997) have pre- sented interesting and useful reviews of the ammonite succession in northern Germany. 83 Submediterranean Standard (Cariou et al. 1991b) Spain – North Africa (Sequeiros 1974; Cariou et al. 1991b) Poland (Tarkowski 1990; Matyja & Glowniak 1994) Bulgaria (Sapunov 1976) Trans- versarium Plicatilis Riazi Antecedens Antecedens Promiscuus Oculatum HelenaePatu- ratten- sis Baccatum Spixi Wartae Antecedens Episcopalis Renggeri Athletoides Riazi Antecedens Cordatum Costicardia Bukowskii Praecordatum Scarburgense Vertebrale (Tenuicostatum) Rotoides Schilli Luciaeformis Parandieri Paturattensis Claro- montanus Claro- montanus Paturattensis Paturattensis Plicatilis Minax Magnouatius MazuricusCordatum Mariae Trans- versarium Fig. 3. A tentative correlation chart for some alternative subdivisions of parts of the Oxfordian Stage in Mediterranean and Submediterranean Europe. Zones The zonal and subzonal subdivision of the Lower Oxfordian Substage was established by Arkell (1941) using Quenstedtoceras mariae and Cardioceras corda- tum as index species; it can be used over large areas of northern and central Europe (Fig. 1) and is also applicable in the Dauphinois basin of south-east France as recently demonstrated by Fortwengler & Marchand (1994a; Fortwengler et al. 1995). In southern Europe, a variety of subdivisions exist; at least three distinct subdivisions testify to the difficulties in erecting a generally accepted zonal scheme if car- dioceratids are missing. In such cases, peltoceratids (Peltomorphites, Peltoceratoides and Parawedekindia), oppeliids (Taramelliceras, Popanites and Creniceras) and perisphinctids (Otosphinctes, Perisphinctes, Prososphinc- tes and Properisphinctes) are important guide fossils (see Fig. 3), e.g. Taramelliceras minax, T. spixi, T. bac- catum, T. oculatum, Popanites paturattensis in Poland (Tarkowski 1990), Peltomorphites athletoides and Creniceras renggeri in Bulgaria (Sapunov 1976), and Prososphinctes mazuricus and P. claromontanus in Spain (Aurell et al. 1990). From the base of the Middle Oxfordian, perisphinctids and peltoceratids become the dominant ammonite groups with respect to index fossils at the substage level in the Submediterranean and Subboreal provinces. The first aulacostephanids (Decipia) also appear at this level. The Perisphinctes plicatilis, Gregoryceras trans- versarium and Dichotomoceras bifurcatum Zones make up the Middle Oxfordian Substage in the Submedi- terranean area, the Perisphinctes plicatilis, P. pumilum and P. cautisnigrae Zones are representative of the Subboreal province. Boreal indexes are Cardioceras tenuicostatum and C. tenuiserratum, Amoeboceras glosense and A. serratum. The correlation between Sub- boreal perisphinctid and amoeboceratid zones was well demonstrated by Wright (1996b). There is a difference in the usage of the P. plicatilis and G. transversarium Zones in Submediterranean Europe. Although Gygi & Marchand (1982) replaced the basal C. vertebrale Subzone with the C. densiplicatum Zone and included the P. antecedens Subzone in the G. transversarium Zone, subsequent authors have not followed the arguments of these authors and have con- tinued to use the P. plicatilis Zone in the sense of Cariou et al. (1991a, b), i.e. with C. vertebrale and P. antecedens Subzones (e.g. Meléndez & Fontana 1993, fig. 4; Cariou et al. 1997). Cariou et al. (1991a) defined the G. trans- versarium Zone to contain the P. parandieri, P. luciae- formis, L. schilli and P. rotoides Subzones. In a more recent publication, Gygi (1995) included in the lower part of the G. transversarium Zone not only the P. antecedens Subzone but also the C. densiplicatum Subzone i.e. the whole P. plicatilis Zone (following the original usage of Oppel & Waagen 1866; R.A. Gygi, personal communication 1997). In further contributions to the Upper Jurassic of Switzerland (Gygi 2000b, c), the G. transversarium Zone is subdivided into the C. densiplicatum, P. antecedens, and P. luciaeformis Subzones; the overlying D. bifurcatus Zone contains in its lower part the L. schilli Subzone, which is consid- ered in Spain and France to represent the upper part of the G. transversarium Zone (see above). The main reason for these differences is the occurrence of L. schilli in Switzerland above the vertical range of G. transversarium. The Upper Oxfordian Substage in Submediterranean Europe comprises the Epipeltoceras bimammatum, Idoceras planula and Sutneria galar Zones. Considering the new correlations of Wright (1996a), Matyja & Wierzbowski (1997) and Cariou et al. (1997), Ringsteadia pseudocordata would be the corresponding index fos- sil for the Subboreal province, whereas Amoeboceras rosenkrantzi would be the index fossil for Boreal Europe. The Amoeboceras serratum Zone of Malinowska (1991) contains Epipeltoceras (uhligi group) and Ringsteadia salfeldi thus indicating, at least partly, equiv- alence with the lower E. bimammatum Zone (E. hypselum Subzone); this demonstrates that the A. ser- ratum Zone of this author is younger in age than the A. serratum Zone of Sykes & Callomon (1979). The A. regulare Subzone of Malinowska (1991) seems to rep- resent the upper E. bimammatum and perhaps the low- ermost I. planula Zones, while the A. lineatum Subzone apparently corresponds to the rest of the I. planula Zone and the S. galar Zone. The most difficult problems associated with these Upper Oxfordian zones concern their correlation in the Subboreal and Submediterranean schemes; this aspect is discussed in detail below. Some minor problems may be caused by the different hierarchical status of zones and subzones in the Subboreal and Boreal provinces. For example, Atrops et al. (1993b) recognised the A. regulare, A. rosenkrantzi and A. bauhini Zones, whereas Malinowska (1991) subdivided the R. pseudocordata Zone into the A. regulare and A. lineatum Subzones or, in Boreal Europe, into the A. regulare and A. rosenkrantzi Subzones. However, comparing the cor- relation chart of Malinowska (1991, table 3) with that of Matyja & Wierzbowski (1997, fig. 3), it becomes evi- 84 dent that the P. pseudocordata Zone of Malinowska corresponds only to the upper part of the A. regulare, the A. rosenkrantzi and the P. baylei Zones. Another example is the variable status of A. bauhini as an index species. There is the A. bauhini horizon in the upper E. bimammatum zone equivalent to the P. densicostata horizon (Schweigert & Callomon 1997), the A. bauhini Subzone of the A. rosenkrantzi Zone (Sykes & Callomon 1979; Cariou et al. 1997), equiva- lent to the P. baylei Zone of Birkelund & Callomon (1985), and the A. bauhini Zone. Although initially equivalent to the P. densicostata horizon (Wierzbowski & Smelror 1993), the A. bauhini Zone was expanded by Matyja & Wierzbowski (1997, 1998) to correlate with the uppermost P. pseudocordata Zone and nearly the whole P. baylei Zone on the one hand and with the whole I. planula Zone and uppermost E. bimamma- tum Zone on the other; a little more restricted was the A. bauhini Zone of Schweigert & Callomon (1997), who excluded the S. galar Subzone of the I. planula Zone (see below). Correlation There have been many proposals and attempts to cor- relate the zonal subdivisions of the Oxfordian of Mediterranean, Submediterranean, Subboreal and Boreal areas of Europe; the most important ones have been already discussed in the text above (see Figs 2, 3). Further informative compilations have been presented by Enay & Meléndez (1984), Mesezhnikov (1988), Cariou et al. (1991a, b; 1997), Malinowska (1991), Aleynikov & Meledina (1993), Meléndez & Fontana (1993), Schweigert (1995b), Wright (1996a, b) and Matyja & Wierzbowski (1997). In short, correlation within the Lower Oxfordian is possible over wide regions of Boreal, Subboreal and Submediterranean Europe, but becomes difficult on approaching the Mediterranean area. At the base of the Middle Oxfordian, ammonites of the Perisphinctes plicatilis Zone provide the last possi- bility for long-distance correlation. Higher up in the Middle Oxfordian, zonal correlations become more and more difficult, best illustrated by the charts of J.H. Callomon (in: Wright 1980), Enay & Meléndez (1984) and Cariou et al. (1991b; see also Fig. 2). The divergent views are also well-documented by the tables of Malinowska (1991), Wright (1996a), Cariou et al. (1997) and Matyja & Wierzbowski (1997). The problems of Upper Oxfordian correlation, con- centrated mainly on the correspondence of the E. bimammatum, I. planula and S. galar Zones to the R. pseudocordata, P. baylei, A. regulare, A. rosenkrantzi and A. bauhini Zones, are under discussion (Wierz- bowski 1991; Atrops et al. 1993b; Atrops & Meléndez 1994b; Schweigert 1995a, b; Cariou et al. 1997; Matyja & Wierzbowski 1997; Schweigert & Callomon 1997). This aspect is especially relevant to the Oxfordian– Kimmeridgian boundary problem and is therefore dis- cussed in more detail below. Chronometric data The duration of the Oxfordian Stage is estimated at 5.3 Ma (Gradstein et al. 1995; Ogg 1995; Ogg & Gutowski 1996); for precise data, see Figure 2. Kimmeridgian (Fig. 4) Following the Luxembourg recommendations of 1962 and 1967 (Maubeuge 1964, 1970), two possibilities existed with respect to usage of the Kimmeridgian Stage, namely either a long version (‘sensu anglico’) or a short version (‘sensu gallico’), both with differing zonal con- tent and boundaries (see below). Use of two different versions of the Kimmeridgian evoked much confusion in following years and led to endless discussion. Therefore a vote of the International Subcommission on Jurassic Stratigraphy (ISJS) on this question was arranged in 1990, simultaneously with the vote on the Tithonian Stage (see below); the members of the ISJS voted for a ‘short’ version of the Kimmeridgian Stage (i.e. ‘sensu gallico’). This meant that in future the upper boundary of the Kimmeridgian Stage should be coin- cident with the lower boundary of the Tithonian Stage and its Boreal equivalent, the Volgian (Zeiss 1991a). The lower boundary of the stage, however, remained ambiguous (see below). Because of the still unresolved problems at the Oxfordian–Kimmeridgian boundary, the lower bound- ary of the Kimmeridgian Stage is drawn in this paper at the base of the Sutneria platynota Zone, following the above-mentioned adoption of a short Kimmeridgian Stage (i.e. ‘sensu gallico’ or according to the ‘conti- nental’ concept; Enay 1980b). The Working Group of the Oxfordian–Kimmeridgian Boundary is mandated to finally define the boundary at a level which allows far- reaching correlations and corresponds to the resolutions of the International Commission on Stratigraphy (ICS); see also the discussions by Wierzbowski (1999, 2001). 