Sorthat Formation Explained

Sorthat Formation
Type:Geological formation
Age:Latest Pliensbachian to Latest Toarcian
~Possible Lower Aalenian layers
Period:Toarcian
Prilithology:Claystone, sandstone[1]
Unitof:Bornholm Group
Subunits:Sorthat & Levka beds
Overlies:Rønne & Hasle Formations
Underlies:Bagå Formation
Thickness:240m (790feet)
Namedfor:Sorthat-Muleby, Bornholm
Namedby:Gry (as part of the Bagå Formation) [2]
Year Ts:1969
Region:
    • Rønne Graben
Country:
    • (ex situ sandstones)
Coordinates:55.09°N 14.42°W
Paleocoordinates:Approx. 35°N

The Sorthat Formation is a geologic formation on the island of Bornholm, Denmark and in the Rønne Graben in the Baltic Sea. It is of Latest Pliensbachian to Late Toarcian age. Plant fossils have been recovered from the formation, along with several traces of invertebrate animals. The Sorthat Formation is overlain by fluvial to lacustrine gravels, along with sands, clay and in some places coal beds that are part of the Aalenian-Bathonian Bagå Formation.[2] Until 2003, the Sorthat Formation was included as the lowermost part of the Bagå Formation, recovering the latest Pliensbachian to lower Aalenian boundary.[3] [4] The Sorthat strata reflect a mostly marginally deltaic to marine unit. Large streams fluctuated to the east, where a large river system was established at the start of the Toarcian.[2] In the northwest, local volcanism that started in the lower Pliensbachian extended along the North Sea, mostly from southern Sweden.[5] At this time, the Central Skåne Volcanic Province and the Egersund Basin expelled most of their material, with influences on the local tectonics.[5] The Egersund Basin has abundant fresh porphyritic nephelinite lavas and dykes of lower Jurassic age, with a composition nearly identical to those found in the clay pits. That indicates the transport of strata from the continental margin by large fluvial channels of the Sorthat and the connected Röddinge Formation that ended in the sea deposits of the Ciechocinek Formation green series.[5]

Stratigraphy

On Bornholm, the lower-middle Jurassic succession is composed of the Rønne (HettangianSinemurian), Hasle (lower–upper Pliensbachian), Sorthat and Bagå Formations. The major Pliensbachian–Bathonian coal-bearing clays and sands that overlie the Lower Pliensbachian Hasle Formation are distributed between both the Sorthat Formation and the overlaying Bagå Formation.[1] The Sorthat Formation is the sister unit of the Röddinge Formation, with both being part of the same fluvial system, as well the regional equivalent of the Ciechocinek Formation of Baltic Germany and Poland, the Fjerritslev Formation of the Danish Basin and the Rya Formation on Scania.[1] The Sorthat Formation beds were referred originally to the Levka, Sorthat and Bagå beds.[2] A major section of the formation is the Korsodde coastal section, located on the southwest part of the island.[2] A detailed stratigraphic interpretation of the beds has been difficult to achieve, in part due to the complicated block faulting, but especially due to the absence of marine fossils and distinct marker beds.[2] The rocks were originally dated as Middle Jurassic using megaspore contents, with the Levka and Sorthat beds being roughly contemporaneous and the Bagå beds possibly slightly younger. Later, when more advanced palynological studies from locations such as the Levka-1 core-well and the Korsodde section Upper Pliensbachian stratum became available,[6] [7] the coals and clays of the Levka beds were removed from the Bagå Formation, as were the coal-dominated beds of the Korsodde and Onsbæk sections.[3] At the time, several megaspores were found to be common in both the Bagå Formation and Sorthat beds, implying the presence of Toarcian–Aalenian strata,[3] although the dating of the megaspore-bearing strata is tentative.[8] With both, the palynological and sedimentological study of all available exposures and cores from the Lower–Middle Jurassic shows that the Hasle Formation (Lower–Middle Pliensbachian) is covered by a succession referable to both the Levka and Sorthat beds, which are composed mostly by bioturbated sands, heteroliths and clays along with abundant coal veins, and contain relatively diverse brackish to marine dinoflagellate assemblages that are indicative of upper Pliensbachian, Toarcian and possibly lower Aalenian strata.[6] The upper stratum is covered by the fluvial gravels and sands, along with lacustrine clays, carbonaceous clays and coals belonging to the Bagå Formation.[1]

Lithology

The Sorthat Formation has a highly variable lithology.[1] The main core studied from the rocks, the Levka-1 well, reveal first sharp-based units fining upwards, 3–14 m thick, consisting of coarse-grained, occasionally pebbly sand, overlain by muddy, coal- and mica-containing, fine- to medium-grained sand that is laminated to homogeneous clay and coal seams with roots.[1] On most of the strata there is a common parallel lamination with subordinate cross-bedding, cross-lamination and Flaser lamination.[1] There are abundant large plant fragments and small bits of quartz. Marine palynomorphs are absent, suggesting that this level was deposited on a coastal or delta plain with fluvial channels, lakes and swamps.[6] This is consistent with finds in the German portion of the Ciechocinek Formation, where a large deltaic system ended: the large ToarcianBajocian deltaic systems were the local shoreline, influenced by the proximity between brackish to freshwater and continental biofacies.[9] [10] The North German Basin shows that on an approximately 14.4 m.y. gap, four third-order sea-level fluctuations led the subsequent formation of four individual delta generations in the Bifrons–Thouarsense (Toarcian), Murchisonae–Bradfordensis (Aalenian) and Humpresianum–Garatiana (Bajocian).[9] The Toarcian section was dominated by regressive elongated river-dominated deltas were due to the fall of the sea level the south to southwest directed delta progradation between the Lower and Upper Toarcian, that was deposited as 40 m of deltaic successions, found in places like Prignitz (east) and Brandenburg (north).[9] Most of the palynomorphs found in the Toarcian stratum are connected with ones found in the Sorthat Formation.[9]

