Palynostratigraphy and palaeoenvironment of the Middle Sortehat Formation (Neill Klinter Group), Jameson Land, East Greenland

Eva B. Koppelhus and Carina F.Hansen

The grey–black mudstones of the Sortehat Formation form part of the Middle Jurassic fill of the Jameson Land Basin in East Greenland. The formation is exposed in the southernmost part of the north–south-trending, rift system in East Greenland that was part of the epeiric sea- way between East Greenland and Norway. Sedimentological observations of the Sortehat Formation indicate deposition in an offshore marine setting that was typically low energy and periodically oxygen-deficient but was influenced by storm currents on occasion. Detailed palynological stud- ies of the Sortehat Formation have resulted in the definition of three palynological assemblage zones recognised at four localities, namely Enhjørningen Dal and Pelion (north Jameson Land), the type section at Sortehat (central Jameson Land) and Albuen at Neill Klinter along Hurry Inlet (south-east Jameson Land). In stratigraphic order, these zones are termed the Botryococcus Assemblage Zone, the Nannoceratopsis gracilis – Nannoceratopsis senex Assemblage Zone, and the Sentusidinium pelionense Assemblage Zone. They are recognised on the basis of the iden- tification of approximately 110 species of palynomorphs, including 45 species of spores, 30 of , 22 of dinoflagellate cysts, 10 acritarch species, two species of algae, and some fungal spores. An Aalenian – ?Early Bajocian age is suggested for the Sortehat Formation on the basis of the palynoflora. Interpretation of the palynomorph assemblages suggests that the formation accumulated in a shallow, brackish marine environment. A significant terrestrial input, including the freshwater green alga Botryococcus, is recorded in the lower part of the formation and interpreted as an allochtho- nous accumulation in an offshore marine environment related to transgression of a low-lying coastal plain. A marked shift in the palynomorph assemblage seen by diversification of marine microplankton above the base of the formation, indicates an increase in the marine signal prob- ably related to the onset of highstand conditions following the marine transgression.

Keywords: East Greenland, Jameson Land Basin, Middle Jurassic, Aalenian – ?Early Bajocian, palynostratigraphy, sedimentology, palaeoenvironment, transgressive–highstand mudstones, allochthonous Botryococcus assemblage

E.B.K., Geological Survey of Denmark and Greenland, Geocenter Copenhagen, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark. Present address: Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller T0J 0Y0, Alberta, Canada. E-mail: [email protected] C.F.H., Geological Institute, University of Copenhagen, Geocenter Copenhagen, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark. Present address: Skovhegnet 4, DK-3460 Birkerød, Denmark.

The Middle Jurassic Sortehat Formation of the East between the sandstone-dominated Ostreaelv Formation Greenland rift basin is a distinctive and laterally per- beneath and the sandstones of the Pelion Formation sistent mudstone-dominated succession sandwiched above (Figs 1, 2). The stratigraphic terminology adapted

Geological Survey of Denmark and Greenland Bulletin 1, 777–811 (2003) © GEUS, 2003 777 I

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Fig. 1. A: Map showing the outcrop of the Sortehat Formation in the Jameson Land Basin; the outcrop pattern is aligned roughly north–south, parallel to the basin axis. B: Structural outline of the southern part of the East Greenland rift basin. The Jameson Land Basin was bounded by faults to the east, west and north, and by the Liverpool Land high towards the east. The nature of the south- ern boundary is unknown. The structural setting suggests an elongate semi-enclosed basin during deposition of the Sortehat Formation. Modified from Engkilde & Surlyk (1993).

in this paper follows the revised scheme of Dam & ular, this paper focuses on establishing the age of the Surlyk (1998). Although the and sequence Sortehat Formation, elucidation of the stratigraphic sig- stratigraphy of the sandy formations have been the sub- nificance of the formation boundaries and contributing ject of considerable study in recent years (Engkilde & to an understanding of the depositional history of the Surlyk 1993; Engkilde 1994; Dam & Surlyk 1995; Engkilde Sortehat Formation. & Surlyk 2003, this volume; Koppelhus & Dam 2003, this volume), the precise age, the nature of the bound- aries and the depositional history of the Sortehat Formation remain poorly understood. The aim of this Regional setting and stratigraphy paper, therefore, is to present the results of a detailed The Middle Jurassic Sortehat Formation is exposed in palynological study of these strata that was undertaken the Jameson Land Basin in the southernmost part of the in close co-operation with a sedimentological and north–south-trending, failed-rift system in East Greenland sequence stratigraphic study (Hansen 1999). In partic- (Fig. 1). Rifting was initiated in the late Palaeozoic, and

778 Upper

Super- Chronostratigraphy group Group Formation

Scoresby Sund Raukelv Volgian

Hall Hareelv Kimmeridgian Bredning

Oxfordian Olympen

Callovian Fossil- Thickness Formation Vardekløft bjerget (m) Bathonian S N urassic J Pelion Fm 100–400 Bajocian Pelion ? Sortehat Fm 60–110 Aalenian Sortehat Jameson Land ? Toarcian Ostreaelv Fm 290–420 Neill Klinter Ostreaelv

Sandstone Gule Horn Pliensbachian Sandstone/mudstone heterolith Rævekløft Lower Middle Mudstone Sinemurian Primulaelv Kap Stewart Hettangian Rhætelv

Rhaetian Innakajik

Fig. 2. Lithostratigraphy of the Jurassic of Jameson Land, East Greenland showing the stratigraphic position of the Sortehat Formation rias. in theT uppermost Neill Klinter Group. Modified from Surlyk (2003, this volume, fig. 5). The boundary between the Ostreaelv Formation and the Sortehat Formation is sharp and erosive in the northern proximal part of the basin and truncates shoreface deposits (indi- cated by the wavy line). At southern localities, the offshore mudstones of the Sortehat Formation gradationally overlie transgressive lower shoreface to offshore deposits. The Sortehat Formation thins towards the south. The heterolithic unit in the lowermost part of the formation also thins towards the south. The upper boundary of the Sortehat Formation is erosional (wavy line) and may record a minor hiatus.

from Late through Mesozoic times the basin land area of Jameson Land and Scoresby Land (Fig. 1). was characterised by relatively uniform thermal subsi- It was deposited in an elongate, semi-enclosed seaway, dence interrupted by periods of faulting (Surlyk et al. connected in the south with the epeiric Jurassic seaway 1981; Surlyk 1990a). The basin was bounded to the west between Greenland and Norway (Surlyk et al. 1981). by a major, approximately north–south-trending fault The Sortehat Formation, as adopted here, was orig- zone and to the east by faults and the elongated inally erected as the lower member (the Sortehat NNE–SSW-trending Liverpool Land high (Fig. 1; Surlyk Member) of the Vardekløft Formation (Surlyk et al. et al. 1981; Surlyk 1990a). To the north, the basin was 1973). It was raised to the status of formation by Surlyk bounded by a number of NW–SE-trending cross-faults (1990b, fig. 3); formal definition of the Sortehat Formation in Kong Oscar Fjord. The southern boundary is unknown as the uppermost formation of the Neill Klinter Group but the basin probably extended further south under the was undertaken by Dam & Surlyk (1998). The type present-day Scoresby Sund. The original extent of the locality of the Sortehat Formation at Sortehat (Fig. 1A) Sortehat Formation is not known in detail, but during is identical to that of the former Sortehat Member (Surlyk Middle Jurassic time it probably covered the present et al. 1973).

779 The Sortehat Formation overlies the sandstone-dom- developed coarse parallel lamination formed by an inated Ostreaelv Formation (Figs 2, 3) which records alternation of thin sand/siltstone layers and mudstone deposition within a shallow wave, storm and tidally layers. The mudstones contain debris and cal- influenced marine embayment (Dam & Surlyk 1995, careous concretions are present locally. Macrofossils 1998). The boundary with the overlying black mud- include belemnites and ostreid bivalves. stones of the Sortehat Formation is distinct at all local- Both mudstone facies are broadly interpreted as hav- ities but changes from a sharp ravinement surface in ing been deposited from suspension below wave base northern localities to a gradational drowning surface in in an offshore environment. The lack of bioturbation southern localities (Figs 3, 4; Hansen 1999). The Sortehat in the black mudstone facies suggests that the sea floor Formation is 60–100 m thick and is overlain by the was periodically inhospitable, probably due to poor sandy, marine Pelion Formation (Fig. 2); the boundary oxygenation. is sharp throughout the basin. Sedimentologically, the boundary between the Ostreaelv and Sortehat Formations represents a land- wards shift in facies and a rise in relative sea level. The Sandstone facies contact is interpreted as a transgressive erosional sur- The interbedded sandstone layers from the heterolithic face (Surlyk 1990a, b; Hansen & Surlyk 1994, Hansen levels are very fine- to medium-grained; they locally 1999) and marks a basinwide flooding event within an show well-developed wave ripples but more commonly overall transgressive period. The upper boundary of appear as lenses and streaks of sandstone, 2–15 mm the Sortehat Formation represents a seawards shift in thick, reflecting incipient ripple development (Fig. 3C; facies related to a fall in sea level and is interpreted as cf. the ‘incipient lenses’ of de Raaf et al. 1977). Laterally a marine erosional surface formed during forced regres- persistent sandstone layers, 5–30 cm thick, are present sion (Surlyk 1990a, b; Engkilde & Surlyk 1993). locally and show hummocky cross-stratification (Fig. 3C). The trace fossils Ophiomorpha nodusa and Pelecypod- ichnus amygdaloides occur in some of the sandstone layers. Facies and depositional setting The basal heterolithic unit records deposition in the The Sortehat Formation consists of dark grey to black upper offshore – offshore transition zone, influenced by mudstones with subordinate heterolithic levels (Figs 3, storm sand deposition on a muddy shelf. The persistent 4). The formation thins southwards away from the north- hummocky cross-stratified sand sheets represent higher ern basin margin, from 100 m at Enhjørningen Dal to energy storm events above storm wave base. The bio- 60 m at Albuen (Fig. 4). A discrete, heterolithic unit of turbation associated with the sandstones indicates well- interbedded mudstones, sandstones and siltstones, 20 m aerated bottom-water conditions, at least on occasion. thick, occurs at the base of the formation at the north- ern locality of Enhjørningen Dal; this unit thins to 8 m at the type locality of the Sortehat Formation and ulti- mately wedges out towards the south (Figs 3, 4). Depositional setting The preliminary interpretation is that the black to dark grey mudstones that dominate the formation were deposited from suspension. In combination with the thin Mudstone facies storm sand layers, the mudstones indicate deposition The mudstones of the Sortehat Formation are darker and in an offshore environment, probably a muddy, shal- more fissile in the lowermost part of the formation and low epeiric sea. The basal thick heterolithic unit at generally become lighter upwards, changing from black Enhjørningen Dal and Sortehat records northwards at the base to dark grey at the top. The dark grey mud- shoreface retreat prior to final drowning of the entire stones, which dominate the formation, commonly appear basin. A detailed sedimentological and sequence strati- structureless but locally show bioturbated fabrics, includ- graphic analysis of the Sortehat Formation was pre- ing subhorizontal traces such as Curvolithus isp. and sented by Hansen (1999). Planolites-like burrows and, near the top of the for- mation, some vertical traces referred to Diplocraterion isp. The black mudstones, in contrast, show a well-

780 A

B C

Fig. 3. A: The Sortehat Formation at the southernmost locality, Albuen, Neill Klinter. The Sortehat Formation consists mainly of dark grey to black mudstones deposited in an offshore environment. The boundary with the underlying fully marine sandy Ostreaelv Formation (arrow) is a particularly distinctive facies boundary within the Mesozoic succession of the East Greenland rift basin and represents a marine flooding event. The profile shown is c. 80 m thick. B: The lower, heterolithic unit of the Sortehat Formation at the northern locality of Enhjørningen Dal (basal boundary marked by arrow; c. 15 m of the Sortehat Formation illustrated). C: In contrast to the shaly appearance at the southern locality of Albuen (see Fig. 3A), this 20 m thick heterolithic unit at Enhjørningen Dal consists of stacked coarsening-upwards units (arrow); the example illustrated here passes up from mudstone (m) to hummocky cross-stratified sandstone (HCS). These stacked units probably record repeated slowing of transgression and shoreline progradation at the northern basin margin. Measuring rule (centre left) is 20 cm long.