85 Until such a definition has been taken by the Oxfordian–Kimmeridgian Boundary Working Group, voted on by the ISJS and approved by ISC, it seems use- ful to maintain the traditional boundaries in both bio- geographic provinces, and it is premature to draw the Oxfordian–Kimmeridgian boundary in the Submedi- terranean area in the upper part of the E. bimamma- tum Zone (cf. Gygi 2000a, b). Lower boundary As the Luxembourg recommendations made it possi- ble to select between two distinct versions of the Kimmeridgian Stage, the lower boundary was also defined twofold. In Subboreal regions of Europe, the boundary was drawn at the lower boundary of the Pictonia baylei Zone, whereas in Submediterranean regions it was placed at the base of the Sutneria platynota Zone (Maubeuge 1964, p. 85–86). At that time, it was supposed that both boundaries were more or less isochronous (Ziegler 1964), although doubts remained (e.g. Zeiss 1965; Cariou et al. 1971). With the publication of Sykes & Callomon (1979), new impetus was given to further studies, which have sug- gested that the assumed time equivalence is erroneous or, at best, only partially true (Matyja & Wierzbowski 1988; Wierzbowski 1991; Atrops et al. 1993b; Schweigert 1995a, b). The main reasons for this view were the dis- covery of new Amoeboceras faunas by these authors 86 Beckeri/ Pressulum Beckeri Eudoxus Eudoxus Mutabilis Cymodoce Cymodoce Achilles Chatelaillon- ensis Cymodoce Baylei Bayi Subkitchini Modestum Ruepellense Mutabilis Lallierianum Orthocera Caletanum Contejeani Autissio- dorensis Irius Mutabilis Autissio- dorensis (Volgensis) Autissio- dorensis (Taimyrense) Elegans Kochi K o ch i N o rv e- gi cu m Autissio- dorensis Eudoxus EudoxusCavouri Acanthicum Acanthicum Divisum Hypselo- cyclum Platynota Polygyratus Desmoides Guilheradense Hippolytense Lothari Crusoliense Uhlandi Balderum Linealis Attenuatus Eulepidus Liparum/ Schilleri Eudoxus Caletanum Subeumela Setatum Ulmense Submediterranean S. Germany Mediterranean N. Italy (S. Alps) Biome Franco-Germanique W. France Subboreal Great Britain Boreal N. Europe Mixed Poland Herbichi Strombecki Silenum Trenerites Raschi Stenonis Divisum Uhlandi Fallax Sub- borealis A ut is si o do re ns is M ut ab ili s C ym o do ce K it ch in i Acanthicum Divisum Hypselo- cyclum Platynota U pp er M id dl e K im m er id gi an Lo w er 15 0. 7 (± 3. 0) 15 4. 1 (± 3. 2) Fig. 4. A tentative correlation chart for the Kimmeridgian Stage in Europe (thick lines as in Fig. 2). Modified after Zeiss (1965), Atrops (1982), Sarti (1988), Hantzpergue et al. (1991), Wierzbowski & Smelror (1993), Kutek & Zeiss (1994), Schweigert & Zeiss (1994) and Matyja & Wierzbowski (1997, 1998). and a re-evaluation of Salfelds (1915) Cardioceras paper as well as that of Koerner (1963), particularly with respect to their remarks concerning the type locality and possible type horizon of Cardioceras (= Amoeboceras) bauhini. The discussion of Wierzbowski (1991) con- cerning the range of the genus Ringsteadia in Poland is also important in this context. It soon became evi- dent that Amoeboceras bauhini has its type horizon just below the upper boundary of the E. bimammatum Zone, (see A. Zeiss in: Enay & Meléndez 1984). The stud- ies of Schweigert (1995a, b; Schweigert & Callomon 1997) resulted in similar conclusions, but led to a more precise faunal horizon subdivision of the Upper Oxfordian in Württemberg, SW Germany and to better correlation possibilities with England, with respect to the A. bauhini and the A. bayi (?= A. subtilicaelatum) horizon. Some problems remained unsolved, however: 1. Does the A. bauhini horizon of southern Germany represent the same time interval as the beds bearing A. bauhini in England, Scotland and the Barents Sea? Or is there a difference, and the vertical range of this species is different in these two areas? What is the situation in Poland, representing an intermediate region? 2. Does the A. subtilicaelatum horizon of southern Germany represent the same time interval as the A. bayi horizon in England? Or is there also a difference in the vertical range of these species in different parts of Europe? 3. Which units in the Subboreal realm correspond to the succession from the base of the I. planula Zone (with three or four faunal horizons) and the top of the lower S. galar Zone, which in Submediterranean Europe occurs between the A. bauhini and the A. subtilicaelatum (?= A. bayi) horizon? It is not easy to answer these questions given the pre- sent state of knowledge; the following points are perti- nent prior to discussion of these problem areas. The usage of A. bauhini as an index ammonite began with its intro- duction by Sykes & Callomon (1979) as a subzone of the A. rosenkrantzi Zone (uppermost Oxfordian); its stratigraphic position was subsequently revised by Birkelund & Callomon (1985), who regarded the A. bauhini Subzone and the P. baylei Zone (Lower Kimmeridgian) as approximate equivalents. One year prior to this latter publication, A. Zeiss (in: Enay & Meléndez 1984, fig. 6) had used A. bauhini informally as a zonal index in a correlation chart to show its approx- imate correspondence with the I. planula Zone sensu lato; this view was also held by Atrops et al. (1993b) and Matyja & Wierzbowski (1997, 1998). Wierzbowski & Smelror (1993) established the A. bauhini Zone formally and suggested that it was equivalent to only the lower part of the P. baylei Zone (the P. densicostata horizon); in more recent papers, Matyja & Wierzbowski (1994, 1995, 1997, 1998) provided charts showing the correlation between the A. bauhini Zone and the I. planula Zone sensu lato as well as with the P. baylei Zone (with the exception of the uppermost part). Finally, in southern Germany, an A. bauhini horizon was described by Schweigert (1995b; Schweigert & Callomon 1997) in the upper part of the T. hauffianum Subzone (uppermost E. bimammatum Zone); the latter authors correlated the Boreal A. bauhini Zone with the I. planula Zone sensu stricto, whereas the S. galar Zone was correlated with the Amoeboceras kitchini Zone. The Amoeboceras bayi horizon was introduced by Birkelund & Callomon (1985) in the upper part of the P. baylei Zone, whereas Wierzbowski & Smelror (1993) reported the species at the base of their A. subkitchini Subzone. Atrops et al. (1993b) found the species, or closely related forms, in the Sutneria platynota Zone of the Submediterranean area. Schweigert (1995b) estab- lished an A. subtilicaelatum horizon in the uppermost part of the Sutneria galar Zone, assuming that A. bayi is only a variant of A. subtilicaelatum, which would then have priority. This conflicts with the opinion of Salfeld (1915), that A. lineatum and A. subtilicaelatum are very close and perhaps synonymous. Schweigert (1995b) also assumed that many specimens determined earlier as ‘A. bauhini’ belong in reality to A. bayi. To verify these assumptions, a comprehensive re-evaluation of the Upper Oxfordian – Lower Kimmeridgian Amoeboce- ras species complex (A. bauhini – A. bayi – A. subtili- caelatum – A. lineatum) would be necessary. Such a study should also illustrate the variation within each species in time and space (see, for example, Klieber 1981; Birkelund & Callomon 1985; Matyja & Wierzbowski 1988, 1994; Schweigert & Callomon 1997). For the time interval of the I. planula Zone, Malinowska (1991) established the A. lineatum Subzone in Poland. It was introduced as the upper subzone of the R. pseudocordata Zone, but the precise correlation with other areas is not clear; from the list of fossils one would conclude that the S. galar Zone is not present. However, as a Sutneria sp. (of the galar/praecursor group?) is mentioned in the text but not figured, a deci- 87 sion is difficult; its low stratigraphic level in the Goldap section would favour the S. praecursor Zone. In addi- tion, Wierzbowski (1978) has described A. lineatum and A. bauhini together from the lower part of the I. planula Zone; thus, the A. lineatum Subzone seems to correspond to the lower part of the P. baylei Zone rather than to the upper part of the R. pseudocordata Zone. Malinowska (1988) reported specimens of A. bauhini only from the Lower Kimmeridgian, but these forms belong to other species such as A. bayi or A. cf. cricki. In the Subboreal province, the Pictonia baylei Zone consists of two or three horizons. The lowermost hori- zon in Great Britain and the Boulonnais area is the Pictonia densicostata horizon; as mentioned above, this probably corresponds to the A. bauhini horizon. In the Boulonnais and Normandy areas, this is followed by the Pictonia baylei horizon sensu stricto, and, more widespread in France, the P. baylei and P. thurmanni horizon. In Dorset, the second horizon is apparently missing (Hantzpergue 1989), while the third one is rep- resented by the P. baylei and P. normandiana hori- zon, which can also be observed in East Greenland (P. aff. normandiana horizon, Birkelund & Callomon 1985). P. normandiana is regarded as a synonym of P. thur- manni by Hantzpergue (1989). This third horizon also contains A. bayi. What conclusions can be made from all these obser- vations? 