Nearly 40 m thick, the upper section of the formation is composed mostly by a series of cross-bedded, cross-laminated, wave-rippled and bioturbated sand and heteroliths with sporadic syneresis cracks, pyrite nodules, the ichnofossils Planolites isp. and Teichichnus isp. and brackish to marine palynomorphs, mostly dinoflagellates. This upper part has a stratum more characteristic of nearshore environments with abundant lagoons, coastal lakes and fluvial channels, with the clean sand at the top probably representing a marine shoreface. The Korsodde section, 93 m thick, is composed mostly of coarse-grained sands with cross-bedding and parallel lamination, being mostly black due to an abundant organic debris. This section has been interpreted as part of the large local fluvial system, probably as a series of minor fluvial channels that were connected with coastal lakes and lagoons where riparian vegetation was abundant, judging by the presence of megaflora remains and palynomorphs. Small ichnofossil burrows and larger burrows, including Diplocraterion isp., are common, indicating that there was at least one subunit that was the fill of an estuarine channel. The uppermost part of the formation in the Korsodde section consists of fine-grained sands of yellowish to brown color with cross-stratification and parallel lamination, along with sandstones with thin bioturbated and wave-rippled heterolithic beds.

Profile

At Korsodde, the environment includes the following:

Stratigraphy of the Korsodde section!Unit !Lithology!Thickness (metres)!Type of environment!Fossil flora!Fossil fauna
Unit AYellow, weakly cemented muscovite quartz sandstone, medium- to fine-grained in the lower part, fine-grained in the upper part. Ripple or herringbone lamination is present in most of the beds, along discontinuous mudstone drapes around 0.5 cm thick and mudstone intraclasts. The mudstones show often ferruginization. A single thin horizon occurs at about 85 cm of the section and also a thin erosional surface with mudstones at 1 m. There is a layer of heterolithic deposits with fine-grained ripple mudstones and sandstones at 1.65–1.75 m.0.45–2.3 mEstuarine channel fill (upper or marginal, less energetic part)None recovered
Unit BIntercalations of muscovite quartz sandstones and dark mudstone drapes, with abundant heteroliths. In the vertical section, the sandstone layers (3 cm thick) are lenticular, with some displaying ripple cross- and herringbone lamination, and the mudstone drapes (0.5 cm thick) have wavy lamination. These last have a few laminae separated by thicker, coarser, mainly silty laminae showing abundant ferruginous cementation. There is a layer over B considered transitional to C.2.3–3.41 mUpper tidal flat deposits surrounding an estuaryNone recovered
  • Planolites isp.
  • Rosselia isp.
  • Palaeophycus isp.
  • B. tortuosus
  • D. parallelum
Unit CTwo main layers: a series of 20 cm dark mudstone with horizontal lamination and silt intercalations and a series of dark heteroliths with intercalated mudstones and ripple limestones. 3.41–3.7 mRestricted bay passing into upper tidal flat depositsNone recovered
  • D. parallelum
Unit DYellow ripple cross sandstone with abundant muscovite, alternating with continuous and discontinuous dark mudstone with abundant organic material. There are pyrite concretions in the lower part.3.7–4.7 mLower tidal flat within an estuaryRoots
  • Planolites isp.
  • Palaeophycus isp.
  • B. tortuosus
  • D. parallelum
Unit EMostly fine-grained sediments with abundant organic matter. Starts with 55 cm of muddy sandstone, dark at the beginning and light in the upper part. A bed of 5 cm of mudstone overlays the sandstone, followed by various levels of fine-grained sandstones interbedded with dark siltstone–mudstone, pyrite concretions and sandy mudstone. Over this is developed a massive coal layer containing Neocalamites stems where pyrite becomes more common. It is overlaid by mudstone and fine sandstone that turn into a poorly sorted yellow ferruginous layer. The upper part, 85 cm thick, is composed of mudstone with allochthonous Neocalamites stems and lignite clasts.4.7–6.9 mLagoonal environment above a coal bed
  • Neocalamites sp. stems
  • Coal
  • Plant cuticles
  • Roots
  • Root structures
Unit FMostly pale, fine-grained, ripple cross muddy sandstone and normal sandstone, separated by thin, pale sandy mudstones or thin mudstone drapes. Pyrite concretions and lignite clasts occur in the sandstones. There are synaeresis cracks noted at 8.15–8.75 m.6.9–9.9 mTidal flat deposits in an estuary
  • Lignites
  • Root structures
  • ?Thalassinoides isp.
  • ?Chondrites isp.
  • Rosselia isp.
  • Palaeophycus isp.
  • Planolites isp.
  • D. parallelum
Unit GA prominent erosional surface at the start, composed of yellow medium- to fine-grained cross-laminated sandstones with muscovite.9.9–11.35 mEstuarine barNone reportedNone reported
Unit HPale, fine-grained ripple and herringbone sandstones and mudstones, with intercalations of sandy mudstones and mudstone drapes with intense ferruginization, and some layers of mudstone–sandstone heteroliths11.35–14.2 mMarginal part of an estuary channel fillNone reported
  • Rosselia isp.
  • Palaeophycus isp.
  • Cylindrichnus isp.
  • Skolithos isp.
  • D. parallelum
Unit I, JBioturbated muddy sandstone14.2–14.4 mShort-lived bay or lagoon
  • Rosselia isp.
  • Teichichnus isp.
  • Teichichnus zigzag
  • Planolites isp.
  • Thalassinoides isp.
  • Palaeophycus isp.