781 S N Enhjørningen Dal

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Sandstone with Pebbles Mudstone clay laminae Logs

Mudstone with Cross-bedded Belemnites Concretions sand lenses sandstone Bivalves Hummocky Degree of bioturbation Sandstone cross-stratified sandstone Ammonites

Fig. 4. Correlation between the three localities Albuen, Sortehat and Enhjørningen Dal showing the palynological zonation (Assemblage Zones 6–9). The upper part of the Sortehat Formation and the overlying Pelion Formation at the type locality (Sortehat) have been removed by recent erosion. The base and top of the Sortehat Formation are indicated by the dotted line.

the basal beds of the Pelion Formation, overlying the Previous work Sortehat Formation, indicating an age not younger than The age of the Sortehat Formation is not well known. early Late Boreal Bajocian for these beds (Surlyk et al. Belemnites have been collected from the formation but 1973; Callomon 1993). to date have not been systematically identified. According The dark shales of the Sortehat Formation have been to Dam & Surlyk (1998), belemnites from the upper lev- subjected to three palynological studies and a geo- els of the underlying Ostreaelv Formation were col- chemical study (Fensome 1979; Lund & Pedersen 1985; lected by Rosenkrantz (1934) and studied by Doyle Krabbe et al. 1994; Underhill & Partington 1994) although (1991); the belemnite ‘Parabrachybelus’ subadunca- none of these studies were based on a comprehensive, tus from this level probably has a range restricted to closely-spaced sampling programme. Three samples the latest Toarcian Levesquei Zone (Doyle 1991). The from the Sortehat Formation were analysed for dinofla- ammonite Cranocephalites borealis has been found in gellate cysts by Fensome (1979). One sample (144112)

782 was from the type section at Sortehat, and two (144229 The organic geochemistry and the palynofacies of the and 144231) were from a locality north of Dusén Bjerg. Sortehat Formation were discussed by Krabbe et al. Sample 144112 yielded a sparse assemblage of well-pre- (1994). Based on the palynofacies study, the succession served palynomorphs, including Nannoceratopsis gra- was divided into three facies: (1) Botryococcus-domi- cilis. Sample 144229 yielded a dinoflagellate cyst nated, (2) spore/pollen and brown/blackwood and (3) assemblage dominated by N. gracilis. The assemblage blackwood-dominated, few spore/pollen. These results, in sample 144231 was dominated by the acritarch together with the geochemical data, suggest an increase Veryhachium sortehatense. Fensome (1979) concluded in the salinity of the depositional environment with that the palynomorph assemblages determined from time (Krabbe et al. 1994). the three samples from the Sortehat Formation did not allow for accurate dating. It is worth noting that the dinoflagellate cyst Sentusidinium pelionense was not found in any of the three samples from the Sortehat Materials and methods Formation, but was common in one sample (144111) This study is primarily based on material collected by from the overlying Pelion Formation. the authors in the 1993 and 1994 field seasons. The Lund & Pedersen (1985) presented the results of a Sortehat Formation was investigated along a north–south- palynological study concerning the Neill Klinter and trending profile through the Jameson Land Basin, par- Vardekløft Groups and the lower part of the Hareelv allel to the basin axis. Sections were sampled and studied Formation. Four samples (142832–35) are from the at four localities: Albuen (Neill Klinter along Hurry Inlet) Sortehat Formation. These samples yielded abundant and Sortehat (the type locality of the formation) in the Sentusidinium pelionense but Nannoceratopsis gracilis south and Enhjørningen Dal and Pelion in the north- was not found, whereas the pollen Perinopollenites ela- ern part of Jameson Land (Fig. 1; Appendix 1). The toides was abundant. Sample 142833 had the lowest boundary between the Ostreaelv and the Sortehat number of marine cysts, whereas Botryococcus was Formations is a well-defined stratigraphic surface and common in 142832 and 142835. On the basis of these was used as a datum for the sections measured at out- data, Lund & Pedersen (1985) suggested a Middle–Late crop; the structural dip of the succession is negligible. Bajocian age for the Sortehat Formation. The outcrop sections were measured by Jacob staff in The Neill Klinter and Vardekløft Groups were also metres relative to this surface. Altimeter readings for the studied palynologically by Underhill & Partington (1994). datum surface are listed in Appendix 1; sample loca- The material on which their study was based was sam- tions are thus referred to height above sea level, being pled at Liaselv (their section 1), Vardekløft (section 2), the sum of the datum altitude (measured by altimeter) and Harris Fjeld/Primulaelv at Neill Klinter, the west- and the measured section thickness above the datum. ern slope of Hurry Inlet (section 3; Fig. 1). Twenty sam- Samples taken from core from the borehole at the type ples from the Sortehat Formation were analysed from locality of the Sortehat Formation are related to an arbi- their sections 1 and 2 (7 samples from section 1, 13 sam- trary datum (base of cored section) within the upper ples from section 2; their fig. 10). An Aalenian–Bajocian Ostreaelv Formation (Appendix 1). age was proposed for the Sortehat Formation. On the Approximately 300 samples were processed at the basis of these data, Underhill & Partington (1994) sug- palynological laboratory of the former Geological Survey gested that the Aalenian–Bajocian record was essentially of Greenland using standard techniques (Nøhr-Hansen complete, without apparent biostratigraphic or sedi- 1993). The palynomorphs were studied using a trans- mentological evidence of the ‘mid-Cimmerian event’ mitted light microscope. For each sample, 200 speci- known from the North Sea area. Underhill & Partington mens were counted, and all species were registered in (1994) concluded that the boundary between the the range chart program SIS. The palynomorphs illus- Sortehat Formation and the underlying Ostreaelv trated in Plates 1–6 are from the borehole at the type Formation does not represent an important uncomfor- locality of the Sortehat Formation and from a section mity. This is confirmed by recent work (Koppelhus & at Lepidopteriselv; the latter has not been used in the Dam 2003, this volume) on the uppermost part of the correlation between the three other localities. All the Ostreaelv Formation which has been referred to the palynomorph taxa recorded in the samples are listed Late Toarcian – early Aalenian on the basis of the paly- in Appendix 2 with author attributions and dates, and nological assemblages. the slides are stored in the collections of the Geological Survey of Denmark and Greenland.

783 Base. The base of the assemblage is placed at the first Palynological zones: definition sample in which Botryococcus spp. overwhelmingly The data presented here form part of a broader study dominates the assemblage and the dinoflagellate cysts encompassing the entire Neill Klinter Group. Nine paly- Nannoceratopsis gracilis and Nannoceratopsis senex nological assemblage zones (1–9) have been recog- and dinoflagellate cysts in general become rare. In the nised in the group; assemblage zones 1–6 from the Albuen section (Fig. 6), this event coincides with the Rævekløft, Gulehorn and Ostreaelv Formations are pre- first co-occurrence of Callialasporites dampieri (pollen) sented in a companion paper (Koppelhus & Dam 2003, and Mendicodinium groenlandicum (dinoflagellate this volume). The three assemblage zones of the Sortehat cyst) although in other sections (e.g. Sortehat, Enhjør- Formation, based on the occurrence of miospores, ningen Dal; Figs 5, 7) these species first occur together dinoflagellate cysts and freshwater algae, are named some metres below the Botryococcus spp. influx. from below: (7) the Botryococcus Assemblage Zone, (8) the Nannoceratopsis gracilis – Nannoceratopsis senex Top. The upper boundary is defined by the uppermost Assemblage Zone and (9) the Sentusidinium pelionense sample showing the Botryococcus-dominated assem- Assemblage Zone. Assemblage Zone 7 is also recognised blage. Above this level, Botryococcus spp. are scarce in the upper levels of the Ostreaelv Formation and is and Nannoceratopsis gracilis and N. senex become described briefly in Koppelhus & Dam (2003, this vol- abundant once more. ume), but is defined herein. Characteristics. The assemblage is characterised by the overwhelming dominance of the freshwater alga Botry- Botryococcus ococcus spp. and the scarcity of dinoflagellates. Pollen Assemblage Zone 7: species such as Perinopollenites elatoides, Cerebro- new assemblage zone pollenites macroverrucosus and bisaccate pollen are also abundant. Occurrence. Albuen 438.5–443.5 m Enhjørningen Dal 425.35–445 m Suggested age. Aalenian (see discussion below). Pelion 550–567 m Sortehat (core) 27.82–36.36 m Palaeoenvironment. The palynomorph assemblage indi- cates a brackish marine environment. The abundant In the cored section from Sortehat, this assemblage Botryococcus and the common spores and pollen reflect occurs in the lower levels of the Sortehat Formation, a significant allochthonous terrestrial input related to the base being immediately above the lower boundary transgression (see later discussion). of the formation (Figs 4, 5). At Albuen, the assemblage is represented in the uppermost few metres of the Remarks. The Botryococcus Assemblage is, as the name Ostreaelv Formation and extends up into the Sortehat indicates, dominated by Botryococcus spp. (Plate 4, Formation (Figs 4, 6; Fig. 6 faces page 794). At Enhjør- fig. 4), but spores and pollen also play an important ningen Dal, Assemblage Zone 7 is restricted to the lower role. The spore flora is diverse, but there are only few Sortehat Formation although here the base is some 6 m specimens of each species, whereas pollen species are above the lower boundary of the Sortehat Formation (Figs less diverse but occur abundantly, such as Perino- 4, 7; Fig. 7 faces page 795). Assemblage Zone 7 occurs pollenites elatoides (Plate 2, fig. 5), bisaccate pollen, within the lower levels of sequence SQ7 of Dam & Cerebropollenites macroverrucosus (Plate 3, fig. 3) and Surlyk (1995, 1998). Corollina torosus (Plate 3, fig. 2). Some of the less abun- dant pollen are stratigraphically significant, such as Reference section. Sortehat (core), 27.82 m (sample Quadraeculina anellaeformis (Plate 3, fig. 5) and 303143-73) – 36.36 m (sample 303143-62; Figs 4, 5). Callialasporites dampieri (Plate 2, fig. 1). Acritarchs and dinoflagellate cysts occur only rarely. Among the latter Additional sections. Albuen, 438.5 m (sample 397452) are Dissilodinium sp. (Plate 6, fig. 4), Mendicodinium – 443.5 m (sample 397468; Figs 4, 6). Enhjørningen sp., Pareodinia halosa (Plate 4, fig. 6), Mancodinium Dal, 425.35 m (sample 398341) – 445 m (sample 398417; semitabulatum (Plate 6, fig. 1) and a few specimens of Figs 4, 7). Nannoceratopsis senex (Plate 5, fig. 2) and N. gracilis (Plate 5, fig. 1).

784 spp. yhachium formosum yhachium sortehatense r r allodinium laganum ipartina variabilis esicaspora fuscus esicaspora asmanites aurocusporites verrucatus T T Tr Uvaesporites puzzlei Ve Ve V W 3

79 12 11 74 75 38 51

spp. Foraminifera 80

spp. 79 asmanites asmanites T

spp. 78 Botryococcus Botryococcus

spp. 77 rare Leiosphaeridia Leiosphaeridia

76

Lecaniella foveata Lecaniella spp.

75 spp. yhachium sortehatense yhachium r Ve

spp.

74 yhachium formosum yhachium r Ve Uncertain determination Ver y Rare Few Common Abundant

73 Limbicysta bjaerkei Limbicysta

72 Acritarch spp. Acritarch 72 ?

R

spp. 71 Scriniocassis Scriniocassis

?