1. It seems likely that A. bauhini has a longer range in south Germany, as suggested by the many records of this ammonite species from the E. hypselum Subzone of the E. bimammatum Zone to the I. plan- ula and S. galar Zones and even from the S. platynota Zone; a number of these determinations, although probably not all, may however be erroneous (Schweigert 1995b). Data from Poland also demon- strate that the range of A. bauhini is not restricted to the upper T. hauffianum Zone (= A. bauhini hori- zon), but extends as in southern Germany from the upper E. hypselum Subzone of the E. bimammatum Zone to the top of the I. planula Zone sensu lato (Matyja & Wierzbowski 1997, 1998). It is likely, there- fore, that the A. bauhini Zone is of longer duration in the Submediterranean area, because it comprises not only the A. bauhini horizon of the upper T. hauf- fianum Subzone, but also three or four horizons of the I. planula Zone sensu stricto and at least one horizon of the lower S. galar Zone. As mentioned above, Malinowska (1991, p.16–17) apparently intro- duced the term A. lineatum Subzone for such an extended A. bauhini Zone. Approximately the same time interval has been called the A. bauhini Subzone (of an unnamed zone) by Matyja & Wierzbowski (1994, 1995) and subsequently elevated to the A. bauhini Zone (Matyja & Wierzbowski 1997, 1998); this zone is now correlated with the upper E. bimam- matum Zone and the I. planula Zone sensu lato. It should also be noted that there is some evidence, at least in Scotland, that above the P. densicostatum bed follows another, younger bed with A. bauhini and Pictonia sp. (Wright 1989). This could be a hint that there are some more beds with A. bauhini, but without P. densicostata, which could correspond to the higher horizons of the P. baylei Zone. In England, in contrast, Cox & Richardson (1982) observed A. bauhini in the uppermost part of the A. rosenkrantzi (= R. pseudocordata) Zone. If these determinations are correct, A. bauhini may occur a little earlier than the P. densicostata horizon. One can conclude from these observations that the range of A. bauhini, even in the Subboreal regions, is not restricted to the P. densicostata horizon or the ‘A. bauhini Zone’ sensu Wierzbowski & Smelror (1993). 2. If it can be confirmed that Amoeboceras bayi and Amoeboceras subtilicaelatum are synonymous, as assumed by Schweigert (1995b), then the upper hori- zon of the Sutneria galar Zone (A. subtilicaelatum horizon) may correspond to the Amoeboceras bayi horizon of the lowermost Kimmeridgian Amoeboceras kitchini Zone. It should be noted, however, that A. bayi has also been reported from the lower (‘Ortho- sphinctes’) horizon of the S. platynota Zone (Atrops et al. 1993b). 3. (a) It can be concluded from the above that corre- lation of the A. bauhini and P. densicostata horizon with the A. subtilicaelatum and A. bayi horizon is pos- sible, but the vertical ranges of the former species may be longer and the correlation may thus be only partial. Consequently, the position of the upper boundary of the A. bauhini Zone and the lower boundary of the A. bayi horizon require more pre- cise definition. (b) In the sequence between the A. bauhini and the A. subtilicaelatum horizons, equivalent to the mid- dle part of the Pictonia baylei Zone, the P. baylei hori- zon of Normandy and the upper A. bauhini-bearing beds in Scotland (e.g. bed 38 with Pictonia sp., Wright 1989) could be expected. They may have 88 their equivalents anywhere in this succession, whereas other parts of the Submediterranean succession are not represented in the Subboreal sections or only by gaps. (c) The inclusion of this part of the Submediterranean subdivision in an A. lineatum Subzone (Malinowska 1991) with its unprecise limits (in southern Germany, the species is known to occur in the Upper Oxfordian and Lower Kimmeridgian) will not help significantly; this subzone can be replaced by the A. bauhini Zone, as used by Matyja & Wierzbowski (1997, 1998). (d) There are apparently different possibilities of cor- relation and further research is necessary to clarify the situation. (e) The Subboreal Oxfordian–Kimmeridgian bound- ary (R. pseudocordata/P. baylei Zone) can, with a high degree of probability, be positioned within the Submediterranean and Mediterranean scheme in the uppermost part of the E. bimammatum Zone on the basis of the correlation of the A. bauhini horizon with the P. densicostata horizon. The Submedi- terranean Oxfordian–Kimmeridgian boundary remains at the base of the S. platynota Zone. Additional remarks on Lower Kimmeridgian correlation As mentioned above, the upper S. galar Zone (A. sub- tilicaelatum horizon) is probably an equivalent of the Amoeboceras bayi horizon (Schweigert 1995a), which extends into the lower part of the S. platynota Zone (Amoeboceras horizon with A. bayi, see Atrops et al. 1993b). This contrasts somewhat with the correlation of Birkelund et al. (1983, table 1), who considered the Pictonia baylei Zone and the Paraspidoceras rupellense Zone of Hantzpergue (1979) to be equivalent. Hantz- pergue (1989), too, correlated the P. baylei Zone with the P. rupellense Zone (horizons R1 and R2); horizons P1–3 of the I. planula Zone sensu lato are considered to be equivalent to the R. pseudocordata Zone sensu lato (Hantzpergue 1989, tables E, F). In his sections, he found the Upper Oxfordian Sutneria galar in the Lithacosphinctes gigantoplex horizon (P3), immediately below his P. rupellense Zone (see Fig. 2). The P. rupellense Zone itself is situated between the gigantoplex horizon (P3) of the uppermost Idoceras planula Zone sensu lato and the Rasenia cymodoce Zone (Fig. 4); it is therefore considered to be equiva- lent to the lowermost Submediterranean Kimmeridgian (S. platynota Zone; Schairer 1970; Atrops 1982; Olóriz & Rodríguez-Tovar 1996); its lower horizon (R1) seems to correspond to the upper part of the lower (‘Ortho- sphinctes’) subzone of the S. platynota Zone, whereas the lower part (Amoeboceras horizon) of this zone is not represented; its upper horizon (R2) contains the index ‘Ardescia virgatoides’, which is similar to forms of the Ardescia desmoides horizon of the Ardescia desmoides Subzone of the middle Sutneria platynota Zone and is therefore very important for correlation to the Submediterranean region. Above the P. ruppelense Zone, Hantzpergue (1989) subdivided the Rasenia cymodoce Zone into nine hori- zons (C1–9); the R. cymodoce horizon (C2) could be traced from western France to Normandy and the Subboreal regions. In northern Europe, the R. cymodoce horizon is rather widespread (Wierzbowski 1989) and in Spitsbergen it represents the only rasenoid horizon within the Amoeboceras succession. In East Greenland, Birkelund & Callomon (1985, fig. 5) recognised two other horizons below the horizon of Rasenia cymodoce (‘17’), namely the ‘Pachypictonia’ horizon (‘16’) and the Rasenia inconstans horizon (‘15’). These horizons of the lower R. cymodoce Zone were considered to be equivalent to the P. altenense horizon (C1; Hantzperque 1989); they are probably equivalent to the lower Ataxioceras hippolytense Subzone of the lower Ataxioceras hypselocyclum Zone of south-east France, whereas the R. cymodoce horizon perhaps has its equiv- alents in the upper part of this subzone. In the middle and upper part of the R. cymodoce Zone, only a few possibilities remain for far-reaching corre- lations in Europe, such as the Eurasenia aulnisa hori- zon (C5), which contains the highly characteristic Submediterranean subzonal index A. lothari, and the Semirasenia askepta horizon (C7), which has been found in Scotland, England, Normandy, western France (Birkelund & Callomon 1985; Hantzpergue 1989) and southern Germany (Heller 1964; Doben & Heller 1968). In northern Germany, Submediterranean ammonites of Early Kimmeridgian age have been found in sediments which had earlier been attributed to the Upper Oxfordian (Fischer 1991). Substages The Kimmeridgian Stage has been subdivided into two or three substages; here a subdivision into three sub- 89 stages is preferred. If the Middle Kimmeridgian is not recognised, then the middle and the upper part are united as Upper Kimmeridgian (Fig. 4). Zones In the Mediterranean and Submediterranean provinces, the Lower Kimmeridgian consists of three zones, which can be correlated approximately as follows: (1) Sower- byceras silenum – Sutneria platynota, (2) Ataxioceras hypselocyclum – Taramelliceras strombecki and (3) Crussoliceras divisum – Mesosimoceras herbichi (Fig. 4). Their further subdivision into subzones is different in both areas (Fig. 4); precise correlation of these units is thus difficult (Pavia et al. 1987; Sarti 1993). Detailed subdivisions into subzones and faunal horizons have been proposed in south-east and western France (Atrops 1982; Hantzpergue 1989); that of south-east France can also be used with some minor changes in southern Germany. The Submediterranean zonal subdivision as established by Geyer (1961) can be used from the Iberian Peninsula to Bulgaria and Turkey (Sapunov 1977a; Lopez Marques 1983; Alkaya 1992). In Poland, the Submediterranean zonal subdivision has been adopted by Malinowska (1988) and Matyja & Wierz- bowski (1998). In Subboreal and Boreal regions, sub- division into two zones is typical (see above): (1) Pictonia baylei and (2) Rasenia cymodoce. These zones can be replaced by the Amoeboceras kitchini Zone in areas where no perisphinctids occur (e.g. Wierzbowski & Smelror 1993); this zone may extend into the lower part of the Aulacostephanus mutabilis Zone (see below). The Middle and Upper Kimmeridgian Substages together consist of three zones in all parts of Europe (Fig. 4). 