Biota

The Sorthat Formation represents one of the most complete floras found in Europe dating to the PliensbachianToarcian boundary, as well as among Jurassic palynological deposits found worldwide.[4] [7] [8] [11]

Environment

Beyond the deposits on the west and south coast of Bornholm, the formation is present in the Stina-1 well, which belongs to the Rønne Graben (a large offshore pull-apart basin that also includes the westernmost fringe of the island of Bornholm), where both the Sorthat and the Bagå Formation are deposited on the hanging wall fault block close to the main eastern bounding fault of this graben along the west coast of the island. This graben was emerged during the deposition of the Sorthat Formation, as proven by the sand and clay with numerous coal horizons from the Stina-1 well.[12] The presence of a high kaolinite content in both coeval marine Danish Basin and local Bornholm, as well the abundant reworked Carboniferous palynomorphs, indicate significant erosion of a Carboniferous regolith, which was almost completely eroded by the Middle Jurassic. This suggests Pliensbachian–Toarcian rivers eroded the Bornholm High, eliminating all of the Carboniferous layers and leaving only older Palaeozoic strata, as proven by the granite of the younger Bagå Formation.[13] Due to a Late Pliensbachian marine regression, deposition of coal-bearing strata in the Sorthat Formation resumed on Bornholm until an Early Toarcian transgression terminated peat formation. The two main deposits of the formation, seen at the Levka-1 well and the lower part of the Korsodde section, were deposited in an environment influenced by the sea, the Levka location being populated by lagoons, lakes, channels and low fluvial areas. Then deposition of the Sorthat Formation in the Latest Pliensbachian–Toarcian demonstrated a rapid subsidence and relative sea level rise of the Rønne Graben, while the adjoining Arnager Block suffered a relative sea level fall. This is because the Rønne Graben experienced a rapid relative sea level rise during the Early Toarcian, coeval with the prominent rise registered in the Danish Basin. This peak transgression of the Ligurian Cycle is found in the coeval layers of the Fjerritslev Formation. The Bifrons to Levesquei zone in the coeval units at the east and west of Prignitz, a sandy coastal-deltaic succession, was replaced by laminated shales with pelagic marine fauna, reflected in the shoreline shifts to the northeast, which contributed to retrogradational stratal pattern architectures.[14] In the Sorthat Formation, a transition occurs from upper to lower shoreface environments, indicating a deepening trend. In the Younger Levesquei subzone, delta plain environments were replaced by shoreface setting with active bioturbation and hummocky cross-stratification.[14] The Rǿnne Graben shows seismic lines with onlapping patterns that have been correlated to these Lower Toarcian marine shoreface deposits with intense bioturbation.[15]

The depositional environments include the following:

Inertinite has been recovered from the coal-bearing levels of the formation, where the palynology shows that the mire vegetation may have been dominated by gymnosperms and also contained ferns characterised by the genera Dicksonia or Coniopteris and the family Osmundaceae. Biomolecules were found in several coal seams there, among which Euulminite and Attrinite were the most abundant huminite macerals recovered.[18] The Levka-1 well section represents fluvial channels, floodplain areas with shallow lakes and lagoons, and small crevasse deltas, with abundant coalified wood fragments and stems, most of them found associated with sandy channel fills and on heavily rooted crevasse and lake deposits in shallow inter-fluvial areas.[19] In the Toarcian at Bornholm, strata indicate a warm, humid climate suggested by the large number of plant species from the interconnected Jameson Land, and thin cutinised leaves of Podozamites and Equisetales comparable in size to modern subtropical bamboos are thought to reflect favourable conditions for plant growth.[19] There is abundant coal, which indicates that wildfires occurred in the bog.[19] Wood particles from this section, both charcoalified and unburned (coalified), with many particles being rounded and worn, imply the influence of greater transportation energies.[20]

Coal

On Korsodde, the Lower Toarcian section records higher temperatures and decreased rainfall and humidity, which led to an increase of the potential for local wildfires, reflected in the increased abundance of charcoal and burnt plants.[21] In the section at Korsodde that includes the Toarcian oceanic anoxic event, thermophilic plant taxa imply that the climate was relatively dry, and presence of micro- and macroscopic charcoal indicates a spike of abundance and increase of the wildfire activity.[22]

Most of the coal seams recovered from the formation come from Levka 1 and the Korsodde section, and are derived in most cases from a densely vegetated, anoxic swamp, which was probably rheotrophic and rich in nutrients. Study of the peat accumulation indicates that it occurred in rather short time intervals (around 2,300 years) and in a warm temperate to subtropical climate, falling short of the rate seen in tropical accumulations, such as the 1.8 mm/yr on the Batang Hari River in Sumatra. Peat accumulation of 1 mm/yr is equal to that of modern Central Kalimantan coastal settings.[23] The deposits have great amounts of thin and clean coal seams, covered by lacustrine–lagoonal flooding peaks, indicating rapid changes in the environment that were controlled by fairly rapid subsidence of the Rønne Graben, which along with eustatic rise in sea level caused decreases and increases in the base level at the coastal plain. The majority of the samples were immature, low-rank coals with generally very high content of humified organic matter, which indicates prevailing anoxic and fully saturated conditions during peat formation, with occasional inundations by freshwater that favoured humification of the plant tissues and also may have increased the gelification processes, raising the pH. Hopanoids are abundant and an indicator of common bacterial activity. The vegetation — both the nearby plants and those of the peat swamp — was probably small in stature, and its diversity suggests a humid, warm-temperate to subtropical climate that favoured prolific vegetation.