70

Phallocysta elongata Phallocysta Scriniocassis Scriniocassis weberii Sentusidinium pelionense Sestrosporites pseudoalveolatus Spheripollenites subgranulatus Staplinisporites caminus Striatella jurassica Striatella parva Striatella Susadinium scrofoides Retitriletes semimuris Retitriletes

? 69

Andreedinium arcticum Andreedinium 4 5 8

68 Phallocysta eumekes Phallocysta 71 53 65 14 32 10 18 26 67

67 Susadinium scrofoides Susadinium

66 Nannoceratopsis plegas Nannoceratopsis

65 Sentusidinium pelionense Sentusidinium

64 eodinia ceratophora eodinia r Pa

spp. 63 Kallosphaeridium Kallosphaeridium

spp. 62 Mendicodinium Mendicodinium

61 Mendicodinium reticulatum Mendicodinium

60 eodinia halosa eodinia r Pa

spp. spp. 59 Dissiliodinium Dissiliodinium

58 Mendicodinium groenlandicum Mendicodinium

57 Dinocyst spp. Dinocyst 57

56 Mancodinium semitabulatum Mancodinium

55 Nannoceratopsis senex Nannoceratopsis

54

Nannoceratopsis ambonis Nannoceratopsis inopollenites elatoides eodinia halosa eodinia ceratophora

r r r

53 olycingulatisporit triangularis Scriniocassis weberii Scriniocassis

Phallocysta elongata Phallocysta eumekes Pinuspollenites minimus P anellaeformis Quadraeculina Retitriletes austroclavatoides Retitriletes clavatoides Pe Pa Pa Nannoceratopsis senex Nannoceratopsis Neoraistrickia

52 Nannoceratopsis gracilis Nannoceratopsis

51 ormation. 6 allodinium laganum allodinium W

70 68 35 23 39 27 34 60 64 55

50 25 Dodekovia tegilla Dodekovia

49 Monosaccate spp. Monosaccate 49

48 Eucommiidites troedsonii Eucommiidites

47 Chasmatosporites apertus Chasmatosporites

46 Araucariacites australis Araucariacites

45 Callialasporites trilobatus Callialasporites

44 Exesipollenites tumulus Exesipollenites

43 Callialasporites turbatus Callialasporites

?

42

Callialasporites dampieri Callialasporites spp.

41 Chasmatosporites major Chasmatosporites

40 Cerebropollenties thiergartii Cerebropollenties

39 Quadraeculina anellaeformis Quadraeculina

38 esicaspora fuscus esicaspora V

37 Corollina torosus Corollina

36 Cerebropollenites macroverrucosus Cerebropollenites

35 Pinuspollenites minimus Pinuspollenites

Nannoceratopsis ambonis Nannoceratopsis gracilis Nannoceratopsis plegas Nannoceratopsis Limbicysta bjaerkei Lycopodiacidites rugulatus Mancodinium semitabulatum Manumia delcourtii Mendicodinium groenlandicum Mendicodinium reticulatum Mendicodinium 34

inopollenites elatoides inopollenites r Pe 33 Bisaccate spp. Bisaccate 33 2

4954 Monosaccate spp. 52 66 73 19 56 2458 spp. Megaspore 61 62

32 Spheripollenites subgranulatus Spheripollenites

31 Chasmatosporites hians Chasmatosporites

R

spp. 30 Densosporites Densosporites

spp. 29 Apiculatisporites Apiculatisporites

28 Kraeuselisporites reissingeri Kraeuselisporites

27 Retitriletes austroclavatoides Retitriletes

spp. 26 spp. Striatella Striatella

spp. 25

Neoraistrickia Neoraistrickia spp.

spp.

spp. 24 Megaspore spp. Megaspore 24

23 olycingulatisporit triangularis olycingulatisporit P

22 Densoisporites velatus Densoisporites

21 minisporis jurassicus minisporis ra Fo

20

Densoisporites scanicus Densoisporites

aminisporis jurassicus ?

19 Lycopodiacidites rugulatus Lycopodiacidites r

18

Striatella parva Striatella Leptolepidites bossus Leptolepidites Ischyosporites variegatus Kallosphaeridium Kraeuselisporites reissingeri Lecaniella foveata Leiosphaeridia Ischyosporites Eucommiidites troedsonii tumulus Exesipollenites Fo

17

Ischyosporites variegatus Ischyosporites

9

spp. 16

Leptolepidites Leptolepidites 16 17 63 28 76 77 15 48 44 8021 spp. Foraminifera

spp. 15 Ischyosporites Ischyosporites

14 Sestrosporites pseudoalveolatus Sestrosporites

spp. 13 Concavissimisporites Concavissimisporites

12

aurocusporites verrucatus aurocusporites

T spp.

11 Uvaesporites puzzlei Uvaesporites

10 Striatella jurassica Striatella

9

Leptolepidites bossus Leptolepidites spp. spp.

spp.

spp. 8 Retitriletes Retitriletes

spp. 7 Deltoidospora Deltoidospora

6 Retitriletes clavatoides Retitriletes

5 Retitriletes semimuris Retitriletes

4 Staplinisporites caminus Staplinisporites

3 ipartina variabilis ipartina Tr

2 Manumia delcourtii Manumia torosus Corollina Deltoidospora Densoisporites scanicus Densoisporites velatus Chasmatosporites apertus Chasmatosporites hians Chasmatosporites major Concavissimisporites Densosporites Dissiliodinium Dodekovia tegilla Dodekovia

spp. 1 Baculatisporites Baculatisporites 7 47 31 41 13 37 20 22 30 57 Dinocyst spp. 59 50 0 4 8

-2 -2 -2

3 3 3 Sample number Sample 4 4 4 1 1 1 3 3 3

0 0 0

303143-11 303143-16 3 3 3 303143-33 303143-37 303143-43 303145-49 303143-55 303143-62 303143-67 303143-69 303143-73 303143-75 303143-77 Sample height Sample

77.98 73.13 69.50 65.24 61.31 57.41 53.86 49.48 45.09 40.11 36.36 32.06 29.56 27.82 26.28 12.65 (m) spp. spp.

75 50 25

spp. Palynological Assembl. Zones Assembl. Palynological

9 8 7 6

Lithostratigraphy Ostreaelv Formation Sortehat Formation Sortehat Formation Ostreaelv

Stage Aalenian oarcian T

Botryococcus Baculatisporites Callialasporites dampieri Callialasporites trilobatus Callialasporites turbatus macroverrucosus Cerebropollenites thiergartiiCerebropollenties Andreedinium arcticum Andreedinium Apiculatisporites Araucariacites australis System Middle Jurassic Middle Jurassic Lower 1 Sortehat 33 Bisaccate spp. 78 42 45 43 36 46 40 72 spp. Acritarch 69 29 Alphabetical species list Fig. 5. Chart showing the distribution of palynomorphs from cored borehole at Sortehat, type locality Sortehat F

785 Assemblage Zone 8: Nannoceratopsis gracilis – Assemblage Zone 9: Sentusidinium pelionense Nannoceratopsis senex new assemblage zone new assemblage zone Occurrence. Albuen 465–502 m Occurrence. Albuen 444.5–460 m Enhjørningen Dal 502–516.75 m Enhjørningen Dal 446–500 m Sortehat (core) 57.41–77.98 m Pelion 573–577 m Sortehat (core) 40.11–53.86 m Assemblage Zone 9 occurs in the upper Sortehat Formation in the Sortehat section; note that the sam- Assemblage Zone 8 occurs within the Sortehat For- pled interval in this section is restricted to the Sortehat mation; it thus falls within sequence SQ7 of Dam & Formation (Figs 4, 5). At Albuen and Enhjørningen Dal, Surlyk (1995, 1998). the assemblage extends through the upper Sortehat Formation and persists up into the overlying Pelion Reference section. Sortehat (core), 40.11 m (sample Formation (Figs 4, 6, 7). Assemblage Zone 9 is thus 303143-55) – 53.86 m (sample 303143-37; Figs 4, 5). characteristic of the upper levels of sequence SQ7 of Dam & Surlyk (1995, 1998), and at Albuen and Additional sections. Albuen, 444.5 m (sample 397469) Enhjørningen Dal spans the sequence boundary at the – 460 m (sample 397474; Figs 4, 6). Enhjørningen Dal, base of the Pelion Formation and extends into sequence 446 m (sample 395625) – 500 m (sample 398442; Figs P1 of Engkilde & Surlyk (2003, this volume). 4, 7). Reference section. Sortehat (core), 57.41 m (sample Base. The lower boundary is placed at the first sample 303143-33) – 77.98 m (sample 303143-11; Figs 4, 5). in which Botryococcus spp. is rare and Nannoceratopsis gracilis and N. senex are abundant. Additional sections. Albuen, 465 m (sample 397475) – 502 m (sample 397498; Figs 4, 6). Enhjørningen Dal, Top. This is defined by the last sample showing the 502 m (sample 395679) – 516.75 m (sample 398448; Figs characteristic assemblage (see below), above which 4, 7). level Nannoceratopsis gracilis becomes less common and Sentusidinium pelionense is the most common Base. The base of the zone is placed at the sample in dinoflagellate cyst. which Sentusidinium pelionense is the most common dinoflagellate cyst; Nannoceratopsis gracilis and N. Characteristics. Botryococcus spp. is rare in this assem- senex are absent or rare. blage whereas Nannoceratopsis gracilis is abundant and there is a general increase in diversity and abundance Top. The upper boundary of the assemblage zone is not of dinoflagellate cysts relative to the underlying zone. defined here. The Sentusidinium pelionense Assemblage Zone extends from the upper part of the Sortehat For- Suggested age. Aalenian (see discussion below). mation into the lowermost beds of the Pelion Formation. The full extent of the zone within the Pelion Formation Palaeoenvironment. The palynological data indicate a is not known. It is likely, however, that the top of the brackish marine environment. zone occurs within the lower levels of the Pelion For- mation (S. Piasecki, personal communication 1997). Remarks. Dinoflagellate cysts are abundant, whereas Botryococcus spp. becomes rare at 40.11 m in the type Characteristics. Sentusidinium pelionense is abundant. section at Sortehat (Fig. 5). There are a few acritarchs, In some of the investigated sections, S. pelionense is such as Veryhachium sorthatense (Plate 4, fig. 2), and accompanied by Nannoceratopsis gracilis in samples at some of the same spores and pollen as observed in the the boundary between the two palynomorph assemblage Botryococcus Assemblage Zone (Plates 1, 2). zones. In other sections, they do not overlap, i.e. N. gra- cilis is replaced by S. pelionense up-section.

786 Suggested age. Aalenian – ?Early Bajocian (see discus- Botryococcus appears and the acme extends for approx- sion below). imately 9 m before Botryococcus disappears and the dinoflagellate cyst Nannoceratopsis gracilis begins to Palaeoenvironment. The palynological data indicate dominate the assemblage. In three samples (303143-75, that most of the organic material came from a brackish 303143-73 and 303143-69) an unidentified dinocyst marine source. (Dinocyst sp.) is common to abundant; this dinoflagel- late cyst is similar to one described as Dinoflagellate sp. Remarks. The assemblage contains fewer spores and indet. 2 from Callovian deposits from Spitsbergen where pollen than the underlying assemblage zone. The most it is said to occur in profusion in some assemblages abundant dinoflagellate cysts are Sentusidinium pelion- (Bjærke 1980). This species was also common in the low- ense (Plate 6, fig. 5) and Pareodinia halosa (Plate 4, ermost sample from the Pelion locality (Fig. 8). fig. 6) although a few specimens of Phallocysta eumekes N. gracilis continues to be the dominant dinoflagel- were recorded. The acritarch Limbicysta bjaerkei was late for 14 m to 53.86 m (sample 303143-37) above also found. which there is an acme of Sentusidinium pelionense together with abundant Pareodinia halosa.