1. In Mediterranean and Submediterranean Europe: (1) Aspidoceras acanthicum Zone, (2) Mesosimoceras cavouri or Aulacostephanus eudoxus Zone and (3) Hybonoticeras pressulum/H. beckeri or H. beckeri Zone. 2. In Boreal and Subboreal Europe: (1) Aulacostephanus mutabilis Zone, (2) A. eudoxus Zone and (3) A. autis- siodorensis Zone. In regions where no perisphinctids are present, these latter zones can be replaced in the lowermost parts by the Amoeboceras kitchini Zone (see above) followed by the A. kochi, A. elegans and Suboxydiscites taimyren- sis Zones (Fig. 4). The latter index has been taken from northern Siberia charts (Birkelund & Callomon 1985), but there is no mention of this species in more west- ern regions, with the exception of a determination from the Middle Kimmeridgian of Greenland. Therefore, for these Boreal regions too, Aulacostephanus autissio- dorensis seems to represent the more appropriate index species. The Middle Kimmeridgian zonal and subzonal sub- divisions can be applied without great difficulty in Boreal, Subboreal and Submediterranean Europe, as there are large regions with overlapping guide fossils, whereas in the Mediterranean province, only a zonal subdivision is possible. Hantzpergue (1989) established a detailed subdivision in western France, which can also be used in northern France (Geyssant et al. 1993; Proust et al. 1993) and traced as far as Germany (Zeiss 1991b; Schweigert 1993a, 1996a), England, Norway and East Greenland (Hantzpergue 1989). An unresolved problem is the lower boundary of the A. mutabilis Zone; it is drawn at the base of the A. lineatum horizon in western and northern France (Hantzpergue 1989; Hantzpergue et al. 1997), but in England, following the revisions of Birkelund et al. (1983), it is placed four horizons deeper, at the base of the S. askepta horizon. Recent investigations in central Poland came to similar results (Matyja & Wierzbowski 1998); these workers traced the boundary to a slightly deeper level in the upper A. hypselocylum Zone. In Germany and the Submediterranean region, the usage from south-east France has been followed (Hantzpergue 1989; Hantzpergue et al. 1991; Zeiss 1991b), which facil- itates correlation with the base of the A. acanthicum Zone; the lower boundary of this zone in Germany is traditionally drawn at the incoming of the first repre- sentatives of the genus Aulacostephanus (lineatum group). In a recent publication by Hantzpergue et al. (1997), the problems of this boundary are well illustrated by their table 12; in the ‘Biome Franco-germanique’, the lower boundary of the A. mutabilis Zone is drawn below its lowermost horizon (linealis horizon), whereas the base of the A. mutabilis Subzone, curiously, is placed two horizons higher (attenuatus horizon). It is evident that the new data from Poland (Matyja & Wierzbowski 1998), which place the base of the A. mutabilis Zone much deeper, will probably necessitate revision of all these correlations. Amoeboceras subdivisions are important from Norway to Spitsbergen (Wierzbowski 1989; Wierzbowski & Århus 1990; Wierzbowski & Smelror 1993) and East Greenland (Birkelund & Callomon 1985). 90 In the Upper Kimmeridgian (upper A. autissiodor- ensis Zone) of Poland and the Russian platform, a Sarmatisphinctes fallax Subzone has been established (Mesezhnikov 1984, 1988; Kutek & Zeiss 1994, 1997). For the lower part (lower A. autissiodorensis Zone), the Discosphinctiodes subborealis Subzone is proposed; D. subborealis is a significant index fossil. In Poland, Aulacostephanus autissiodorensis has been found only in the lower and middle parts of the S. fallax Subzone. In western Siberia, a zone of Virgataxioceras dividuum is the equivalent of the S. fallax Subzone (Mesezhnikov 1988). In northern Germany, Schweigert (1996a) stated, based on re-study of previous collections, that the A. autissiodorensis Zone is probably present. In southern Germany, where subdivision into two subzones was previously adopted, new discoveries of ammonites have made it possible to organise the H. beck- eri Zone into three subzones: (1) Sutneria subeumela, (2) Virgataxioceras setatum and (3) Lithacoceras ulmense (Schweigert & Zeiss 1994, 1999); further subdivision into several faunal horizons is possible (Schweigert 1996b, 1998). Furthermore, Schweigert (1993a, b, 1994) discovered ammonites in the Upper Kimmeridgian of Swabia with a Subboreal habitus, providing better cor- relation possibilities between the Subboreal A. autissio- dorensis and Submediterranean H. beckeri Zones (see below). For the Upper Kimmeridgian of western France, a useful subdivision has been proposed by Hantzpergue (1989), who subdivided the A. autissiodorensis Zone into two subzones, the A. autissiodorensis and the Gravesia irius Subzones, each with two faunal horizons. The succession in the Boulonnais area and farther north has been worked out in detail by Geyssant et al. (1993) and Geyssant (1994); the succession in southern England was reported by Cox & Gallois (1981), Birkelund et al. (1983) and Callomon & Cope (1996). Correlation Many difficulties are encountered in correlating zones (and subzones) of the Lower Kimmeridgian in Europe, mainly between the Submediterranean and Subboreal regions, but also between the Submediterranean and Mediterranean areas (Fig. 4). Many correlations are arbi- trary and well-constrained correlation is only possible at certain levels. Such correlation possibilities in the Lower Kimmeridgian Substage have already been explained in connection with the problems of the Oxfordian–Kimmeridgian boundary. Some problems exist around the Lower–Middle Kimmeridgian boundary, as the base of the A. mutabilis Zone is variably defined in different parts of Europe (see above). Considering the most recent results from Poland (Matyja & Wierzbowski 1998), the lower boundary of the Subboreal A. mutabilis Zone lies within the uppermost part of the Submediterranean A. hypselocyclum Zone, i.e. one zone deeper than previously assumed. In the Middle Kimmeridgian Substage, correlations within the A. acanthicum/A. mutabilis Zones and the A. eudoxus Zone pose no great problems although the uppermost part of the A. eudoxus Zone of western France (A. contejeani Subzone) seems to correspond to the lower part of the H. beckeri Zone in south Germany (Schweigert 1993b). Correlation of the A. kochi Zone with the upper part of the A. mutabilis and/or the lower part of the A. eudoxus Zone (Wierzbowski & Smelror 1993) is still tentative, as is the correlation of the A. ele- gans Zone with most of the A. eudoxus Zone. Correlation of the Upper Kimmeridgian Substage (Submediterranean H. beckeri Zone with the Subboreal A. autissiodorensis Zone) was hitherto only possible by indirect arguments. The elaboration of a new zonal and subzonal subdivision in western France by Hantz- pergue (1989) and the new discoveries by Schweigert (1993a, b, 1994) in Germany and by Kutek & Zeiss (1997) in Poland now permit correlation of parts of the Upper Kimmeridgian of western, central and eastern Europe and perhaps also western Siberia. Chronometric data The duration of the Kimmeridgian Stage has been esti- mated to be 3.4 Ma (Gradstein et al. 1995; Ogg 1995; Ogg & Gutowski 1996); for precise data, see Figure 4. Tithonian and Volgian (Fig. 5) The Tithonian, and its Boreal equivalent the Volgian, have been confirmed as stage names by a vote of the International Subcommission on Jurassic stratigraphy in 1990 (Zeiss 1991a). A further stage name ‘Bononien’ (for the ‘Upper Kimmeridgian sensu anglico’, proposed by Cope 1993) seems unnecessary and could result in each region with a differing zonal subdivision claim- ing its own stage name, leading only to more confu- sion rather than to international agreement concerning uniform nomenclature. Furthermore, due to the differ- ent meanings of the stage ‘Portlandian’ in different 91 countries, it was voted in 1990 that usage of this name should be discontinued. The most recent review of the Tithonian Stage and its ammonites is that provided by Geyssant (1997); for the Volgian Stage and ammonite biostratigraphy, see Gerasimov et al. (1995), Callomon & Cope (1996) and Kutek & Zeiss (1997). Lower boundary The base of the Tithonian Stage is defined by the base of the Hybonoticeras hybonotum Zone. It is generally supposed that the base of the coeval Gravesia gigas, Virgatosphinctoides elegans and Ilowaiskya klimovi Zones are drawn at approximately the same time level (see also below). Substages The Tithonian is subdivided into two or three substages; here preference is given to a tripartite Tithonian Stage (Fig. 5). If only two substages are used, then the lower and middle part are united as the lower substage (‘Danubian’), the upper substage corresponds to the ‘Ardescian’ substage. Type regions for the Lower and Middle Tithonian Substages have been proposed by Barthel (1975) and Zeiss (1975). The type region for the Upper Tithonian Substage, the Ardescian, has been revised by Cecca et al. (1989a, b). The subdivision of the Volgian is threefold, into lower, middle and upper substages. The lower and middle substages (‘Gorodishchian’) correspond roughly to the Tithonian Stage (Fig. 5), whereas the upper substage 92 N. Italy (S. Spain) E. Austria, Moravia Russian PlatformS. Germany Central Poland England Greenland Vulgaris (Durangites) [Crassicollaria] Subpalmatus ? Palatinus Vimineus Triplicatus Tagmersheimense Moernsheimense Rueppellianus Riedense Palmatus Glaber Ciliata Rothpletzi/ Penicillatum Albertinum (Darwini) Hybonotum Volanense (Ponti, ‘Burck- hardticeras’) Trans- itorius Transitorius [Crassicollaria] [Granulosa p.p.] [Dunkeri] Pseudoscythica Pseudoscythica Sokolovi Sokolovi Klimovi Klimovi Puschi Regularis Zarajskensis Scythicus Quenstedti Oppressus Nikitini Blakei Rosanovi Virgatus Zarajskensis Pavlovi (Disprosopa, Contradictionis) Oppressus Anguiformis Kerberus Okusensis Glaucolithus Albani Fittoni Rotunda Pallasioides Elegans Scitulus Eastlecottensis Para- virgatus Dorse- tensis Smedmorensis Wheatleyensis Reisiformis Encombensis Scitulus Wheatleyensis Hudlestoni Pectinatus Primus Iatrensis Rugosa Communis Liostraca Gracilis Pseudaperta Anguinus Groenlandicus Elegans Vogulicus (Pseudoscythica) Mucronatum Lithographicum Admirandum/ Biruncinatum Semiforme/ Verruciferum Richteri Richteri Volanense Austriacus [rugosa] Tenui- costata occiden- talis Tenui- costata Tenui- costata Magnum [Boneti] Scruposus Simplisphinctes M ic ra ca nt um U pp er M id dl e T it ho ni an Lo w er Fa lla ux i Fa lla ux i Sc yt hi cu s Pa nd er i Pe ct in at us H ud le - st o ni W he at - le ye ns is Pa ra vi rg at us V ir ga tu s M id dl e Vo lg ia n Lo w er V o lg ia n N ik it in i H yb o no tu m M uc ro na tu m V im in eu s Se m i- fo rm e 14 4. 