Phytoplankton

In the Lower Jurassic of Bornholm there were several successions of nearshore peat formations with dinoflagellates.[24] Coal-bearing strata were deposited in an overall coastal plain environment during the HettangianSinemurian, and then during the Early Pliensbachian deposition was interrupted until the late Pliensbachian–Lowermost Toarcian due to a sea regression.[24]

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Baltisphaeridium
  • Baltisphaeridium infulatum
  • Korsodde section
Very rare and limited to the middle layers
  • Cysts
An algal acritarch, probably related to freshwater red algae, similar to extant Florideophyceae (for example, Hildenbrandia) or Batrachospermales (Batrachospermum) and Thoreales.
Botryococcus
  • Botryococcus sp.
  • Korsodde section
  • Sorthat beds
  • Levka-1 borehole
Abundant to very abundant towards the upper sections
  • Miospores
Type genus of the Botryococcaceae in the Trebouxiales. A colonial green microalga of freshwater and brackish ponds and lakes around the world, where it often can be found in large floating masses. Sorthat Formation Botryococcus lived in an environment interpreted as a coastal lake, permanently vegetated and shallow, that was occasionally flooded by the sea.
Chomotriletes[27]
  • Chomotriletes minor
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Very rare and limited to the lower layers
  • Miospores
Affinities with the family Zygnemataceae. A genus derived from freshwater filamentous or unicellular, uniseriate (unbranched) green algae.
Crassosphaera
  • Crassosphaera coccinia
  • Crassosphaera hexagonalis
  • Korsodde section
Very rare and limited to middle section
  • Miospores
Affinities with the family Pycnococcaceae.
Cymatiosphaera
  • Cymatiosphaera sp.
  • Korsodde section
  • Sorthat beds
Very rare with peak in middle-upper layers in Levka-1 borehole; Very rare and limited to the lowermost layers in Sorthat beds
  • Miospores
Affinities with the family Pterospermopsidaceae.
Haplophragmoides
  • Haplophragmoides tryssa
  • Haplophragmoides platus
  • Haplophragmoides sp.
  • Sorthat beds
Common
  • Calcareous skeletons
A foraminifer, member of the family Lituoloidea in the Lituolida.
Korystocysta
  • Korystocysta sp.
  • Korsodde section
Very rare and only present in the middle section
  • Cysts
A dinoflagellate, member of the Cribroperidinioideae.
Lecaniella
  • Lecaniella foveata
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Rare to Abundant but limited to the lower-middle section
  • Miospores
Affinities with the family Zygnemataceae.
Leiosphaerida
  • Leiosphaerida spp.
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Very abundant either on the lower middle or upper sections, very rare or absent in all other layers
  • Miospores
Affinities with the family Prasinophyceae.
Luehndea
  • Luehndea spinosa
  • Korsodde section
Abundant but only present in the middle section
  • Cysts
A dinoflagellate, member of the Luehndeoideae. It establishes the Luehndea spinosa zone; the age of this zone is late Pliensbachian to early Toarcian.
Mancodinium
  • Mancodinium semitabulatum
  • Korsodde section
Very rare and only present in the middle section
  • Cysts
A dinoflagellate, type genus of the Mancodinioideae.
Mendicodinium
  • Mendicodinium groenlandicum
  • Mendicodinium reticulatum
  • Korsodde section
  • Sorthat beds
Rare to Abundant but limited to the lower-middle section
  • Cysts
A dinoflagellate, member of the family Gonyaulacales.
Micrhystridium
  • Micrhystridium fragile
  • Micrhystridium intromittum
  • Micrhystridium lymensis
  • Micrhystridium spp.
  • Korsodde section
  • Levka-1 borehole
Abundant but limited to the middle section
  • Cysts
An acritarch, familia incertae sedis
Nannoceratopsis
  • Nannoceratopsis senex
  • Nannoceratopsis gracilis
  • Nannoceratopsis ridingui
  • Nannoceratopsis triangulata
  • Nannoceratopsis triceras
  • Nannoceratopsis dictyanbonis
  • Nannoceratopsis sp.
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Abudant in the lower section in Levka-1; Very abundant in the middle-upper sections in Sorthat beds and Korsodde
  • Cysts
A dinoflagellate, member of the family Nannoceratopsiaceae. It is characteristic of marine deposits. The presence of N. gracilis, N. senex and N. triceras, and common occurrence of Botryococcus is interpreted as indicating a lagoon succession overlying a transgressive surface and signals a rise in relative sea level.
Ovoidites
  • Ovoidites sp. A
  • Ovoidites sp. B
  • Ovoidites sp. C
  • Ovoidites spp.
  • Korsodde section
  • Levka-1 borehole
Very rare with peaks in the middle layers
  • Miospores
Affinities with the family Zygnemataceae. A genus derived from freshwater filamentous or unicellular, uniseriate (unbranched) green algae.
Pterospermella
  • Pterospermella spp.
  • Korsodde section
Very rare and limited to middle section
  • Miospores
Affinities with the family Pterospermataceae.
Rotundus
  • Rotundus granulatus
  • Sorthat beds
Abundant to very abundant but limited to the lower section
  • Miospores
An algal palynomorph unique to the setting and probably related to freshwater red algae; similar to extant Batrachospermales.
Spirillina
  • Spirillina sp.
  • Sorthat beds
Rare
  • Calcareous skeletons
A foraminifer, type genus of the Spirillinidae in the Spirillinida.
Striatella
  • Striatella jurassica
  • Striatella parva
  • Striatella seebergensis
  • Striatella scanica
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Very rare to abundant in the upper sections
  • Miospores
Brown algae, type genus of the family Striatellaceae in the Striatellales. These brown algae diatoms are associated with either brackish or marginal marine environments.
Tasmanites
  • Tasmanites sp.
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Abudant to very rare, limited to the middle layers
  • Miospores
Affinities with the family Pyramimonadaceae. Found on shoreface and shoreface–offshore transition zone deposits.
Tetraporina
  • Tetraporina compressa
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Abundant but limited to the lowermost layer in Levka-1 borehole; Abudant to rare in Sorthat beds & Korsodde
  • Miospores
Affinities with the family Zygnemataceae.
Veryhachium
  • Veryhachium spp.
  • Levka-1 borehole
Very rare and limited to the lower layers
  • Cysts
A dinoflagellate, member of the Dinophyceae.