Palynological results Albuen Sortehat The Sortehat Formation at Albuen covers about 60 m At the type locality, the Sortehat Formation is a mini- and 41 samples have been investigated from this inter- mum of 50 m thick; the top of the formation is not seen val (Fig. 6). The boundary between the Ostreaelv and due to recent erosion. Seventy samples were collected Sortehat Formations is at 440 m, but the palynomorph and seventeen of these are shown on the distribution assemblages change between sample 405449 at 434 m chart (Fig. 5). and sample 397452 at 438.6 m, several metres below The palynological assemblages from the entire sec- the top of the Ostreaelv Formation, where Botryococcus tion fall into three distinct assemblages. There is a dis- becomes common, dinoflagellate cysts become rare tinct change from the Perinopollenites elatoides Zone (Mendicodinium groenlandicum, Nannoceratopsis gra- (Assemblage Zone 6, described by Koppelhus & Dam cilis and Nannoceratopsis senex are present but rare), 2003, this volume) in the underlying Ostreaelv Formation and pollen are more common than spores. At 443.5 m to the Botryococcus Assemblage Zone (Assemblage in sample 397468, 3.5 m above the base of the Sortehat Zone 7), which is overwhelmingly dominated by the Formation, Botryococcus is abundant for the last time; freshwater alga Botryococcus. This change is first above this level N. gracilis and N. senex become abun- observed at 27.82 m (sample 303143-73) just above the dant and are accompanied by Sentusidinium pelionense. lithological boundary between the Sortehat and Ostreaelv S. pelionense is only common in one sample (397471), Formations at 27.6 m. The Perinopollenites elatoides and Pareodinia halosa is common in two other sam- Zone is characterised by the first appearance of Stap- ples, 397475 and 347477. This assemblage changes linisporites caminus, Sestrosporites pseudoalveolatus, between 460 m and 465 m, above which level S. pelion- Phallocysta eumekes and Wallodinium laganum; ense is the only common dinoflagellate; this species Callialasporites dampieri makes its first appearance disappears above 496 m, within the lower levels of the near the top of the zone. The data from the three Pelion Formation. The overall dinoflagellate cyst diver- Ostreaelv Formation samples from the Sortehat section sity decreases from the upper part of the Nannoceratopsis do not show exactly the same pattern although S. cami- gracilis – Nannoceratopsis senex Assemblage Zone nus and C. dampieri have their first appearance in the through the Sentusidinium pelionense Assemblage Zone uppermost sample from the Ostreaelv Formation and in the upper part of the Sortehat Formation. A few the dinoflagellate cyst W. laganum is very common in dinoflagellate cysts have been found in most of the sample 303143-80 at 6.64 m (not shown on Fig. 5), samples, and pollen are more common than spores. The approximately 20 m below the boundary between the samples that span the boundary between the Sortehat Ostreaelv and Sortehat Formations. Formation and the overlying Pelion Formation yielded In sample 303143-73 at 27.82 m, 0.22 m above the an impoverished palynomorph assemblage without any base of the Sortehat Formation, the freshwater alga age-diagnostic species.

787 gellate cysts are present and locally common (Nanno- Enhjørningen Dal ceratopsis gracilis, Mendicodinium groenlandicum and Enhjørningen Dal is a composite section spanning the Mendicodinium sp.). The two uppermost samples (sam- top of the Ostreaelv Formation, the entire Sortehat ple 339710 at 573 m and sample 339711 at 577 m) Formation and the lowermost part of the Pelion yielded an assemblage rich in dinoflagellate cysts includ- Formation (Fig. 7). The boundary between the Ostreaelv ing Nannoceratopsis gracilis, Nannoceratopsis triceras, and Sortehat Formations is at 418 m; the first change Nannoceratopsis plegas, Nannoceratopsis triangulata in the palynomorph assemblages is observed in sam- and Mancodinium semitabulatus and some acritarchs. ple 398341 at 425.35 m. This lower interval (belonging Spores and pollen are common to abundant through- to the uppermost part of the Ostreaelv Formation and out the 27 m section. the lowermost part of the Sortehat Formation) is com- pletely dominated by the dinoflagellate species Nanno- ceratopsis gracilis and Nannoceratopsis senex, and by the pollen Perinopollenites elatoides. This interval is Discussion accordingly referred to the Perinopollenites elatoides The palynological study reported here has implications Assemblage Zone (Assemblage Zone 6; Koppelhus & both for the age and regional correlation of the Sortehat Dam 2003, this volume) which is of Late Toarcian – early Formation and for the environmental understanding of Aalenian age. At 425.35 m (sample 398341), Botryococcus the formation, in association with the detailed sedi- appears in abundance and persists in large numbers up mentological and sequence stratigraphic study (Hansen to sample 398417 at 445 m. From sample 395625 at 1999). 446 m to sample 398442 at 500 m, the palynomorph assemblage contains few Botryococcus; N. gracilis and N. senex are very common in the lower part of this interval. In the upper part of the section, however, from Age and correlation sample 395657 at 476 m to the uppermost sample On the basis of macrofauna in the underlying and over- (398448) at 516.75 m, N. gracilis and N. senex are absent lying formations (see previous discussion), the age of whereas other dinoflagellates are present but rare (e.g. the Sortehat Formation is constrained between the lat- Mancodinium semitabulatum, Phallocysta eumekes, est Toarcian and the mid-Bajocian; previous palyno- Susadinium scrofoides, Parvocysta barbata, Dissilo- logical work has indicated an Aalenian–Bajocian dinium sp. and Mendicodinium sp.). Sentusidinium (Underhill & Partington 1994) or a Middle–Late Bajocian pelionense is abundant in samples 395679 and 395671, age (Lund & Pedersen 1985). With a view to a more Pareodinia halosa is common in sample 395671 and well-founded understanding of the stratigraphy of the Kallosphaeridium sp. is abundant in the uppermost Sortehat Formation, the results of this study are com- sample (398448). Among the pollen at this level, bisac- pared with published palynological data from the North cates and Cerebropollenites macroverrucosus are espe- Atlantic region. Although palynological assemblages cially abundant. Trilete spores seem to decrease in from the Toarcian–Bajocian interval have been widely diversity and density compared to the lower part of the reported from this region, few of the sections have section. been independently dated using ammonites and the Aalenian assemblages, in particular, are commonly dom- inated by terrestrial material. The Aalenian stage is named after Aalen in Germany, Pelion where the lowest part of the ‘Braunjura’ crops out at Ten samples were investigated from the Pelion local- the northern edge of the Swabian Alps. Although no ity (Fig. 8). They cover the lowermost 27 m of the palynological papers have been published on material Sortehat Formation, which is approximately 110 m thick from the type locality, a dinoflagellate cyst assemblage at this locality. Generally, the palynomorph assemblages has been described from two cores from Hausen in are rich but the preservation is poor. The assemblage south-western Germany, 40 km from the type locality, from the lowermost 17 m of the formation (sample and the lithological units have been correlated with the 339702 at 550 m to sample 339709 at 567 m) is domi- ammonite zonation from the Eastern Swabian Alb (Feist- nated by Botryococcus. In sample 339709, Botryococcus Burkhardt 1990). The palynological assemblages are has its last abundant appearance. In this interval, dinofla- rich in spores and pollen, and only 5–20% of the total

788

66 Fungal spp. Fungal 66

65 Miscellaneous spp. Miscellaneous 65

spp. 64 Botryococcus Botryococcus

63 Acritarch spp. Acritarch 63

spp. 62 Leiosphaeridia Leiosphaeridia

61 Nannoceratopsis triangulata Nannoceratopsis

60 Mancodinium semitabulatum Mancodinium

59 Nannoceratopsis triceras Nannoceratopsis

58 Nannoceratopsis plegas Nannoceratopsis

spp. 57 Mendicodinium Mendicodinium

56 Nannoceratopsis senex Nannoceratopsis

55 Mendicodinium groenlandicum Mendicodinium

54 Nannoceratopsis gracilis Nannoceratopsis

53 Dinocyst spp. Dinocyst 53

spp. 52 Nannoceratopsis Nannoceratopsis

spp. 51 Phallocysta Phallocysta

50 Araucariacites australis Araucariacites

49 Callialasporites segmentatus Callialasporites

48 Callialasporites trilobatus Callialasporites

47 Cerebropollenties thiergartii Cerebropollenties

spp. 46 Monosulcites Monosulcites

45 Chasmatosporites hians Chasmatosporites

spp. 44 Striate Striate

43 Chasmatosporites apertus Chasmatosporites

42 Alisporites robusta Alisporites

41 Chasmatosporites major Chasmatosporites

40 Exesipollenites tumulus Exesipollenites

39 Spheripollenites psilatus Spheripollenites

38 Callialasporites dampieri Callialasporites

37 Vitreisporites pallidus Vitreisporites

36 Callialasporites minus Callialasporites

35 Quadraeculina anellaeformis Quadraeculina

34 Corollina torosus Corollina

spp. 33 Pinuspollenites minimus Pinuspollenites

spp.

32

inopollenites elatoides inopollenites r Pe

spp. 31 Bisaccate spp. Bisaccate 31

spp.

30 Cerebropollenites macroverrucosus Cerebropollenites

29 Callialasporites turbatus Callialasporites

ipartina variabilis spp. 28

Callialasporites Callialasporites odisporites major aurocusporites segmentatus

Retitriletes austroclavatoides Retitriletes clavatoides Retitriletes semimuris Retitriletes Rogalskaisporites cicatricosus Sestrosporites pseudoalveolatus Spheripollenites psilatus Spheripollenites subgranulatus Staplinisporites caminus Stereisporites stereoides Striate Striatella jurassica Striatella parva Striatella seebergensis Striatella T T Tr Uvaesporites Vitreisporites pallidus 27 Spheripollenites subgranulatus Spheripollenites

6 7 3 2 spp. 26

Uvaesporites Uvaesporites 17 14 10 39 27 13 16 44 23 11 20 25 15 22 26 37

25 aurocusporites segmentatus aurocusporites T

24 Leptolepidites bossus Leptolepidites

23 Striatella jurassica Striatella

22 ipartina variabilis ipartina Tr

21 Densoisporites scanicus Densoisporites

20 Striatella seebergensis Striatella

19 olycingulatisporit triangularis olycingulatisporit P

spp. 18 Neoraistrickia Neoraistrickia

17 Retitriletes austroclavatoides Retitriletes

16 Stereisporites stereoides Stereisporites

15 odisporites major odisporites T

14 Rogalskaisporites cicatricosus Rogalskaisporites

spp. 13 Staplinisporites caminus Staplinisporites spp.

spp.

12 spp. Manumia delcourtii Manumia spp.

spp.

11

Striatella parva Striatella spp.

10 Sestrosporites pseudoalveolatus Sestrosporites

9 Ischyosporites variegatus Ischyosporites

spp. 8 Leptolepidites Leptolepidites

7

Retitriletes semimuris Retitriletes inopollenites elatoides

r

6 ekryphalospora distincta

olycingulatisporit triangularis Retitriletes clavatoides Retitriletes

Ischyosporites variegatus K Leiosphaeridia Leptolepidites bossus Leptolepidites Pe Nannoceratopsis gracilis Nannoceratopsis plegas Nannoceratopsis senex Nannoceratopsis Nannoceratopsis triangulata Nannoceratopsis triceras Nannoceratopsis Neoraistrickia Phallocysta Pinuspollenites minimus P anellaeformis Quadraeculina Mancodinium semitabulatum Manumia delcourtii Mendicodinium groenlandicum Mendicodinium Monosulcites

spp. 5

Baculatisporites Baculatisporites

9 1 8

spp. 4 62 24 32 54 58 56 52 61 59 18 51 33 19 35 65 Miscellaneous spp. 60 12 55 57 46 Deltoidospora Deltoidospora

spp. 3 Retitriletes Retitriletes

spp. 2 Striatella Striatella

1 ekryphalospora distincta ekryphalospora K Sample number Sample

339711 339710 339709 339708 339707 339706 339705 339704 339703 339702 Sample height Sample

577.00 573.00 567.00 565.00 563.00 559.00 557.00 556.00 555.00 550.00 (m) spp.