0 (± 2. 5) 15 0. 7 (± 3. 0) Mediterranean Submediterranean Subboreal eastern western Boreal Fig. 5. A tentative correlation chart for the Tithonian and Volgian Stages in Europe (thick lines as in Fig. 2). Modified after Barthel (1964), Zeiss (1968, 2001), Cope et al. (1980), Callomon & Birkelund (1982), Kutek & Zeiss (1988, 1997), Mesezhnikov (1988), Sarti (1988), Zeiss & Bachmayer (1989), Mitta (1993), Kutek (1994) and Geyssant (1997). Non-ammonite taxa are indicated in square brackets. (‘Kashpurian’) belongs to the Cretaceous System (Sasonova & Sasonov 1979; Zeiss 1983, 1986; Sey & Kalacheva 1993a; W.A. Wimbledon in: Callomon & Cope 1996). A type section for the Volgian Stage has been proposed by Gerasimov & Mikhailov (1966). Zones and Subzones Whereas the two lower stages of the Upper Jurassic have two main zonal subdivisions, at least four subdivisions are necessary in the upper stage (Fig. 5). This is due to the extreme provincialism of ammonites caused by the increasing isolation of late Jurassic marine basins, which seem to have only rarely been directly connected; inter- basinal migration was apparently only favoured during the lowermost zone of the stage. The most important lower zone is that of Hybo- noticeras hybonotum, which can be followed over long distances in Mediterranean and Submediterranean Europe (Zeiss 1968; Olóriz 1978; Sapunov 1979; Sarti 1988); in southern Germany it is possible to recognise three subzones and seven horizons in the H. hybono- tum Zone (Schweigert & Zeiss 1999). In central Europe, the latter overlaps with the Gravesia gigas Zone, which has a rather wide distribution regionally in central and western Europe. During the last decades, many new dis- coveries have been reported and the genus Gravesia and the stratigraphy of the beds with Gravesia have been revised (Hahn 1963; Zeiss 1974; Hantzpergue 1989; Schweigert 1994, 1996a, b; Schweigert et al. 1996; Zeiss et al. 1996; Dimke & Zeiss 1997). In the Subboreal sub- province, the genus Gravesia is also present, but less numerous, so that other index fossils have been given priority, such as Virgatosphinctoides elegans in north- western and Ilowaiskya klimovi in eastern Europe (Cope 1967; Cope et al. 1980; Kutek & Zeiss 1974, 1994, 1997; Callomon & Birkelund 1982; Mesezhnikov 1988). According to Callomon & Cope (1996), Gravesia cf. gravesiana occurs in the lower part of the Virgato- sphinctoides scitulus Zone, thus demonstrating the cor- relation with the upper H. hybonotum Zone (containing G. gravesiana). In northern Germany, beds with Gravesia gigas intermedia are apparently the youngest beds containing Jurassic ammonites (Schweigert 1996a) and are succeeded by brackish and freshwater sedi- ments up to the Jurassic–Cretaceous boundary. In these beds, ostracodes have proved to be the best guide fos- sil (Bischoff & Wolburg 1963; Schudack 1994, fig. 24), permitting subdivision of the Tithonian Stage in north- west Germany into four zones. In other areas, such as eastern England and Denmark, subdivision into nine zones is possible using ostracodes (Christensen 1988; Schudack 1994, fig. 24). The upper zone of the Lower Tithonian in Mediterranean Europe, the zone of Semiformiceras dar- wini (or of Virgatosimoceras albertinum), is apparently equivalent to the Neochetoceras mucronatum and Franconites vimineus Zones (each of them with two sub- zones and some horizons) of Submediterranean Europe, as they have numerous faunal elements in common (Enay & Geyssant 1975; Olóriz 1978; Cecca et al. 1986; Sarti 1984, 1988; Cecca 1990a, b). Precise correlations have still to be worked out, however, and at present this is difficult as no subzones or even horizons have been recognised in the Tethyan realm. The Submedi- terranean zones have been traced from south-east France via southern Germany to Hungary as well as in Bulgaria and perhaps also Turkey (Zeiss 1968; Sapunov 1977b, 1979; Vigh 1984; Fözy 1988, 1993; Alkaya 1989; Atrops 1994; Fözy et al. 1994). Correlation with the Subboreal regions is only tentative and different proposals have been published (Fig. 5; Zeiss 1977; Mesezhnikov 1988; Kutek & Zeiss 1997). In Subboreal Europe, the situation is not much bet- ter and correlations between the different subprovinces of Northwest and eastern Europe are only approximate. Consequently, different zonal subdivisions are also applied in these subprovinces. In eastern Europe, for example, species of the genus Ilowaiskya are used (e.g. the Ilowaiskya sokolovi and I. pseudoscythica Zones; Mesezhnikov 1988; Kutek & Zeiss 1997), whereas in Northwest Europe, representatives of the genera Virgato- sphinctoides, Arkellites and Pectinatites have been selected (e.g. the Virgatosphinctoides scitulus, W. wheat- leyensis, Arkellites hudlestoni and Pectinatites pectina- tus Zones); each of these latter zones can be subdivided into two subzones (Cope et al. 1980; Callomon & Birkelund 1982; Geyssant 1997). For the Middle Tithonian Substage, the subdivisions in Mediterranean and Submediterranean Europe are rather distinct (Fig. 5). Furthermore, minor faunal differentiations exist within the Mediterranean area, and different zonal indexes are used for the same time inter- val (Enay & Geyssant 1975; Olóriz 1978; Cecca & Santantonio 1988; Sarti 1988): (1) Semiformiceras semi- forme or Haploceras verruciferum, (2) Semiformiceras fallauxi or (2a) Richteria richteri and (2b) Simoceras admirandum/biruncinatum (or S. biruncinatum), and (3) Simoceras volanense or ‘Burckhardticeras’ peroni or Micracanthoceras ponti. Note that Burckhardticeras Olóriz 1978 is a junior homonym of Burckhardticeras 93 Flores Lopez 1967 (Schweigert & Zeiss 1998). In the Submediterranean area of southern Germany, the fol- lowing guiding ammonites have been observed (Barthel 1975; Zeiss 1986): (1) Virgatosimoceras rothpletzi and Sublithacoceras penicillatum, (2) Lemencia ciliata, (3a) Sublithacoceras(?) glaber, (3b) Isterites palmatus, and (3c) Isterites subpalmatus. According to Scherzinger & Schweigert (1999), a horizon with Sublithacoceras cal- lodiscus has been observed above the level with Lemencia ciliata. In eastern Europe, the equivalents of the Middle Tithonian Substage are probably the upper part of the Lower Volgian (upper Ilowaiskya pseudoscythica and Ilowaiskya tenuicostata Zones). The latter unit is dis- cernable in Poland but has not been recognised in Russia to date (Kutek & Zeiss 1974, 1988, 1994, 1997; Mesezhnikov 1988; Kutek 1994). In its upper part, the Pseudovirgatites puschi horizon is important due to its mixed fauna (Kutek & Zeiss 1974, 1988, 1997). A local time equivalent in north-eastern Austria is probably the Isterites austriacus Zone with Buchia rugosa as an important guide fossil (Fig. 5). A quite different zonal subdivision exists in Great Britain and the adjoining Subboreal and Boreal regions as far as Greenland (Cope 1978, 1980; Wimbledon 1980; Callomon & Birkelund 1982; Kejsi et al. 1988); the Middle Tithonian perhaps corresponds to the main part of the Pectinatites pecti- natus Zone and perhaps to the Pavlovia pallasioides Zone of England or to the Dorsoplanites primus and Pavlovia iatrensis Zone of East Greenland. The Upper Tithonian Substage consists of two or three zones in the Mediterranean area. In southern Spain, the lowermost zone has been identified as the Simplisphinctes Zone (Tavera 1985). This unit has not been identified in northern Italy (Sarti 1988), but could be recognised as far as north-eastern Austria, where the same ammonite fauna (containing the genus Oloriziceras) occurs (Zeiss & Bachmayer 1989). In the absence of the rather peculiar index genus Simpli- sphinctes, this zone was called the Oloriziceras mag- num Zone for this region (Zeiss 2001). Above the Sim- plisphinctes (or S. abnormis or O. magnum) Zone, the Paraulacosphinctes transitorius Zone (with the first Crassicollaria) occurs. A Micracanthoceras micracan- thum Zone is sometimes adopted instead of the P. tran- sitorius Zone; this zone apparently also contains the equivalents of the Simplisphinctes (better S. abnormis) Zone (Enay & Geyssant 1975; Sarti 1988; Geyssant 1997). Some authors consider the Simplisphinctes and P. tran- sitorius Zones as subzones of the M. micracanthum Zone (Benzaggagh & Atrops 1997; Geyssant 1997) although the former authors, based on Moroccan data, only partially substituted the Simplisphinctes Subzone, replacing its upper part and the P. transitorius Subzone by two new subzones, that of ‘Micracanthoceras (Corongoceras) spp.’ and that of ‘Moravisphinctes spp.’