In Early Toarcian carbonates, local bulk organic matter and wood fragments have been associated with carbon cycle perturbations, shedding light on the reaction of the continental biota to the Toarcian oceanic anoxic event, which accompanied large-scale volcanism.[28] There are several changes to the woody vegetation in the wood-derived carbon, with pollen assemblages dominated by pollen types in the Sciadopityaceae and Miroviaceae, such Cerebropollenites associated with cycad pollen types (Chasmatosporites) and the hirmeriellaceous Corollina.[28] The local palynology has shown the terrestrial changes of the local flora. In the Pliensbachian the dominant palynofacies were ones in the Cupressaceae such as Perinopollenites, along with cycads such as Cycadopites, found in mid-latitude Mediterranean climates.[29] Then, at the start of the event the local pollen assemblages show a shift to spore-rich layers, showing a long-term increase in ferns and lycophytes, an indicator of more humid conditions.[29] Finally, after the Toarcian anoxic event, the Sorthat Formation showed an abrupt rise of pollen of Hirmeriellaceae such as Corollina and specially Spheripollenites, both indicators of semidesertic to dry Mediterranean climates, implying an abrupt warming event coeval with the changes happening at sea.[29] The main deposits of macroflora are the Hasle clay pit and the Korsodde section. The flora was originally stated to be coeval with the RhaetianHettangian floras of Sweden, but found later to be Pliensbachian–Toarcian. Möller did the two major studies on the local flora, with 68 species described, 50% of them ferns. The Late Pliensbachian section is dominated by ferns, suggesting a warm and humid climate, which fits with the palaeolatitude of Bornholm, firmly within the Jurassic warm biome. But the presence of Ginkoaleans and absence of large-leafed Bennettites suggest this warm climate was seasonal. Ferns and sphenophytes in the assemblage are interpreted to have occupied the forest floor. Bennettites were mid-storey shrubs, and conifers, such as Pagiophyllum, together with ginkgoaleans, make up the main arboreal flora. All the flora developed on a meandering river system with well-vegetated marshy flood plains. The Toarcian section shows a radical change on the local flora, as Hirmeriellaceae conifers take over the role of dominant flora, representing 95% of the pollen recovered, along with the rise of seed ferns, Bennettites and Czekanowskiales. The dominance of Pagiophyllum and its related pollen Corollina torosus indicate high temperature and aridity with seasonal wildfires (though some sections show a low coal ratio and are derived from slightly more humid environments), with rare occurrences of Brachyphyllum and one Cyparissidium. Is also common to found wood from the nearshore deposits of Korsodde, with two sets: macroscopic wood, recognizable to the naked eye, up to branch-sized; and microscopic wood (0.25 to 1 mm average dimension).[30]

Bryophyta

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Cingutriletes
  • Cingutriletes infrapunctus
  • Cingutriletes oculus
  • Korsodde section
Very rare and present in certain intervals
  • Spores
Incertae sedis; affinities with Bryophyta. This spore is found in Jurassic sediments associated with the polar regions. The Sorthat Formation is among its southernmost locations.
Foraminisporis
  • Foraminisporis jurassicus
  • Korsodde section
Very rare and only in a few layers, with a few layers of very abundant presence in the middle
  • Spores
Affinities with the family Notothyladaceae in the Anthocerotopsida. Hornwort spores.
Polycingulatisporites
  • Polycingulatisporites circulus
  • Polycingulatisporites liassicus
  • Polycingulatisporites triangularis
  • Levka-1 borehole
  • Korsodde section
Very rare and only in the middle layers
  • Spores
Affinities with the family Notothyladaceae in the Anthocerotopsida. Hornwort spores.
Sculptisporis
  • Sculptisporis aulosenensis
  • Sorthat beds
  • Korsodde section
Abudant in the lowermost layer to very rare or absent in the upper ones
  • Spores
Affinities with the family Sphagnaceae in the Sphagnopsida.
Staplinisporites
  • Staplinisporites caminus
  • Korsodde section
Very rare and only in a few layers of very abundant presence in the uppermost section
  • Spores
Affinities with the family Encalyptaceae in the Bryopsida. Branching moss spores, indicating high water-depleting environments.
Stereisporites
  • Stereisporites antiquasporites
  • Stereisporites stereoides
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare with an abundant peak in the middle in Levka-1; very rare and limited to lower layers in Sorthat beds and Korsodde
  • Spores
Affinities with the family Sphagnaceae in the Sphagnopsida. "Peat moss" spores, related to genera such as Sphagnum that can store large amounts of water.
Taurocusporites
  • Taurocusporites verrucatus
  • Korsodde section
Very rare and only in a few layers, with a few layers of very abundant presence in the middle
  • Spores
Affinities with the family Sphagnaceae in the Sphagnopsida.
Rogalskaisporites
  • Rogalskaisporites cicatricosus
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare in the lower layers and absent in the youngest layers, with peak moderately abundant in the middle in Levka-1; very rare and limited to lower middle and uppermost layers in Sorthat beds and Korsodde
  • Spores
Affinities with the family Sphagnaceae in the Sphagnopsida.