575 565 555 spp.

spp. Palynological. Assembl. Zones Assembl. Palynological. spp.

8 7

n o i t a Lithostratigraphy m r o F

t a h e t r o S

Stage n a i n e l a A

Middle Jurassic Middle System Alisporites robusta Araucariacites australis Baculatisporites Botryococcus Callialasporites dampieri Callialasporites minus Callialasporites segmentatus Callialasporites Callialasporites trilobatus Callialasporites turbatus macroverrucosus Cerebropollenites thiergartiiCerebropollenties Chasmatosporites apertus Chasmatosporites hians Chasmatosporites major torosus Corollina Deltoidospora Densoisporites scanicus tumulus Exesipollenites elion 5 4 Fig. 8. Chart showing the distribution of palynomorphs from locality at Pelion. P Alphabetical species list 63 spp. Acritarch 42 50 31 Bisaccate spp. 64 38 36 49 28 48 29 30 47 43 45 41 34 21 53 Dinocyst spp. 40 66 Fungal spp.

789 palynomorph content is microplankton. In the Sortehat The first dinoflagellate cyst zonation of the Jurassic Formation, where the assemblages are also rich in of the Canadian Arctic was made by Johnson & Hills spores and pollen, microplankton form 25–40% of the (1973). Their Nannoceratopsis gracilis Range Zone cov- total palynomorph content. In the material from south- ers the Toarcian–Bajocian. They recorded the ammonite western Germany, twelve of the eighteen dinoflagel- Leioceras opalinum, but the only dinoflagellate cyst in late cysts recognised have also been identified in the common with the East Greenland material is Nanno- material from the Sortehat Formation. However, none ceratopsis gracilis. Bujak & Williams (1977) erected the of the four stratigraphically significant species for the Nannoceratopsis gracilis Zone for the Pliensbachian– Aalenian/Bajocian boundary from Hausen, Carpatho- Aalenian and the Mancodinium semitabulatum Zone dinium sp., Dissilodinium giganteum, Dissilodinium for the Bajocian, from successions offshore south-east- sp. A, and Durotrigia daveyi, have with certainty been ern Canada. Several species are common to the assem- found in the material from the Sortehat Formation. blages from the Sortehat Formation and from the The closest succession to Jameson Land with known Canadian Arctic but the age has not been confirmed by palynological data of Aalenian age is the Stø Formation any marine micro- or macrofauna. Davies (1983) estab- (Unit C) from the Møre Basin, offshore mid-Norway lished eight zones covering the Upper Pliensbachian – (Smelror et al. 1994). Ammonites have not been found Callovian, also from the Canadian Arctic, of which the to confirm the age, but a few foraminifera of little strati- Dapcodinium coalitum – Phallocysta eumekes Zone, of graphic value have been used, together with the pres- Late Toarcian – Early Bajocian age, contains species in ence of the dinoflagellate cyst Phallocysta eumekes, common with the Sortehat Formation assemblages. which is restricted to the latest Early Toarcian – Aalenian Davies (1983) stated that the macrofauna possibly indi- in Europe (Riding & Thomas 1992). Spores, pollen and cates a Toarcian – Early Bajocian age. dinoflagellate cysts from Unit C are similar to those In the Barents Sea area, seven dinoflagellate cyst from the Botryococcus and Nannoceratopsis gracilis – zones have been recognised in the Toarcian – Lower Nannoceratopsis senex Assemblage Zones of the Sortehat Oxfordian (Smelror & Below 1992). One of them, the Formation. Two thin levels with Botryococcus have Dodekovia bulla – Nannoceratopsis senex Concurrent been observed in the Stø Formation although not as rich Range-Zone, has a number of species (Nannoceratopsis in abundance as in the Sortehat Formation (Smelror et gracilis, Nannoceratopsis senex, Nannoceratopsis triceras, al. 1994). Moreover, the abrupt shift seen in the Sortehat Scriniocassis weberi, Susadinium scrofoides, Pareodinia Formation from an assemblage dominated by halosa and Phallocysta eumekes) in common with the Botryococcus to one dominated by the dinoflagellate assemblages from the Sortehat Formation. The pres- cysts Nannoceratopsis gracilis, Nannoceratopsis senex ence of the earliest Aalenian Opalinum Zone on Svalbard and Pareodinia halosa is not recognised in the Møre has been confirmed on the basis of ammonites. Basin. It should be noted, however that the Botryococcus In the United Kingdom, the Aalenian palynomorph Assemblage Zone of the Sortehat Formation is not strati- assemblages are divided into Sub-biozone c (Opalinum graphically significant but reflects only the palaeo- Zone) and Sub-biozone d (Murchisonae and Concavum environmental conditions within the Jameson Land Zones) of the Nannoceratopsis gracilis Zone (Riding & Basin at the time of deposition (see discussion below). Thomas 1992; the DSJ10 and DSJ11 Zones of Poulsen Aalenian palynomorph assemblages have also been & Riding 2003, this volume). These zones are based on identified from Arctic Canada and south-east Canada the FAD (first appearance datum) and the LAD (last (Johnson & Hills 1973; Davies 1983; Bujak & Williams appearance datum) of specific dinoflagellate cyst species. 1977), Barents Sea (Hammerfest Basin, Nordkap Basin, Of the stratigraphically significant species appearing and Franz Josef Land; Smelror & Below 1992; Smelror in these zones, Susadinium scrofoides, Wallodinium 1994), north-west Scotland, north Yorkshire and laganum and other species of the genus Parvocysta have Gloucestershire in England (Riding 1983, 1984a, b, 1987; also been found in the material from the Sortehat Riding et al. 1991; Riding & Thomas 1992), Sweden Formation. Other common but stratigraphically less (Guy-Ohlson 1994; Guy-Ohlson & Norling 1994), the restricted species are Nannoceratopsis gracilis, Nanno- Danish Subbasin (Dybkjær 1991; Seidenkrantz et al. ceratopsis senex, Mancodinium semitabulatum and 1993; Poulsen 1994), Øresund and the Baltic Sea Scriniocassis weberi. The palynological zonation in the (Koppelhus & Nielsen 1994; Koppelhus & Batten 1996), UK area has been related to the standard ammonite- and north-west and south-west Germany (Prauss 1989). based zones (Riding & Thomas 1992).

790 In Skåne, southern Sweden, the upper part of the Rya Nannoceratopsis plegas Zone of Aalenian age, has a Formation and the lower part of the Vilhelmsfält number of species in common with the three zones Formation contain palynomorph assemblages domi- from the Sortehat Formation, such as Nannoceratopsis nated by terrestrial material deposited in a freshwater gracilis, Mancodinium semitabulatum, Pareodinia environment, although thin marine and brackish levels halosa, Dodekovia tegillata, Scriniocassis weberi, occur. Palynological zone III is suggested to be of Phallocysta eumekes and Kallospharidium sp. Unfor- Aalenian age (Guy-Ohlson & Norling 1994), and shows tunately the suggested Aalenian age is not confirmed similarities to the palynomorph assemblages from the by ammonites. three zones of the Sortehat Formation. No ammonites To conclude this review, the Sortehat Formation paly- have been found in the Middle Jurassic of southern nomorph assemblages clearly show close similarities to Sweden but a foraminifera zonation has been established. assemblages of inferred Aalenian age in the North In the Danish Basin, rocks of Toarcian and early Atlantic region. Few of these studies, however, include Aalenian age have been identified based on the occur- independent ammonite data with which to accurately rence of species of the genus Parvocysta (Poulsen 1994). constrain the age of the strata. A notable exception, A miospore and dinoflagellate cyst zonation was erected upon which the age assignment of the Sortehat for the Lower and Middle Jurassic of the Danish Basin Formation depends, is the work of Riding (1982), (Dybkjær 1991). The Perinopollenites elatoides miospore Woollam & Riding (1983) and Riding & Thomas (1992) zone of Dybkjær (1991) covers sediments of Aalenian from the Jurassic of the UK. According to the zonation – Early Bajocian age, whereas the equivalent dinofla- presented by these workers, the stratigraphically impor- gellate cyst zone includes Nannoceratopsis gracilis and tant palynomorphs recorded from the Sortehat Formation comprises sediments of Late Pliensbachian – Bajocian are Mancodinium semitabulatum, Nannoceratopsis age. Similar palynomorph assemblages have been ambonis, Phallocysta eumekes and Nannoceratopsis ple- reported from the Øresund area (Koppelhus & Batten gas. M. semitabulatum, which was recorded through- 1996). A miospore zonation was recommended for the out the Sortehat Formation, has a range of Late Lower–Middle Jurassic of Bornholm in the Baltic area, Pliensbachian – Early Bajocian (Woollam & Riding 1983; where the Callialasporites–Perinopollenites Zone from Riding & Thomas 1992). N. ambonis, which occurs in the Bagå Formation has many species in common with the lower half of the formation, was initially thought to the Sortehat Formation (Batten et al. 1994; Koppelhus range from the Aalenian to the Early Bajocian (Riding & Nielsen 1994). Dinoflagellate cysts (Nannoceratopsis 1982; Woollam & Riding 1983) but its range has sub- gracilis) have only been found in the lowermost part sequently been extended to Late Pliensbachian – Early of this zone. The Callialasporites–Perinopollenites Zone Bajocian (Riding & Thomas 1992). P. eumekes, which is defined by the first appearance of Callialasporites and occurs throughout the Sortehat Formation, has a the dominance of Perinopollenites elatoides. Unfor- restricted range of Late Toarcian – Aalenian in Europe tunately, confirmatory ammonite or microfossil data is (Riding & Thomas 1992). However, the species was lacking in the Danish area to support the proposed first described from Bathonian strata in Arctic Canada Aalenian–Bathonian age, which is based entirely on where it has a range of Toarcian–Bathonian (Dörhöfer comparison to other palynological studies in Europe. & Davies 1980) and is generally considered a more In the Sentusidinium pelionense Assemblage Zone of long-ranging species in Boreal regions (Riding 1984c). the Sortehat Formation, spores and pollen are abundant, N. plegas is rare in the Sortehat Formation although its and dinoflagellate cysts are locally present. The dinofla- presence in the lower levels of the formation indicates gellate cyst Sentusidinium pelionense has also been an early Aalenian age for this part of the succession found in the Jydegård Formation on Bornholm in the (Riding & Thomas 1992). Baltic Sea (Piasecki 1984; Noe-Nygaard et al. 1987). In association, therefore, these palynomorphs indi- However, with the exception of the presence of S. pelion- cate an Aalenian – ?Early Bajocian age for the Sortehat ense, the Sentusidinium pelionense Assemblage Zone Formation. This study thus confirms the age determi- of the Sortehat Formation is more closely comparable nation of Underhill & Partington (1994), although it is to assemblages from the Norwegian and UK areas than clear from the above review that more detailed studies to those of the Danish Basin and Bornholm. of both the miospore and the dinoflagellate cyst strati- Palynomorph assemblages from the Pliensbachian– graphies of the early Middle Jurassic are urgently needed Callovian of north-west Germany were described by to further resolve the stratigraphy of the Sortehat For- Prauss (1989). The palynomorph assemblage from the mation.