. It is very important that these new subzones can be cor- related rather precisely with the calpionellid subdivi- sion; the Chitinoidella boneti Subzone (of the Chitinoid- ella spp. Zone) corresponds to the first two subzones. The base of the Crassicollaria spp. Zone (Zone A) approximately coincides with the base of the Moravisphinctes spp. Subzone, which corresponds to the lower part of this zone (= Subzone A1). The Durangites Zone follows above the P. transitor- ius Zone. In northern Italy, this zone was named the Durangites vulgaris Zone by Sarti (1988); this term has also been adopted by other authors. In some countries, this zone has not been recognised; the equivalents of this zone are then apparently included in the P. tran- sitorius Zone, which sometimes even includes parts of the Lower Cretaceous (e.g. Sapunov 1977b). The fauna of this zone has been mainly described by Tavera (1985), Tavera et al. (1994) and Enay et al. (1998a, b). During the Middle Volgian, central Poland belonged to the eastern Subboreal subprovince, but only the low- ermost unit, the Zaraiskites scythicus Zone (with the lower Z. scythicus and upper Z. zarajskensis Subzones) is represented (Kutek 1994). Brackish sediments pre- vail higher in the Polish section and yield ostracodes; the Cypridea dunkeri and the Cypridea granulosa Zones can be recognised. On the Russian Platform, the low- ermost horizon of the Z. scythicus Subzone (Z. quen- stedti horizon in Poland) is probably represented by beds containing Zaraiskites disprosopa and Isterites(?) con- tradictionis (Ilovaiskij & Florenskij 1941). On the Russian Platform, a Dorsoplanites panderi Zone is now used instead of the Z. scythicus Zone (Mesezhnikov 1988; Kutek 1994); above follows the Virgatites virgatus Zone (with three subzones: V. gerasi- movi, V. virgatus and C. ivanovi; Gerasimov et al. 1995). The V. virgatus Zone is succeeded by the Epivirgatites nikitini and Lomonossovella blakei Zone (separated by Callomon & Birkelund (1982), and, in reverse order, by Mesezhnikov (1988) but adopted as a single zone by other Russian authors (e.g. Gerasimov et al. 1995)). The uppermost Middle Volgian is represented by the Paracraspedites oppressus Zone (Mesezhnikov 1988). In the Baltic area, Middle Volgian ammonites are rare although a few specimens from Lithuania were men- tioned by Rotkyte . (1976, 1987). In Scandinavia, Middle Volgian ammonites have been found in Denmark 94 (Birkelund & Pedersen 1980) and in Norway (Birkelund et al. 1978). In England and East Greenland, Dorso- planitidae are prevalent, but in both these regions, the subdivisions are distinct; in England, Pavlovia palla- sioides, Pavlovia rotunda and Virgatopavlovia fittoni characterise the lower part of the Middle Volgian whereas Progalbanites albani and the giants Glaucolithites glau- colithus, Galbanites okusensis, Kerberites kerberus and Titanites anguiformis characterise the upper part (Cope 1978; Wimbledon & Cope 1978). As in Russia, the upper- most zone is the Paracraspedites oppressus Zone (Casey 1973; Kejsi & Mesezhnikov 1986; Kejsi et al. 1988), but not all authors adopt this zone. In East Greenland, there are some similarities with Siberian ammonite successions, but in general the subdivision there has its own char- acter and, with three exceptions, its distinct index species (Callomon & Birkelund 1982; Mesezhnikov 1988): Dorso- planites primus, Pavlovia iatrensis, Pavlovia rugosa, Pavlovia communis and Dorsoplanites liostracus char- acterise the lower part of the Middle Volgian, whereas Dorsoplanites gracilis, Epipallasiceras pseudapertum, Crendonites anguinus, Laugeites groenlandicus and Epilaugeites vogulicus are represented in the upper part. The lower part of the Upper Volgian Praechetaites tenuicostatus Zone of East Greenland may correspond to the uppermost part of the Middle Volgian, the upper Paracraspedites oppressus Zone of England and the lower Praechetaites exoticus Zone (= lowermost Cras- pedites okensis Zone sensu lato) of northern Siberia. Correlation As explained above, the basal zones of the Tithonian (and Volgian) can be correlated over long distances, but correlation becomes very difficult in the higher parts of these stages. Not only is it difficult to correlate between the Boreal and Mediterranean regions, but also within these regions. Distinct lineages of ammonites were evolving throughout the area and consequently it is necessary to develop and apply different ammonite zonal subdivisions; correlation possibilities are thus only few and mostly tentative. Many attempts have been made to correlate the different zonal subdivisions of Europe (Cope & Zeiss 1964; Zeiss 1965, 1974a, 1979, 1983, 1986; Enay 1972; Enay & Geyssant 1975; Olóriz 1978; Callomon & Birkelund 1982; Jeletzky 1984, 1989; Tavera 1985; Cecca et al. 1986; Hoedemaker 1987, 1991; Kejsi et al. 1988; Kutek & Zeiss 1988, 1997; Geyssant & Enay 1991; Sey & Kalacheva 1993a; Kutek 1994; W.A. Wimbledon in: Callomon & Cope 1996; Geyssant 1997). Due to problems of provinciality, such correlation schemes are necessarily speculative and ultimately unsat- isfactory. A tentative summary correlation scheme is given in Figure 5, based on developments since earlier attempts by the author (Zeiss 1983, 1986). A similar, although in detail somewhat different, correlation chart has recently been published by Hantzpergue et al. (1998). Concerning the Middle and Upper Tithonian (upper Lower and Middle Volgian) Substages, a number of observations are pertinent. Although correlation between the Mediterranean and Submediterranean area is quite possible in the lowermost Middle Tithonian Substage (S. semiforme/R. richteri – V. rothpletzi/S. pennicilatum Zones), a number of different proposals have been made for the higher zones (Enay & Geyssant 1975; Olóriz 1978; Jeletzky 1984, 1989; Cecca et al. 1986; Kutek 1994). A satisfactory answer to this problem requires complete revision of the famous Submedi- terranean Neuburg fauna and sections, in which some levels with distinct ammonite faunas have already been recognised by Barthel (1964, 1975). In eastern central Europe (north-eastern Austria, Moravia, central and southern Poland), some Submedi- terranean and Mediterranean ammonites genera of Middle and Late Tithonian age are represented by char- acteristic forms. They sometimes interfinger with Sub- boreal elements, thus providing good potential for correlation (Kutek & Wierzbowski 1986; Kutek & Zeiss 1988, 1997; Kutek 1994). The I. tenuicostata and Z. scythicus Zones of central and southern Poland, for example, display interesting forms with affinities to both the Submediterranean and Subboreal provinces. Combined with observations from other localities, this facilitates better correlation between these two regions: (1) the Pseuvirgatites puschi horizon of the uppermost Ilowaiskya tenuicostata Zone contains Isterites species described from the higher parts of the Neuburg beds, i.e. of late Middle Tithonian age, and (2) the Z. regu- laris horizon of the lower Z. zarajskensis Subzone (upper Z. scythicus Zone) contains Pseudovirgatites scruposus and calpionellids indicative of the calpionellid Zone A, such that correlation is possible with the lower part of the Paraulacosphinctes transitorius Zone. In the Boreal and Subboreal provinces, quite differ- ent zonal subdivisions exist, mainly based on different perisphinctid groups, such as the Pectinatitinae and Dorsoplanitinae in England, Denmark, Norway and Greenland and the Ilowaiskyinae, Virgatitinae and Dorsoplanitinae in Poland and Russia. The correlation of these zones is rather arbitrary, as demonstrated by Callomon & Birkelund (1982), Mesezhnikov (1988) and 95 W.A. Wimbledon (in: Callomon & Cope 1996), and is based mainly on similar, but non-identical species of Dorsoplanitinae. Chronometric data The approximate duration of the Tithonian has been estimated to be 6.7 Ma (Gradstein et al. 1995; Ogg 1995); for precise data, see Figure 5. Biochronological importance of non- ammonite fossil groups: a review The Jurassic System is the classic one for subdivision by ammonites. This fossil group has been used with much success since the pioneering work in the last century by workers such as L. von Buch, A. d’Orbigny, A. Oppel, F.A. Quenstedt, K.A. von Zittel and S. Buck- man. Indeed, this contribution on the chronological subdivision of the Upper Jurassic of Europe has been compiled primarily using ammonites (see above). However, Upper Jurassic marine sediments of epicon- tinental shelves, the habitat of ammonites, are not pre- sent everywhere in Europe, so that ammonites are not always available. It is often necessary, therefore, to utilise other fossil groups with proven stratigraphic value such as bivalves, brachiopods, foraminifera, ostra- codes and distinct plant mega-, micro- and nannofos- sil groups. Radiolarians, calpionellids, conchostracans, insects and vertebrates should also be added to this list; the first two groups are very useful in pelagic sedi- mentary basins whereas the last ones are used with much success in the stratigraphic subdivision of conti- nental sediments, such as those of central and eastern Asia and of North America. The challenging task of correlating between the dif- ferent fossil subdivision schemes has been addressed for individual groups (e.g. Le Hégarat & Remane 1968; Surlyk & Zakharov 1982). Multidisciplinary correlation charts, typically for microfossil groups, have only been successfully developed within the last two decades. Useful though incomplete examples of such schemes, including micro- and macrofossils, have recently been published by Tavera et al. (1994), R. Enay (in: Cariou & Hantzpergue 1997), Gramann et al. (1997) and Remane (1997). It remains as one of the more important tasks, however, to establish European multidisciplinary cor- relation charts that incorporate all fossil groups impor- tant for biochronology and also include radiometric ages and palaeomagnetic reversal data. During the editorial work, it was brought to the attention of the author that charts fulfilling many of these expectations have recently been published by Hardenbol et al. (1998, charts 6–7); of special interest are the chronometric data for most of the biochronostratigraphic units (see below). Invertebrate megafossil groups Cephalopods – other than ammonite conchs Aptychi In the Tethyan regions, aptychi have proven to be a use- ful addition to ammonites for the subdivision of Upper Jurassic sediments. Following the studies of Durand & Gąsiorowski (1970) and Gąsiorowski (1962, 1985), it is possible to differentiate eleven zones of aptychi using four larger groups of aptychi, the lamellaptychi and laev- aptychi and to a lesser degree the laevilamellaptychi and punctaptychi. Correlation between aptychi and ammonite zones still poses problems (A. Wierzbowski, personal communication 1998). Eliás̆ et al. (1996) also used apty- chi ranges for biostratigraphy, but without a zonal sub- division; they preferred a multidisciplinary correlation method using the calpionellid subdivision as reference. Belemnites The most recent review of this fossil group is that of Doyle & Bennett (1995) which includes a section on Middle and Upper Jurassic belemnite groups, includ- ing those of Europe. This publication presents a com- prehensive review of the subject, including the work of Saks & Nalnyaeva (1964, 1966), Riegraf (1980, 1981), Combémorel & Mariotti (1986), and Doyle & Kelly (1988); a range chart of the most useful taxa for biostrati- graphy of the Middle and Upper Jurassic is included by Doyle & Bennett (1995). The stratigraphic ranges of some more important Polish species have been pub- lished by Pugazewska (1988) and Malinowska (1997) and those of Sicily by Combémorel & Mariotti (1990). Recently, Combémorel (1997) compiled all data avail- able for the Tethys and the Boreal region of Europe and for each of them presented a correlation scheme with the subdivisions based on ammonites and belemnites; see also Hardenbol et al. (1998, chart 7). 