Lycophyta

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Anapiculatisporites
  • Anapiculatisporites spiniger
  • Anapiculatisporites sp.
  • Levka-1 borehole
  • Korsodde section
Peak in the upper middle, very abundant to very rare in the lowermost and uppermost layers in Levka-1; absent to abundant in upper layers in KorsoddeSporesAffinities with the Selaginellaceae in the Lycopsida. Herbaceous lycophyte flora, similar to ferns, found in humid settings. This family of spores are also the most diverse in the formation.
Cadargasporites
  • Cadargasporites granulatus
  • Korsodde section
Abundant only in the middle upper section; absent in all other levelsSporesAffinities with the Selaginellaceae in the Lycopsida.
Camarozonosporites
  • Camarozonosporites golzowensis
  • Camarozonosporites rudis
  • Korsodde section
Very rare SporesAffinities with the family Lycopodiaceae in the Lycopodiopsida.
Kraeuselisporites
  • Kraeuselisporites reissingeri
  • Korsodde section
Very rare and only in a few layers, with an exceptional peak in the middle upper sectionSporesAffinities with the Selaginellaceae in the Lycopsida.
Neoraistrickia
  • Neoraistrickia gristhorpensis
  • Neoraistrickia sp.
  • Levka-1 borehole
  • Sorthat beds
  • Korsode section
Very rare and only in the middle layersSporesAffinities with the Selaginellaceae in the Lycopsida.
Retitriletes
  • Retitriletes austroclavatidites
  • Retitriletes clavatoides
  • Retitriletes semimurus
  • Retitriletes spp.
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Very rare in the lower layers to moderately abundant in the upperSporesAffinities with the family Lycopodiaceae in the Lycopodiopsida.
Selaginellites[31]
  • Selaginellites falcatus
  • Hasle clay pit
Limited to a few specimensFine stemsAffinities with Selaginellaceae and Lycopodiidae in the Lycopodiales. It was originally described as Lycopodites falcatus. The leaves of this species are more prominently anisophyllous than in the Raheto-Hettangian S. coburgensis from Franconia.[32]
Sestrosporites
  • Sestrosporites pseudoalveolatus
  • Sorthat beds
  • Korsodde section
Very rare and limited to the lowermost layerSporesAffinities with the family Lycopodiaceae in the Lycopodiopsida. Lycopod spores, related to herbaceous to arbustive flora common in humid environments.
Uvaesporites
  • Uvaesporites argenteaeformis
  • Uvaesporites microverrucatus
  • Uvaesporites puzzlei
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare and in a few samples in Levka-1; abundant but only in the lowermost layer in Sorthat beds; very rare in Korsodde sectionSporesAffinities with the Selaginellaceae in the Lycopsida.

Equisetales

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Calamospora
  • Calamospora tener
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Abudant in the upper section, rare to not present in the underliying layers in Levka-1; very rare and only in the middle in Sorthat beds; abundant to very abundant in the middle Korsodde sectionSporesAffinities with the Calamitaceae in the Equisetales. Horsetails are herbaceous flora found in humid environments and are flooding-tolerant. In the sections of the formation such as Korsodde, this genus has small peaks in abundance in the layers where more Equisetites stems are found.
Equisetites
  • Equisetites munsteri
  • Equisetites lyelli
  • Equisetites sp.
  • Bagagraven clay pit
  • Nebbeodde
  • Stina-1 well
Extremely CommonStemsAffinities with Equisetaceae in the Equisetales. Related equisetalean stems are found in the Hettangian strata along Skane, Sweden. In the lagoonar sections there is correlation between bioturbation and transported Equisetites stems. Local Equisetales reached a considerable size, comparable to modern subtropical bamboos, close to lakes and in the wettest environments.
Neocalamites[33] [34] [35] [36] [37] [38]
  • Neocalamites hoerensis
  • Neocalamites sp.
  • Korsodde section
  • Bagagraven clay pit
Rare
  • Three incomplete axes
  • Isolated Incomplete fragments
Affinities with Calamitaceae in the Equisetales. Related equisetalean stems are found in strata of the same age along Skane, Sweden. Based on analogies with morphologically similar extant Equisetum species, it is interpreted to represent a plant of consistently moist habitats, such as marshes, lake margins or forest understorey, normally developing dense thickets.
Phyllotheca
  • Phyllotheca cf. equisetiformis
  • Hasle clay pit
RareLeaf whorlsAffinities with Equisetidae in the Equisetales.