791 A notable conclusion of this study is that there is no of dinocysts throughout the Sortehat Formation, albeit evidence, on palynological grounds, for a significant hia- in reduced numbers in the Botryococcus Assemblage tus at, or near, the lower boundary of the Sortehat For- Zone, suggests a persistent marine influence. Indeed, mation (see also Koppelhus & Dam 2003, this volume), the reduced number of marine dinoflagellate cysts in nor within the Sortehat Formation itself. These obser- this zone may be in part an artefact related to the count- vations are particularly pertinent to regional studies of ing procedure such that the abundance of Botryococcus Middle Jurassic uplift in the North Atlantic region (see and terrestrial sporomorphs tends to dilute the marine Underhill & Partington 1994). The stratigraphic signif- dinoflagellate cysts. Given the co-occurrence of the icance of the upper boundary of the Sortehat Formation freshwater alga Botryococcus and the marine dinofla- is less well-constrained palynologically, although there gellate cysts, the abundance of Botryococcus is here is no direct evidence from this study of a major strati- interpreted as an allochthonous accumulation. Seawards graphic break at this surface. transport of material from inland areas suggests either a major freshwater input from rivers or inundation and erosion of coastal areas with seawards transport of land- derived material during transgression. During marine Environmental implications of the flooding, the initial rise of base-level causes expansion palynological data of lakes on low-lying areas within the coastal plain The three palynomorph assemblage zones of the Sortehat (Wells & Coleman 1987; Dominguez & Wanless 1991; Formation can be recognised and correlated through- Surlyk et al. 1995). As transgression proceeds, physical out the Jameson Land Basin (Fig. 4). Regionally, the communication with the open sea is established, result- Botryococcus Assemblage Zone ranges in thickness from ing in the possibility for a large influx of freshwater algae 9–17 m and spans the uppermost part of the Ostreaelv into the marine system. Formation and the lower part of the Sortehat Formation. Such an allochthonous origin related to transgression The zone starts several metres below the boundary is consistent with the fact that the top of the Botryococcus between the Ostreaelv and Sortehat Formations in the Assemblage Zone coincides with the maximum flood- southernmost locality at Albuen, whereas the appear- ing surface and the end of the transgressive period ance of the zone coincides with the boundary at the (Hansen 1999). A similar influx of Botryococcus, related Sortehat type locality. Farther north, at Enhjørningen to a flooding event, has been described from the Middle Dal, this change in the palynomorph assemblage does Jurassic Brent Group of the North Sea by Williams not occur until 7 m above the formation boundary (Figs (1992), who interpreted this as an allochthonous accu- 4, 7). At the northernmost locality of Pelion, the change mulation resulting from the flushing of a freshwater takes place 5 m above the boundary. environment into a marine environment. The green alga Botryococcus is a freshwater form The diversification of the dinoflagellate cysts in the (Guy-Ohlson 1992) with no stratigraphic value as it basal part of the Nannoceratopsis gracilis – Nanno- ranges from the to the present (Tyson ceratopsis senex Assemblage Zone is suggestive of a 1995; Batten & Grenfell 1996). However, the abundance general increase in marine influence. This may be related of Botryococcus in the organic material from the low- to an increase in salinity during sea-level highstand fol- ermost Sortehat Formation in Jameson Land is notable. lowing the transgression, as it is generally suggested that Sedimentologically, the depositional shift from the an increased diversity of dinoflagellate cysts points to Ostreaelv Formation to the Sortehat Formation marks more open marine conditions (Gorin & Steffen 1991; a basinwide flooding event within an overall trans- Leckie et al. 1992). gressive period. As the abundance of Botryococcus is Upwards within the Nannoceratopsis gracilis – Nan- restricted to the section around the flooding surface noceratopsis senex Assemblage Zone and through the (base of Sortehat Formation) and extends up to the Sentusidinium pelionense Assemblage Zone, especially maximum flooding surface, it seems to be linked to the at Albuen and partly at Enhjørningen Dal, there is a flooding event (Hansen 1999). The question is whether marked decrease in the diversity of dinoflagellate cyst the abundance of Botryococcus reflects in situ deposi- species, although the assemblage still shows marine tion under freshwater conditions that prevailed through- influence. A decrease in diversity indicates more stressed out the basin or if it represents an allochthonous and unfavourable environmental conditions, often with accumulation of algae transported out to sea from inland unstable salinities (Gorin & Steffen 1991; Leckie et al. freshwater environments. The continuous occurrence 1992; Tyson 1995).

792 Although the strength of the marine signal partly blages become poor in diversity and richness, suggest- increases up-section, the dinoflagellate cysts are char- ing more stressed conditions, but dinoflagellate cysts acterised by forms tolerant of reduced salinities (e.g. continue into the sandy Pelion Formation. Nannoceratopsis gracilis, Nannoceratopsis senex and Sentusidinium pelionense; Piasecki 1986; Prauss & Riegel 1989; Krabbe et al. 1994). The palynological data there- fore indicate a marine environment, yet restricted in Acknowledgements terms of salinity such that brackish conditions prevailed This study was supported by a grant from the Danish during deposition of the Sortehat Formation. Energy Research Programme (EFP-93, projects 1313/93- 0010, -0017) to the Geological Survey of Greenland. We are grateful to David J. Batten, Karen Dybkjær, Jon R. Ineson, Henrik Nøhr-Hansen, Stefan Piasecki and Conclusions Finn Surlyk for discussion and critical reading of the man- The age of the Sortehat Formation is Aalenian to ?Early uscript and to Henrik Nøhr-Hansen in particular for Bajocian. This age assignment is based on comparison patient assistance with the distribution charts. of the three palynomorph assemblage zones (the Botryo- coccus Assemblage Zone, the Nannoceratopsis gracilis – Nannoceratopsis senex Assemblage Zone and the Sentusidinium pelionense Assemblage Zone) with paly- References nomorph assemblages in the North Atlantic region that Batten, D.J. & Grenfell, H.R. 1996: Botryococcus. In: Jansonius, have been recorded either from sections dated directly J. & McGregor, D.C. (eds): Palynology: principles and appli- by ammonites or from sections that can be reliably cor- cations. American Association of Stratigraphic Palynologists Foundation 1, 205–214. related to successions with good ammonite control. 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Manuscript received 16 April 1997; revision accepted 18 January 1999.

795 West Enhjørningen Dal Appendix 1: Locality information and Geological Location. 71°26′53′′N, 23°25′41′′W; the northernmost of two small Survey of Greenland (GGU) numbers ravines. of samples used in this study Sample numbers. GGU 398203–398299, 398308–398372. Albuen

Location. 70°34′10′′N, 22°38′54′′W. East Enhjørningen Dal

Section datum. The boundary between the Ostreaelv and the Location. 71°26′14′′N, 23°21′52′′W; the northernmost of two small Sortehat Formations is the datum for the measured section and ravines. is located 440 m above sea level. Sample numbers. GGU 395616–395668, 398417–398448. Sample numbers. GGU 397452–397498, 398501, 398507, 398509, 398513. Pelion

Sortehat borehole Location. 71°28′N, 23°19′W.

Location. 70°53′50′′N, 22°49′56′′W. Sample numbers. GGU 339702–339711. These samples were col- lected by Lars Stemmerik in 1990. Section datum. For the drill-core, an arbitrary datum was defined 80 m below the top of the borehole. The boundary between the Ostreaelv and the Sortehat Formations occurs 27.6 m above this datum.

Sample numbers. GGU 303143-11–303143-80.

Lepidopteriselv

Location. 71°15′12′′N, 22°37′04′′W.

Section datum. The boundary between the Ostreaelv and the Sortehat Formations, which forms the datum for the section, lies at c. 810 m above sea level.

Sample numbers. GGU 398151–398199.

Enhjørningen Dal

Two sections were measured in Enhjørningen Dal; Figure 7 is a composite of these sections. The boundary between the Ostreaelv and the Sortehat Formations, at 418 m above sea level, is the datum for the composite section.

796 Kraeuselisporites reissingeri (Harris) Morbey 1975 Appendix 2: Leptolepidites bossus (Couper) Schulz 1967 List of all recorded palynomorph taxa L. major Couper 1958 L. spp. Miospores: Limbosporites lundbladii Nilsson 1958 Lycopodiacidites rugulatus (Couper) Schulz 1967 Alisporites robustus Nilsson 1958 Manumia delcourtii (Pocock) Dybkjær 1991 (Plate 1, fig. 8) Apiculatisporites spp. Megaspore fragments Araucariacites australis Cookson 1947 Monosaccate pollen Baculatisporites comaumensis (Cookson) Potonié 1956 Monosulcites spp. (Plate 1, fig. 3) Murospora spp. B. spp. Neoraistrickia gristhorpensis (Couper) Tralau 1967 Bisaccates indeterminate N. taylorii Playford & Dettmann 1965 Callialasporites dampieri (Balme) Dev 1961 (Plate 2, fig. 1) N. spp. C. microvelatus Schulz 1966 (Plate 2, fig. 3) Perinopollenites elatoides Couper 1958 (Plate 2, fig. 5) C. minus (Tralau) Guy 1971 Pinuspollenites minimus (Couper) Kemp 1970 (Plate 3, fig. 6) C. segmentatus (Balme) Dev 1961 Polycingulatisporites circulus Simoncsics & Kedves 1961 C. trilobatus (Balme) Dev 1961 Polycingulatisporites triangularis (Bolkhovitina) Playford & C. turbatus (Balme) Schulz 1967 (Plate 2, fig. 2) Dettmann 1965 C. spp. Quadraeculina anellaeformis Malyavkina 1949 (Plate 3, fig. 5) Calamospora tener (Leschik) Mädler 1964 Retitriletes austroclavatidites (Cookson) Döring et al. 1963 Camarozonasporites spp. (Plate 1, fig. 1) Cerebropollenites macroverrucosus (Thiergart) Schulz 1967 R. clavatoides (Couper) Döring et al. 1963 (Plate 1, fig. 2) (Plate 3, fig. 3) R. semimuris (Danzé-Corsin & Laveine) McKellar 1974 C. thiergartii Schulz 1967 R. spp. Chasmatosporites apertus Nilsson 1958 Ricciisporites tuberculatus Lundblad 1954 C. hians Nilsson 1958 (Plate 2, fig. 4) Rogalskaisporites cicatricosus (Rogalska) Danzé-Corsin & Laveine C. major Nilsson 1958 (Plate 2, fig. 6) 1963 Chomotriletes minor (Kedves) Pocock 1970 (Plate 4, fig. 1) Sestrosporites pseudoalveolatus (Couper) Dettmann 1963 Cibotiumspora jurienensis (Balme) Filatoff 1975 Spheripollenites psilatus Couper 1958 Conbaculatisporites mesozoicus Klaus 1960 S. spp. (Plate 3, fig. 4) C. spp. S. subgranulatus Couper 1958 Concavissimisporites spp Staplinisporites caminus (Balme) Pocock 1970 (Plate 1, fig. 5) Corollina torosus (Reissinger) Cornet & Traverse 1975 S. spp. (Plate 3, fig. 2) Stereisporites stereoides (Potonié & Venitz) H.D. Pflug in: Thomson C. spp. & Pflug 1953 Deltoidospora minor (Couper) Pocock 1970 Striatella jurassica Mädler 1964b (Plate 1, fig. 4) D. spp. S. parva (Li & Shang) Filatoff & Price 1988 D. toralis (Leschik) Lund 1977 S. seebergensis Mädler 1964b Densoisporites scanicus Tralau 1968 S. spp. D. velatus Weyland & Krieger 1953 Striate spp. Densosporites spp. Taeniasporites rhaeticus Schulz 1967 Eucommiidites troedsonii Erdtman 1948 T. spp. Exesipollenites tumulus Balme 1957 (Plate 3, fig. 1) Taurocusporites verrucatus Schulz 1967 Foraminisporis jurassicus Schulz 1967 T. spp. Fungal spores Tigrisporites spp. Iraqispora labrata Singh 1964 Todisporites major Couper 1958 Ischyosporites crateris Balme 1957 T. minor Couper 1958 I. spp. (Plate 1, fig. 6) Triancoraesporites spp. I. variegatus (Couper) Schulz 1967 Trilete spp. Kekryphalospora distincta Fenton & Riding 1987 Tripartina variabilis Malyavkina 1949 (Plate 1, fig. 7)