96 Bivalves The most important group of bivalves for biostrati- graphic purposes in the Upper Jurassic of Europe is the genus Buchia. In the Boreal regions of Eurasia and North America, it is of particular importance as a sup- plement to ammonites. The genus has been the focus of many papers in the last decades such as Zakharov (1981, 1987, 1990), Surlyk & Zakharov (1982), Jeletzky (1984), Kelli (1990), Sey & Kalacheva (1993b) and Sha & Fürsich (1994). An interesting interpretation of the dif- ferent ranges of Buchia species in America and Eurasia has been presented by Hoedemaker (1987). Stratigraphic range lists of selected bivalve species from Poland have been published by Karczewski & Pugaczewska (1988) and Malinowska (1997). A correlation chart that is mainly based on buchiid bivalves but also includes other bivalve genera (e.g. Retroceramus) has been compiled for northern Russia and the circum-Pacific regions by Damborenea et al. (1992). In the Upper Jurassic, the stratigraphic resolu- tion of bivalve taxa, with the exception of buchiids, seems to be rather limited and/or needs further research (Damborena et al. 1992). For some regions, stratigraphic range lists of selected bivalve species have been pub- lished, for example for Poland (Malinowska 1997; Karczewski & Pugaczewska 1998) and for northern Germany by Kaever et al. (1976). Gastropods The biostratigraphic resolution of this group in the Jurassic is not very high, but in special cases, when other guide fossils are not present, some representatives of the group may be used. An example from the Upper Jurassic of France (Nerineaceae) has been published recently by Barker (1994). Range lists of selected species from Poland have been published by Karczewski (1988) and Malinowska (1997). Brachiopods The most recent reviews of this group with respect to Upper Jurassic brachiopods are those of Ager (1994) and Alméras et al. (1991, 1994), especially for France and Britain, and Boullier & Laurin (1997) for the Tethys and the ‘Domaine NW européen français’. Ager (1994) con- sidered the group within a global context. Alméras et al. (1994) discussed the facies dependence of brach- iopods, concluding that distinct zonal species of brach- iopods are often necessary for different facies. For biostratigraphical purposes, it is possible to subdivide the Upper Jurassic of England and north-west France into nine zones and some subunits. The Polish species have been figured and described by Barczyk (1988); range charts are given in Malinowska (1997). Prozo- rovskaja (1993) presented an overview of the brachio- pod subdivision of the Upper Jurassic of the southern part of the former USSR. Echinoderms To date, there is no subdivision scheme of the Upper Jurassic with respect to echinoderms. Some genera have biostratigraphic value; Saccocoma, for example, has been used in some multidisciplinary schemes. Thierry et al. (1997) presented range charts of the Upper Jurassic regular and irregular echinoid genera and species of France, with the expectation that with detailed research it would be possible to create a subdivision scheme com- parable to that developed for the brachiopods of France. Corals (scleractinians) This group has poor biostratigraphic resolution. Its use- fulness for stratigraphic purposes is therefore rather limited, also because of the close dependence of corals on ecological factors (Rosendahl 1988). Nevertheless, Beauvais (1988) subdivided the Upper Jurassic Series (except the Lower Oxfordian) into six zones based on madreporians (scleractinians). Polish species with range charts have been presented by Roniewicz & Morycowa (1988) and range charts were published by Malinowska (1997). Sponges This fossil group is poorly suited to regional correla- tion, but some species may be useful for local subdi- vision; examples from France have been presented by Gaillard (1997). Vertebrate megafossils Jurassic vertebrate fossils are too scarce to be used as guide fossils. Nevertheless, if vertebrate remains are 97 studied thoroughly, they frequently provide valuable biostratigraphic information (e.g. elasmobranchian teeth, Gramann et al. 1997). It should be mentioned that the Jurassic Period in Europe saw the early evolution of mammals, the flourishing of the first true birds and the first wave of the acme of the dinosaurs. In other parts of the globe, vertebrates have been used for stratigraphy; in North America, for example, Turner & Peterson (1998) subdivided the Upper Jurassic Morrison Formation into four biozones on the basis of dinosaurs, whereas in China, fish are used for subdivi- sion (Chen 1990). Invertebrate microfossils Foraminifera In the 1950–60s, foraminifera were one of the most important microfossil groups, together with ostracodes, for relative age determinations of marine sediments in boreholes; their importance has decreased in more recent times. Studies of foraminifera faunas from out- crops in southern Germany were reviewed by Groiss (1984). An account of epistominian zonation was given by Ascoli (1988), who also presented zonations and correlations between east Canadian offshore wells and the East European Platform (Grigelis & Ascoli 1995). Foraminifera from northern Germany were presented by Klingler et al. (1962) and Gramann et al. (1997). The guide fossils and characteristic species of the Upper Jurassic foraminifera of Poland have been published by Bielecka (1988) and Styk (1997), those of the Russian Platform by A.Y. Azbel (in: Mesezhnikov 1989). Fora- minifera of Sweden were studied by Norling (1972) and Guy-Ohlson & Norling (1988). A short compilation of Upper Jurassic foraminifera in Britain has been published by Shipp & Murray (1981), together with a range chart and figures of index species. The most recent reviews of foraminifera of Europe have been compiled by Ruget & Nicollin (1997) on the small benthic forms, and by Bassoulet (1997a) on the large forms; see also Hardenbol et al. (1998, chart 7). Radiolaria This microfossil group, which has been the subject of much scientific research in recent years in Europe, is of particular importance in the Tethyan region. A com- prehensive monograph was recently published by Baumgartner et al. (1995) on the radiolarians of the Tethys, including a catalogue of all Tethyan species. The biochronological potential for subdividing the Upper Jurassic Series into ‘Unitary Association Zones’ (U.A.Z.) is well-demonstrated; there are six such zones cover- ing the whole Upper Jurassic. They have a duration of between 2–6 Ma. This monograph demonstrates the significant advances in research into this group, espe- cially if new quantitative concepts, such as the ‘Unitary Association Zones’, are applied to the biochronologi- cal subdivision of the Upper Jurassic. Research into radiolarians and their stratigraphic potential has also been on the increase outside the Tethys, as demonstrated by recent publications con- cerning the Submediterranean province (Riegraf 1987; Kießling 1997; Zügel 1997; Zügel et al. 1998), and even the Subboreal and Boreal provinces, including the North Sea (Dyer & Copestake 1989), the Russian Platform and the Barents Sea (Vishnevskaya 1993, 1997, 1998; Kozlova 1994). Dyer & Copestake (1989) introduced a biozona- tion based on a succession of ten radiolarian events in the Kimmeridgian and Tithonian. Important attempts are also underway to correlate the new peri-Tethyan radi- olarian assemblages with different micro- and macro- fossil biozonations (Vishnevskaya & De Wever 1997); owing to strong provincialism, direct correlation between the peri-Tethyan and Tethyan zonations is still very dif- ficult, but has been undertaken recently (Hardenbol et al. 1998, chart 7). Ciliata This group is important only in the Tethyan region and the surrounding shelf deposits; the most comprehen- sive studies of the ciliata in recent years have been published as a result of the Sümeg meeting (Fülöp 1986; Remane et al. 1986). Polish forms have been reported by Nowak (1988) and those of Spain by Tavera et al. (1994) and Olóriz et al. (1995). Remane (1997, 1998) recently published informative reviews of the state-of- the-art of the group, providing tables which include the stratigraphic succession of calpionellid species and the correlation of calpionellid, nannofossil and ammonite subdivisions with magnetostratigraphic events. Nearly simultaneously, Blau & Grün (1997a, b) and Grün & Blau (1996, 1997) proposed a revision of the calpionellid zonal and subzonal division. For the Tithonian Stage, they introduced and formally defined two zones and seven subzones; the duration of zones in the Jurassic 98 is less than one million years, that of subzones about 300 000 years. Important results from the southern Tethyan margin have been contributed by Benzaggagh & Atrops (1995, 1997). These workers provided precise correlation and species range charts for ammonites and calpionellids for the lower part of the calpionellid succession, which previously was poorly known, and