Pteridophyta

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Annulispora
  • Annulispora folliculosa
  • Korsodde section
Very rare and only in the lower layersSporesAffinities with the genus Saccoloma, type representative of the family Saccolomataceae. This fern spore resembles those of the living genus Saccoloma, being probably from a pantropical genus found in wet, shaded forest areas.
Baculatisporites
  • Baculatisporites comaumensis
  • Baculatisporites primarius
  • Baculatisporites wellmanii
  • Baculatisporites sp.
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Moderately abundantSporesAffinities with the family Osmundaceae in the Polypodiopsida. Near fluvial current ferns, related to the modern Osmunda regalis.
Cladophlebis
  • Cladophlebis nebbensis
  • Cladophlebis roesserti
  • Cladophlebis svedbergii
  • Cladophlebis hirta
  • Vellengsby
  • Bagagraven clay pit
  • Hasle clay pit
  • Nebbeodde
Abudant
  • Isolated pinnae
Affinities with Osmundaceae in the Osmundales. Related to species commonly reported from the Triassic–Jurassic of southern Sweden.
Cladotheca[39]
  • Cladotheca undans
  • Bagagraven clay pit
Rare
  • Fertile pinna fragments
Affinities with Osmundaceae in the Osmundales. Specimens assigned to this morphothype have been found in the Middle Jurassic flora of Yorkshire, associated with Todites miospores, and were originally described as Asplenites cladophleboides.
Cibotiumspora
  • Cibotiumspora jurienensis
  • Sorthat beds
  • Korsodde section
Very rare and only in a few upper layersSporesAffinities with the family Cyatheaceae in the Cyatheales. Arboreal fern spores.
Clathropteris
  • Clathropteris meniscioides
  • Clathropteris platyphylla
  • Bagagraven clay pit
  • Hasle clay pit
  • Nebbeodde
Abudant
  • Isolated pinnae
Affinities with Dipteridaceae in the Polypodiales.
Conbaculatisporites
  • Conbaculatisporites mesozoicus
  • Conbaculatisporites spinosus
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Moderately to abudant in the middle layers in Levka-1; very rare and only in middle to upper layers in Sorthat beds and KorsoddeSporesIncertae sedis; affinities with the Pteridophyta
Coniopteris
  • Coniopteris hymenophylloides
  • Coniopteris acutidens
  • Bagagraven clay pit
RareIncomplete frond fragmentAffinities with Polypodiales in the Polypodiidae. Common cosmopolitan Mesozoic fern genus. Recent research has reinterpreted it a stem group of the Polypodiales (closely related to the extant genera Dennstaedtia, Lindsaea, and Odontosoria).[40]
Deltoidospora
  • Deltoidospora minor
  • Deltoidospora toralis
  • Deltoidospora spp.
  • Korsodde section
  • Levka beds
  • Sorthat beds
Abudant throughout the interval in Levka-1; moderately common under to abundant in the upper Sorthat beds; rare or not present in lower to moderlately common in upper KorsoddeSporesIncertae sedis; affinities with the Pteridophyta
Dicksonia
  • Dicksonia pingelii
  • Dicksonia pauciloba
  • Bagagraven clay pit
  • Hasle clay pit
RareLeafletsAffinities with Dicksoniaceae in the Cyatheales. It show similarities with Sphenopteris longipinnata in the morphological outline of the leaflets and the keels of the pinnate axis.
Dictyophyllum
  • Dictyophyllum acutilobium
  • Dictyophyllum munsteri
  • Dictyophyllum barthollini
  • Dictyophyllum cf. nilssonii
  • Dictyophyllum cf. spectabile
  • Vellengsby
  • Bagagraven clay pit
  • Hasle clay pit
Abudant
  • Isolated pinnae
Affinities with Dipteridaceae in the Polypodiales. Dictyophyllum is a common dipteridacean genus of the mid-Mesozoic.
Eboracia
  • Eboracia lobifolia
  • Eboracia sp.
  • Bagagraven clay pit
  • Hasle clay pit
  • Vellengsby
Dominant
  • Isolated pinnae
Affinities with Dicksoniaceae in the Cyatheales. The Lund material is dominated by ferns belonging to the genus Eboracia (28 specimens of E. lobifolia and 14 of another Eboracia sp.). The latter has smaller pinnules than E. lobifolia.
Gleicheniidites
  • Gleicheniidites senonicus
  • Levka-1 borehole
  • Korsodde section
Very rare and in a few samplesSporesAffinities with the Gleicheniales in the Polypodiopsida. Fern spores from low herbaceous flora.
Gutbiera
  • Gutbiera angustiloba
  • Vellengsby
  • Nebbeodde
Very rareIsolated pinnaeAffinities with Matoniaceae in the Gleicheniales.
Hausmannia
  • Hausmannia crenata
  • Hausmannia dichotoma
  • Hausmannia dentata
  • Hausmannia lasciniata
  • Hausmannia acutidens
  • Bagagraven clay pit
  • Hasle clay pit
AbudantIsolated pinnaeAffinities with Dipteridaceae in the Polypodiales. Specimens from the same species have been found in the Hettangian Höör Sandstone at southern Sweden.
Intrapunctisporis
  • Intrapunctisporis toralis
  • Sorthat beds
Very rare and only in a few upper layersSporesIncertae sedis; affinities with the Pteridophyta
Iraqispora
  • Iraqispora labrata
  • Korsodde section
Very rare and only in a few layers with a few layers of very abundant presence in the middleSporesAffinities with the Gleicheniales in the Polypodiopsida. Fern spores from low herbaceous flora.
Ischyosporites
  • Ischyosporites crateris
  • Ischyosporites variegatus
  • Levka-1 borehole
  • Korsodde section
Very rare and only in a few layersSporesIncertae sedis; affinities with the Pteridophyta
Klukisporites
  • Klukisporites lacunus
  • Korsodde section
Very rare and only in a few layersSporesAffinities with the family Lygodiaceae in the Polypodiopsida. Climbing fern spores.
Laevigatosporites
  • Laevigatosporites mesozoicus
  • Korsodde section
  • Levka-1 borehole
Rare and in concrete samples in Korsodde; very rare and only in the middle in Levka-1SporesIncertae sedis; affinities with the Pteridophyta
Leptolepidites
  • Leptolepidites bossus
  • Leptolepidites macroverrucosus
  • Leptolepidites major
  • Leptolepidites sp.
  • Korsodde section
Very rare and only in a few layersSporesAffinities with the family Dennstaedtiaceae in the Polypodiales. Forest fern spores.
Lycopodiacidites
  • Lycopodiacidites infragranulatus
  • Lycopodiacidites infragranulatus
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare and in concrete samples in Levka-1 & Sorthat beds; abundant but limited to lower layers in KorsoddeSporesAffinities with the Ophioglossaceae in the Filicales. Fern spores from lower herbaceous flora.
Manumia
  • Manumia delcourtii
  • Levka-1 borehole
  • Sorthat beds
  • Kosodde section
Abundant in the lower middle section, very rare in upper Levka-1; very rare and only in the middle in Sorthat bedsSporesAffinities with the Pteridaceae in the Polypodiopsida. Forest ferns from humid ground locations.
Marattia
  • Marattia munsteri
  • Vellengsby
Very rareIsolated pinnaeAffinities with Marattiaceae in the Marattiopsida.
Marattisporites
  • Marattisporites scabratus
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare and in the lower layers only in Levka-1; very rare but also in upper layers in Sorthat beds and KorsoddeSporesAffinities with the Marattiaceae in the Polypodiopsida. Fern spores from low herbaceous flora.
Phlebopteris
  • Phlebopteris schouwii
  • Phlebopteris elegans
  • Phlebopteris mirovensis
  • Phlebopteris woodwardii
  • Phlebopteris affinis
  • Phlebopteris polypodioides
  • Vellengsby
  • Bagagraven clay pit
  • Hasle clay pit
  • Nebbeodde
Abundant
  • Isolated pinnae
Affinities with Matoniaceae in the Gleicheniales.
Skarbysporites
  • Skarbysporites crassexinius
  • Sorthat beds
  • Korsodde section
Very rare and only in a few layersSporesIncertae sedis; affinities with the Pteridophyta
Spiropteris
  • Spiropteris sp.
  • Bagagraven clay pit
Rare
  • Single impression
Incertae ordinis in the Pteridophyta. Spiropteris is the name given to the fossil of a coiled, unopened fern leaf.
Tigrisporites
  • Tigrisporites halleinis
  • Tigrisporites microrugulatus
  • Sorthat beds
  • Korsodde section
Very rare and only in a few uppermost layersSporesIncertae sedis; affinities with the Pteridophyta
Thaumatopteris
  • Thaumatopteris brauniana
  • Vellengsby
  • Bagagraven clay pit
Very rareIsolated pinnaeAffinities with Dipteridaceae in the Polypodiales.
Todisporites
  • Todisporites major
  • Todisporites minor
  • Levka-1 borehole
  • Sorthat beds
Very rare and only in the middle layers in Levka-1; very rare and only in the uppermost section in Sorthat bedsSporesAffinities with the family Osmundaceae in the Polypodiopsida.
Tripartina
  • Tripartina variabilis
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare down to moderately abundant in Levka-1; very rare and only in upper layers in Sorthat beds and Korsodde
  • Spores
Affinities with the genus Dicksoniaceae in the Polypodiopsida. Tree fern spores.
Verrucosisporites
  • Verrucosisporites obscurilaesuratus
  • Korsodde section
Very rare and limited to the middleSporesIncertae sedis; affinities with the Pteridophyta
Vesicaspora
  • Vesicaspora fuscus
  • Korsodde section
Only in the uppermost layers and very rareSporesAffinities with the Callistophytaceae in the Callistophytales. Spores from large arboreal to arbustive ferns.
Zebrasporites
  • Zebrasporites interscriptus
  • Korsodde section
Very rare and only in the lowermost layersSporesAffinities with the family Cyatheaceae in the Cyatheales. Arboreal fern spores.