797 Uvaesporites puzzlei Guy 1971 Miscellaneous U. spp. Nannoceratopsis ambonis Drugg 1978 (Plate 5, fig. 3) Vesicaspora fuscus (Pautsch) Morbey 1975 N. gracilis Alberti emend. van Helden 1977 (Plate 5, fig. 1) Vitreisporites pallidus (Reissinger) Nilsson 1958 N. plegas Drugg 1978 (Plate 5, fig. 4) Zebrasporites laevigatus (Schulz) Schulz 1967 N. ridingii Poulsen 1992 N. senex van Helden 1977 (Plate 5, fig. 2) N. spp. Phytoplankton: N. triangulata Prauss 1987 N. triceras Drugg 1978 Acritarch spp. Pareodinia ceratophora Deflandre 1947 Andreedinium arcticum Below 1987 P. halosa (Filatoff) Prauss 1989 (Plate 4, fig. 6) Beaumontella caminuspina (Wall) Below 1987 P. spp. Botryococcus spp. (Plate 4, fig. 4) Parvocysta barbata Bjærke 1980 Chytroeisphaeridia chytroeoides (Sarjeant) Downie & Sarjeant P. spp. 1965 Phallocysta eumekes Dörhöfer & Davies 1980 Cymatiosphaera spp. P. thomasi Smelror 1991 Dinocyst spp. P. spp. Dissilodinium spp. (Plate 6, figs 2, 4) Pterospermella spp Dodekovia tegillata Prauss 1989 Scrinocassis weberi Gocht 1964 Foraminiferal linings (Plate 4, fig. 3) S. spp. Fungal spp. Sentusidinium pelionense Fensome 1979 (Plate 6, figs 5, 6) Kallosphaeridium spp. S. spp. Lecaniella foveata Singh 1971 Susadinium scrofoides (Dörhöfer & Davies) Below 1987 Leiosphaeridia spp. Tasmanites spp. Limbicysta bjaerkei (Smelror) MacRae et al. 1996 Valensiella ovulum (Deflandre) Eisenack 1963 Mancodinium semitabulatum Morgenroth 1970 (Plate 6, fig. 1) Veryhachium collectum Wall 1965 M. spp. V. formosum Stockmans & Williere 1960 Mendicodinium groenlandicum (Pocock & Sarjeant) Davey 1979 V. sortehatense Fensome 1979 (Plate 4, fig. 2) (Plate 6, fig. 3) V. spp. M. reticulatum Morgenroth 1970 Wallodinium laganum Feist-Burkhardt & Monteil 1994 M. spp. (Plate 4, fig. 5)

798 Plates 1–6

799 Plate 1

Figs 1–4 and 6–8 are from the Lepidopteriselv section, fig. 5 is from the borehole at Sortehat, the type locality of the Sortehat Formation. Scale bar is 10 microns. For each of the illustrated specimens, the EFR (England Finder Reference) is given.

Fig. 1. Retitriletes austroclavatidites. Sample 398181, slide 3, EFR E36.

Fig. 2. Retitriletes clavatoides. Sample 398181, slide 3, EFR F50.

Fig. 3. Baculatisporites comaumensis. Sample 398158, slide 4, EFR S26.

Fig. 4. Striatella jurassica. Sample 398158, slide 5, EFR K271.

Fig. 5. Staplinisporites caminus. Sample 303143-26, slide 3, EFR J51.

Fig. 6. Ischyosporites sp. Sample 398189, slide 4, EFR M40.

Fig. 7. Tripartina variabilis. Sample 398189, slide 4, EFR R43.

Fig. 8. Manumia delcourtii. Sample 398192, slide 4, EFR W383.

800 2

1

4 3

6 5

7 8

801 Plate 2

Figs 1 and 4–6 are from the borehole at Sortehat, figs 2 and 3 are from the Lepidopteriselv section. Scale bar is 10 microns.

Fig. 1. Callialasporites dampieri. Sample 303143-46, slide 3, EFR F222.

Fig. 2. Callialasporites turbatus. Sample 398158, slide 4, EFR P25.

Fig. 3. Callialasporites microvelatus. Sample 398158, slide 4, EFR H382.

Fig. 4. Chasmatosporites hians. Sample 303143-46, slide 3, EFR L42.

Fig. 5. Perinopollenites elatoides. Sample 303143-13, slide 3, EFR N25.

Fig. 6. Chasmatosporites major. Sample 303143-46, slide 3, EFR K43.

802 1

2

4

3

5 6

803 Plate 3

Figs 1, 2, 5 and 6 are from the Lepidopteriselv section, figs 3 and 4 are from the borehole at Sortehat. Scale bar is 10 microns.

Fig. 1. Exesipollenites tumulus. Sample 398181, slide 3, EFR J362.

Fig. 2. Corollina torosus. Sample 398158, slide 5, EFR M33.

Fig. 3. Cerebropollenites macroverrucosus. Sample 303143-40, slide 3, EFR F24.

Fig. 4. Spheripollenites sp. Sample 303143-46, slide 3, EFR W20.

Fig. 5. Quadraeculina anellaeformis. Sample 398158, slide 4, EFR S384.

Fig. 6. Pinuspollenites minimus. Sample 398158, slide 5, EFR D522.

804 1 2

4

3

56

805 Plate 4

Figs 1, 2 and 6 are from the borehole at Sortehat, figs 3–5 are from the Lepidopteriselv section. Scale bar is 10 microns.

Fig. 1. Chomotriletes minor. Sample 303143-46, slide 3, EFR K46.

Fig. 2. Veryhachium sortehatense. Sample 303143-46, slide 3, EFR H22.

Fig. 3. Foraminiferal inner lining. Sample 398158, slide 4, EFR H31.

Fig. 4. Botryococcus sp. Sample 398189, slide 4, EFR E492.

Fig. 5. Wallodinium laganum. Sample 398194, slide 4, EFR Z39.

Fig. 6. Pareodinia halosa. Sample 303143-26, slide 3, EFR M51.

806 1 2

3

4

56

807 Plate 5

Fig. 1 is from the Lepidopteriselv section, figs 2–4 are from the borehole at Sortehat. Scale bar is 10 microns.

Fig. 1. Nannoceratopsis gracilis. Sample 398158, slide 4, EFR H54.

Fig. 2. Nannoceratopsis senex. Sample 303143-40, slide 3, EFR S363.

Fig. 3. Nannoceratopsis ambonis. Sample 303143-46, slide 3, EFR J40.

Fig. 4. Nannoceratopsis plegas. Sample 303143-40, slide 3, EFR M482.

808 1 2

3 4

809 Plate 6

Figs 1, 2 and 4–6 are from the borehole at Sortehat, fig. 3 is from the Lepidopteriselv section. Scale bar is 10 microns.

Fig. 1. Mancodinium semitabulatum. Sample 303143-46, slide 3, EFR S264.

Fig. 2. Dissilodinium sp. Sample 303143-30, slide 3, EFR K324.

Fig. 3. Mendicodinium groenlandicum. Sample 398158, slide 5, EFR R39.

Fig. 4. Dissilodinium sp. Sample 303143-26, slide 3, EFR C54.

Fig. 5. Sentusidinium pelionense. Sample 303143-20, slide 3, EFR M51.

Fig. 6. Sentusidinium pelionense. Sample 303143-26, slide 3, EFR E53.

810 1 2

3 4

5 6

811

M Albuen (A) Alphabetical species list 31 Anapiculatisporites sp.

sp. 32 Anapiculatisporites telephorus sp.

sp. 68 Araucariacites australis sp.

sp. 8 Baculatisporites sp. sp. sp. spp. sp. sp. 56 Bisaccate spp. sp.

sp. 66 Callialasporites dampieri sp. sp. sp. 73 Callialasporites microvelatus spp. 74 Callialasporites minus spp. sp. 67 Callialasporites sp. 76 Callialasporites trilobatus inopollenites elatoides aminisporis jurassicus 77 Callialasporites turbatus r r ipartina variabilis iletes ekryphalospora distincta esicaspora fuscus esicaspora olycingulatisporites triangularis olycingulatisporites circulus odisporites minor odisporites major odisporites aurocusporites verrucatus aeniasporites 72 Campenia sp. Rogalskaisporites cicatricosus Conbaculatisporites mesozoicus Lycopodiacidites rugulatus Deltoidospora Baculatisporites Tigrisporites microrugulatus Fo Kraeuselisporites reissingeri Tr Retitriletes clavatoides Retitriletes semimuris Retitriletes Stereisporites stereoides Densoisporites scanicus Retitriletes austroclavatoides T K Cibotiumsporites jurienensis Striatella parva T Striatella jurassica Leptolepidites Zebrasporites interscriptus Stereisporites antiquasporites Uvaesporites argenteaeformis Tr Striatella Striatella seebergensis Chomotriletes Anapiculatisporites Anapiculatisporites telephorus Neoraistrickia T Densoisporites velatus Sculptisporites aulosenensis Uvaesporites T Manumia delcourtii Ischyosporites variegatus Ischyosporites crateris Staplinisporites caminus P Sestrosporites pseudoalveolatus P Neoraistrickia taylorii Leptolepidites major Limbosporites lundbladii Murospora Chasmatosporites hians thiergartiiCerebropollenites V Quadraeculina anellaeformis Quadraeculina Pe Pinuspollenites minimus macroverrucosus Cerebropollenites Chasmatosporites major Chasmatosporites apertus torosus Corollina Monosulcites punctatus Chasmatosporites Chasmatosporites elegans Vitreisporites pallidus Callialasporites dampieri Callialasporites Araucariacites australis Eucommiidites troedsonii Cerebropollenites Spheripollenites subgranulatus Campenia Callialasporites microvelatus Callialasporites minus tumulus Exesipollenites Callialasporites trilobatus Callialasporites turbatus Eucommiidites major Ricciisporites tuberculatus T Striate 57 Cerebropollenites macroverrucosus 70 Cerebropollenites sp. System Stage Lithostratigraphy Assembl. Zones Palynological (m) Sample height Sample number 1 2 3 4 8 9 10 11 12 5 6 7 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 fragments Megaspore 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 Bisaccate spp. 57 58 59 60 61 62 63 64 Monosaccate spp. 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 51 Cerebropollenites thiergartii 59 Chasmatosporites apertus 439.00 341248 63 Chasmatosporites elegans A 7 438.50 397452 50 Chasmatosporites hians 58 Chasmatosporites major 434.00 405449 ?

432.00 341247 ? 62 Chasmatosporites sp. 431.50 405450 29 Chomotriletes sp. 428.00 405451 ? 18 Cibotiumsporites jurienensis 425.90 405452 R 2 Conbaculatisporites mesozoicus 60 Corollina torosus 4 Deltoidospora spp. 420 14 Densoisporites scanicus 416.50 405453 35 Densoisporites velatus

efjord Bjerg Mb efjord 78 Eucommiidites major 69 Eucommiidites troedsonii Tr 411.50 405454 75 Exesipollenites tumulus 10 Foraminisporis jurassicus 6 41 Ischyosporites crateris 40 Ischyosporites variegatus 402.00 405455 R 17 Kekryphalospora distincta 400 11 Kraeuselisporites reissingeri 47 Leptolepidites major 22 Leptolepidites sp. 392.00 405456 48 Limbosporites lundbladii 3 Lycopodiacidites rugulatus 39 Manumiadel courtii 30 Megaspore fragments 64 Monosaccate spp. 380 61 Monosulcites punctatus 379.40 405457 49 Murospora sp. 377.50 341243 oarcian 33 Neoraistrickia sp.