clarified the suc- cession of zones and subzones from the Middle Titho- nian Semiformiceras fallauxi/Chitinoidella dobeni Subzone to the Upper Tithonian Durangites vulgaris/ Crassicollaria A3 Subzone. An important contribution on the calpionellid faunas of the southern and eastern Tethyan region of Europe was presented by Reháková & Michalík (1997); the western Carpathians and their foreland in Moravia were treated by ̌Rehánek (1990) and Reháková (1995, 2000). In all these last-mentioned pub- lications, the Middle/Upper Tithonian boundary has apparently been drawn a little too high. Following the results of Benzaggagh & Atrops (1995, 1997), this bound- ary lies between the Dobeni and Boneti Subzones of the Chitinoidella Zone and not above this zone. Ostracodes This group has a rather high stratigraphic resolution and has therefore been used frequently and success- fully for the subdivision of sediments in northern Germany, Poland, England, the Netherlands, the North Sea Basin, France and Russia. In a recent monograph, Schudack (1994) revised the ostracodes of the Upper Jurassic in north-west Germany, documenting the cor- relation possibilities of this group in western, central and northern Europe. The Upper Jurassic of north-west Germany was subdivided into nineteen ostracode zones, representing variable durations (0.25–2.5 Ma; Schudack 1996a; Gramann et al. 1997). This study also presents a comprehensive list of all important publications on ostracodes. In northern Europe, the papers of Herngreen et al. (1988), Herngreen & Wang (1989) and Guy-Ohlson & Norling (1994) deal with this group in the Nether- lands and Sweden, respectively. In Poland, Bielecka et al. (1988) treated the group, and range charts have been published by J. Szteijn (in: Marek & Pajchlova 1997); Danish faunas were described by Christensen (1988). The most recent reviews of European ostra- codes are those of Bodergat (1997) on marine ostracodes and Colin (1997) on non-marine ostracodes; see also Hardenbol et al. (1998, chart 7). Plant microfossils Dinoflagellata Dinoflagellate cysts have become a widely used sup- plement to ammonites and are of particular importance in the subsurface. In a recent study, Poulsen (1996) emphasised the important role of dinoflagellates in Jurassic stratigraphy while comparing the Upper Jurassic of Denmark and Poland. The marine Upper Jurassic of Denmark was divided into seven zones and fifteen sub- zones whereas that of Poland was divided into four zones and twelve subzones (Poulsen 1996); the dinofla- gellate cyst zonation of the Jurassic of Subboreal Europe is reviewed in Poulsen & Riding (2003, this volume). In Great Britain, Riding & Thomas (1992) have deliv- ered the most recent compilation of dinoflagellates. Other important papers are those of Sarjeant (1979), Riley (1980), Riley & Fenton (1982) and Riding & Sarjeant (1984); one concerning Russia is that of Lentin & Vozzhennikova (1990). In the Netherlands, Herngreen et al. (1988; see also Herngreen & Wang 1989) pre- sented a report on the stratigraphic bioevents based on the first and last appearance of dinoflagellate cyst species which made possible a subdivision into nine zones. In north-west Germany, the Oxfordian and Kimmeridgian has been subdivided into three dinoflagellate zones and eight subzones (Gramann et al. 1997). Detailed subdivisions for the Boreal and Tethyan regions have recently been published by Hardenbol et al. (1998, chart 7). Calcareous nannofossils (coccoliths, nannolith groups) Recent advances in Jurassic calcareous nannofossil research have been reviewed by Bowen (1996), who dealt with several general aspects of this group, such as evolutionary succession, species diversity and longevity, distribution and provincialism, which are all important when regarding the utility of the group for biostratigraphic purposes. If conditions are favourable, then it is possible to subdivide the Upper Jurassic into five Boreal nannofossil zones (with six subzones) or three Submediterranean nannofossil zones (with seven subzones); correlation between these two regions is thus still problematic. The calcareous nannofossil bio- events were recently reviewed by Gardin (1997). Subdivisions and correlations for the Boreal/Subboreal 99 and the Tethyan/Submediterranean Provinces can be found in Hardenbol et al. (1998, chart 7). Charophyaceae This group of calcareous algae has received new impe- tus with respect to its potential for biostratigraphy. In a recent publication, the results of a local zonal subdi- vision based on charophytes in the Lower Saxony Basin of north-west Germany (Schudack 1996b) has been correlated firstly with the new European Mesozoic charophyte biozonation (Riveline et al. 1996), secondly with the subdivisions of other microfossil groups in north-west Germany, such as ostracodes and dinocysts, and thirdly with the old micropalaeontological subdi- visions for the Upper Jurassic (Malm) of north-west Germany (e.g. Klingler et al. 1962; Wick & Wolburg 1962). In north-west Germany, from the Upper Oxfordian to the top of the Tithonian, five charophyte zones are now recognised, whereas in other parts of western Europe there are only three (Schudack 1991, 1993). The stratigraphic resolution of this group is not very high in the Upper Jurassic. Each biozone represents a dura- tion of between 0.5 and over 2 million years. The Charophyaceae of western Europe have been revised in detail by Schudack (1993), and a useful compilation of all new data in Europe has been compiled by Riveline et al. (1996); see also Hardenbol et al. (1998, chart 7). Dasycladaceae This group seems to be only locally important for bio- stratigraphy (e.g. Portugal, Italy, Dinarids); a short review was presented by Bassoulet (1997b). Spores and pollen The value of pollen and spore grains for stratigraphic subdivision is not very high in the Upper Jurassic. The palynostratigraphy of Sweden (north-west Skåne) was discussed by Guy-Ohlson & Norling (1988) in connec- tion with a study of the microflora of some boreholes. It was revealed that “detailed correlation without the presence of dinoflagellates or other significant taxa appears difficult if not impossible” (Guy-Ohlson & Norling 1988, p. 15). In the Central Graben of the south- ern North Sea, Upper Jurassic sediments were subdi- vided into four zones on the basis of sporomorphs (Herngreen et al. 1988; Herngreen & Wang 1989). In north-west Germany, the Upper Jurassic was divided into four zones using spores and pollen (Gramann et al. 1997). The group apparently has its greatest importance at the system boundaries; it has been used successfully at the Triassic–Jurassic boundary and, to a lesser degree, at the Jurassic–Cretaceous boundary. Magnetostratigraphy This important method has become more directly applic- able for stratigraphic purposes in the last few decades, especially when combined with radiometric and bio- stratigraphic data. Some of the more important papers on this topic are: (1) Mesozoic in general, Harland et al. (1990), Gradstein et al. (1995); (2) Upper Jurassic – Lower Cretaceous, Ogg (1983), Ogg et al. (1984), Odin et al. (1994); (3) Oxfordian, Steiner et al. (1985), Ogg & Steiner (1988a), Ogg et al. (1991), Ogg & Coe (1998); (4) Oxfordian – Lower Kimmeridgian, Ogg & Gutowski (1996); (5) Kimmeridgian–Tithonian, Ogg et al. (1994), (6) Jurassic–Cretaceous boundary, Ogg et. al. 1984), Ogg & Lowrie (1986), Ogg & Steiner (1988b), Ogg et al. (1991, 1994). In a recent publication on sequence chronostratigraphy of European Mesozoic basins, charts with magnetochronostratigraphic units have been com- piled together with sequence chronostratigraphic and biochronostratigraphic data (Hardenbol et al. 1998, see below); the time span of the Upper Jurassic contains polarity chronozones M35 (upper part) – M19. Sequence chronostratigraphy Sequence stratigraphy is gaining in importance in chronostratigraphic correlation, as illustrated recently by the presentation of a framework for the European Mesozoic and Cenozoic basins (Hardenbol et al. 1998). Many data have been used and compiled in charts, two of which are important for the Upper Jurassic. They demonstrate the sequence chronostratigraphy (Sequences, T-R Facies Cycles, Major Transgressive–Regressive Cycles) for the Boreal and Tethyan realms combined with the ammonite biochronostratigraphy and magne- tostratigraphy, plotted against the time scale. The Upper Jurassic of Europe starts in the upper half of the transgressive part of the Second Major T-R Cycle (1st order cycle, named North Sea Cycle) in the Jurassic and ends within the regressive phase of this cycle. A 100 total of 21 sequences (3rd order cycles) have been recognised (Ox 0–8, Ki 1–7, Ti 1–6) and three 2nd order T-R cycles (T8b–R10b) in the Boreal area, whereas the number in the Tethyan area is somewhat lower. A detailed overview of the North Sea Cycle in Europe (from the North Sea to south-east France) has been pre- sented by Jacquin et al. (1998); marginal areas have been studied as follows: East Greenland (Surlyk 1991; 2003, this volume), Portugal, Lusitanian Basin (Leinfelder & Wilson 1998), Portugal and Spain, South Iberian Margin (Olóriz et al. 1991), south-east France (Jan du Chêne et al. 2000), Switzerland (Gygi et al. 1998), West Carpathians (Reháková 2000) and Russia (Sahagian et al. 1996). Acknowledgements The author expresses his sincere thanks to Jon R. Ineson who kindly improved the English text and provided many useful suggestions. Two critical readers, Beris M. Cox and Andrzej Wierzbowski, contributed much to the advancement of this paper by their useful propos- als and comments. Many colleagues helped with lite- rature; in particular, I should like to mention Jean Guex, Franç̧ois Atrops, Elie Cariou, Raymond Enay, Reinhart Gygi, Vassili Mitta, Zdenek Vasicek and Andrzej Wierzbowski. Technical help has been provided by F. Boehm, H. Forke, W. Kießling and E. 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