T 375.50 405458 46 Neoraistrickia taylorii Skævdal Member 54 Perinopollenites elatoides 369.50 405459 55 Pinuspollenites minimus

5 367.50 341245 ? 45 Polycingulatisporites circulus 366.20 405460 43 Polycingulatisporites triangularis Ostreaelv Formation Ostreaelv 365.00 341241 53 Quadraeculinaanellae formis 360 363.00 405462 15 Retitriletes austroclavatoides 361.30 405464 5 Retitriletes clavatoides 359.00 405466 6 Retitriletes semimuris R 7 Retitriletes sp. 79 Ricciisporites tuberculatus Interval not sampled 1 Rogalskaisporites cicatricosus 36 Sculptisporites aulosenensis 300 44 Sestrosporites pseudoalveolatus 297.80 405434 71 Spheripollenites subgranulatus Lower Jurassic Lower 295.40 405433 42 Staplinisporites caminus 24 Stereisporites antiquasporites 4 294.20 405432 292.20 405431 13 Stereisporites stereoides 290.75 405430 81 Striate sp. 290.20 405429 21 Striatella jurassica 287.80 341236 19 Striatella parva 287.00 405428 28 Striatella seebergensis 27 Striatella sp. 280 284.70 405427 R

Nathorst Fjeld Member 283.40 405426 80 Taeniasporites sp. 34 Taurocusporites verrucatus 273.80 405425 9 Tigrisporites microrugulatus 271.00 341235 20 Todisporites major 16 Todisporites minor 267.40 405424 38 Todisporites sp. 3 26 Triletes sp. 264.27 405423 R 264.25 405422 12 Tripartina variabilis 260 25 Uvaesporites argenteaeformis 37 Uvaesporites sp. 256.50 341234 52 Vesicaspora fuscus 253.40 405421 65 Vitreisporites pallidus 23 Zebrasporites interscriptus 250.50 405420 249.00 341233 247.40 405419 246.00 405418 240 241.00 405417 240.00 405416 238.20 405414 2 238.00 405413 Uncertain

Albuen Mb As ? 236.80 405411 determination

231.70 405410 ? 229.80 405408 R Ver y rare 229.50 341232 229.40 405406 Rare 220 229.00 405405 Few 222.50 405404 1 215.50 405403 Common 213.00 405402 Abundant 211.00 405401 R Fig. 4A. Terrestrial palynomorph distribution chart for the Gule Horn and Ostreaelv Formations at Albuen (for location, see Fig. 1). M, Middle Jurassic; A, Aalenian; As, Astartekløft Member. R R Gule Horn Formation Elis Bjerg Member R R L. PliensbachianL. Upper Pliensbachian Albuen (B)

Alphabetical species list 27 Acritarch spp. 5 Beaumontella caminuspina spp.

sp. 11 Beaumontella delicata sp. sp. sp. spp. spp. 45 Beaumontella sp. spp. spp. sp. sp. spp. spp. sp. 47 Botryococcus spp. sp. sp.

spp. 51 Celyphus spp. spp. 44 Cymatiosphaera sp.

A 14 Dinocyst sp. M vocysta barbata vocysta yhachium formosum yhachium collectum yhachium reductum yhachium irregulare yhachium trispinosum eodinia halosa r r r r r r r r 18 Dissiliodinium sp. allodinium alvaeodinium armatum alvaeodinium etraporina compressa asmanites Nannoceratopsis senex Nannoceratopsis Nannoceratopsis gracilis Nannoceratopsis Mendicodinium reticulatum Beaumontella caminuspina triangulata Nannoceratopsis plegas Nannoceratopsis Mancodinium semitabulatum Pa Mendicodinium groenlandicum Beaumontella delicata Mendicodinium Luehndea spinosa V V Nannoceratopsis ambonis Nannoceratopsis Dissiliodinium Leiofusa jurassica Leiosphaeridia Pa Phallocysta eumekes triceras Nannoceratopsis Pa Kallosphaeridium Scriniocassis Phallocysta elongata W Micrhystridium lymensis Micrhystridium Ve Limbicysta bjaerkei Ve T Lecaniella foveata Ve Ve Micrhystridium intromittum Micrhystridium wattonense Micrhystridium fragile Ve Micrhystridium stellatum Cymatiosphaera Beaumontella Pterospermella Pterospermella Botryococcus T Lecaniella Celyphus Haplophragmoides Haplophragmoides 50 Fungal spores 52 Haplophragmoides spp. System Stage Lithostratigraphy Assembl. Zones Palynological (m) Sample height Sample number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Dinocyst sp. 15 16 17 18 27 spp. Acritarch 28 29 19 20 21 22 23 24 25 26 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 Fungal spores 51 52 53 Miscellaneous spp. 23 Kallosphaeridium sp. 36 Lecaniella foveata

439.00 341248 ? 7 49 Lecaniella spp. 438.50 397452 28 Leiofusa jurassica efjord Bjerg Mb efjord 434.00 405449 29 Leiosphaeridia spp. Tr 432.00 341247 ? 33 Limbicysta bjaerkei 431.50 405450 13 Luehndea spinosa 428.00 405451 ? 53 Miscellaneous spp. 425.90 405452 8 Mancodinium semitabulatum 10 Mendicodinium groenlandicum 420 4 Mendicodinium reticulatum 12 Mendicodinium sp. 416.50 405453 41 Micrhystridium fragile 39 Micrhystridium intromittum 411.50 405454 30 Micrhystridium lymensis 31 Micrhystridium spp.

oarcian 43 Micrhystridium stellatum T Skævdal Member 40 Micrhystridium wattonense 6 402.00 405455 17 Nannoceratopsis ambonis 400 3 Nannoceratopsis gracilis

Ostreaelv Formation Ostreaelv 7 Nannoceratopsis plegas 1 Nannoceratopsis senex 2 Nannoceratopsis sp. 392.00 405456 6 Nannoceratopsis triangulata 21 Nannoceratopsis triceras 22 Pareodinia halosa 9 Parvocysta barbata 19 Parvocysta sp. 380 379.40 405457 25 Phallocysta elongata

377.50 341243 ? ? 20 Phallocysta eumekes 375.50 405458 46 Pterospermella spp. 24 Scriniocassis sp. 48 Tasmanites sp. 369.50 405459 35 Tetraporina compressa 5 367.50 341245 ? 15 Valvaeodinium armatum ? Nathorst Fjeld Member 366.20 405460 16 Valvaeodinium spp. 365.00 341241 34 Veryhachium collectum 360 363.00 405462 32 Veryhachium formosum 361.30 405464 38 Veryhachium irregulare 359.00 405466 37 Veryhachium reductum 42 Veryhachium trispinosum Interval not sampled 26 Wallodinium spp.

300 Lower Jurassic Lower

4 292.20 405431

Albuen Mb As 287.80 341236 287.00 405428 284.70 405427 280 283.40 405426

273.80 405425 271.00 341235

267.40 405424 3 264.27 405423 264.25 405422 260

256.50 341234 253.40 405421 250.50 405420 249.00 341233 Upper Pliensbachian 247.40 405419 246.00 405418 240 241.00 405417 240.00 405416 Gule Horn Formation 238.20 405414 2 238.00 405413 Uncertain Elis Bjerg Member ? 236.80 405411 determination 231.70 405410 R Ver y rare 229.80 405408 229.50 341232 Rare 229.40 405406 220 229.00 405405 Few 222.50 405404 1 Common 215.50 405403 213.00 405402 Abundant 211.00 405401

Fig. 4B. Marine palynomorph distribution chart for the Gule Horn and Ostreaelv Formations at Albuen (for location, see Fig. 1). M, Middle Jurassic; A, Aalenian; As, Astartekløft Member. L. Pliensbachian L. Lepidopteriselv Alphabetical species list sp. 23 Anapiculatisporites sp. sp. sp. 26 Annulispora folliculosa sp.

sp. 15 Apiculatisporites parvispinosus sp. spp. spp. sp. 27 Apiculatisporites sp. sp. sp. sp. sp. 45 Araucariacites australis sp. spp. 1 Baculatisporites sp. spp. sp. 34 Bisaccate spp. 65 Botryococcus spp. 3 Calamospora tener yhachium trisulcum yhachium inopollenites elatoides 48 Callialasporites minus r r r ipartina variabilis

ekryphalospora distincta 47 Callialasporites turbatus esicaspora fuscus esicaspora odisporites major asmanites Baculatisporites Deltoidospora Calamospora tener Calamospora T Rogalskaisporites cicatricosus Striatella parva Retitriletes austroclavatoides Retitriletes clavatoides Neoraistrickia Stereisporites stereoides Retitriletes semimuris Kraeuselisporites reissingerii Retitriletes Tigrisporites microrugulatus Apiculatisporites parvispinosus Conbaculatisporites mesozoicus Densoisporites scanicus Lycopodiacidites rugulatus Iraquispora Tigrisporites Tr K Anapiculatisporites Marattiisporites scabratus minor Deltoidospora folliculosa Annulispora Apiculatisporites Striatella jurassica Densosporites Densosporites variabilis rudis Camarozonosporites Murospora anellaeformis Quadraeculina Pe Chasmatosporites hians Pinuspollenites minimus Chasmatosporites apertus thiergartiiCerebropollenites torosus Corollina Monosulcites punctatus macroverrucosus Cerebropollenites V Chasmatosporites major Araucariacites australis Spheripollenites psilatus Callialasporites turbatus Callialasporites minus Eucommiidites troedsonii Corollina Vittatina Mendicodinium reticulatum Mancodinium semitabulatum senex Nannoceratopsis triangulata Nannoceratopsis gracilis Nannoceratopsis Nannoceratopsis Micrhystridium intromittum Ve Micrhystridium fragile Lecaniella Micrhystridium lymensis Ve Botryococcus T 31 Camarozonosporites rudis 42 Cerebropollenites macroverrucosus 39 Cerebropollenites thiergartii Lithostratigraphy Assembl. Zones Palynological (m) Sample height Sample number System Stage 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Bisaccate spp. 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 Monosaccate spp. 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 38 Chasmatosporites apertus 36 Chasmatosporites hians 700 700.00 139146 R 44 Chasmatosporites major 16 Conbaculatisporites mesozoicus 51 Corollina sp. 40 Corollina torosus 695.00 139145 25 Deltoidospora minor 2 Deltoidospora spp. 17 Densoisporites scanicus 29 Densosporites sp. 30 Densosporites variabilis 49 Eucommiidites troedsonii 19 Iraquispora sp. 3 22 Kekryphalospora distincta 12 Kraeuselisporites reissingerii

685.00 139144 R ? 62 Lecaniella spp. 18 Lycopodiacidites rugulatus 54 Mancodinium semitabulatum 24 Marattiisporites scabratus 53 Mendicodinium reticulatum 61 Micrhystridium fragile 679.00 139143 59 Micrhystridium intromittum 63 Micrhystridium lymensis 50 Monosaccate spp. 675 R 41 Monosulcites punctatus 674.00 139142 32 Murospora sp. 57 Nannoceratopsis gracilis 55 Nannoceratopsis senex 58 Nannoceratopsis sp. 56 Nannoceratopsis triangulata 9 Neoraistrickia sp. 35 Perinopollenites elatoides 37 Pinuspollenites minimus 33 Quadraeculina anellaeformis 7 Retitriletes austroclavatoides

Lower Jurassic Lower 8 Retitriletes clavatoides 11 Retitriletes semimuris Elis Bjerg Member 13 Retitriletes sp. Upper Pliensbachian

Gule Horn Formation 5 Rogalskaisporites cicatricosus 46 Spheripollenites psilatus 10 Stereisporites stereoides 28 Striatella jurassica 6 Striatella parva 654.00 139141 66 Tasmanites sp. 653.00 139140 14 Tigrisporites microrugulatus 20 Tigrisporites sp. 650 4 Todisporites major 21 Tripartina variabilis

648.00 139139 ? 64 Veryhachium sp. 60 Veryhachium trisulcum 2 43 Vesicaspora fuscus 52 Vittatina sp.

642.00 139138 ? Uncertain ? determination R Ver y rare 638.00 139137 ? Rare R Few Common Abundant

Fig. 13. Palynomorph distribution chart for the Gule Horn Formation (Elis Bjerg Member) at Lepidopteriselv (for location, see Fig. 1).