Peltaspermales

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Alisporites
  • Alisporites grandis
  • Alisporites radialis
  • Alisporites robustus
  • Alisporites thomasii
  • Alisporites microsaccus
  • Alisporites diaphanus
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare and only in the lower to middle layers in Levka-1; abundant to very abundant in Sorthat beds and KorsoddePollenAffinities with the families Peltaspermaceae, Corystospermaceae or Umkomasiaceae in the Peltaspermales. Pollen of uncertain provenance that can be derived from any of the members of the Peltaspermales. The lack of distinctive characters and poor conservation make this pollen difficult to classify. Arboreal to arbustive seed ferns.
Carpolithes
  • Carpolithes cinctus
  • Carpolithes nebbensis
  • Carpolithes nummularius
  • Vellengsby
  • Bagagraven clay pit
  • Nebbeodde
Common
  • Plant propagules
Plant propagules that may be from Pteridospermatophyta, Vladimariales, Bennettitales or Pinales. Fruits or seeds of uncertain placement.
Ctenozamites[41]
  • Ctenozamites leckenbyi
  • Bagagraven clay pit
Very rare
  • Isolated pinnae
Affinities with Umkomasiaceae in the Pteridospermatophyta.
Cycadopteris
  • Cycadopteris heterophylla
  • Cycadopteris brauniana
  • Bagagraven clay pit
Very rareIsolated pinnaeAffinities with Corystospermaceae in the Pteridospermatophyta.
Feildenia
  • Feildenia cuspiformis
  • Hasle clay pit
Very rareLeaf compressionsAffinities with Umaltolepidaceae in the Vladimariales. These belong to a group parallel to Gingkoaceans and derived probably from Umkomasiaceae.
Kekryphalospora
  • Kekryphalospora distincta
  • Sorthat beds
  • Levka-1 borehole
  • Korsodde section
Very rare and in concrete layersPollenAffinities with the families Peltaspermaceae, Corystospermaceae or Umkomasiaceae in the Peltaspermales.
Komlopteris[42]
  • Komlopteris nordenskioeldii
  • Vellengsby
  • Korsodde section
Very rareIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta.
Pachypteris
  • Pachypteris laceolata
  • Pachypteris papillosa
  • Vellengsby
  • Korsodde section
Very rareIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta. Less common than other arboreal plants.
Ptilozamites
  • Ptilozamites falcatus
  • Ptilozamites cycadea
  • Bagagraven clay pit
CommonIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta.
Pteridospermae
  • Pteridospermae indet.
  • Korsodde section
Very rare
  • Coalified fragments
  • Cuticles
Affinities with Pteridospermae in the Pteridospermatophyta.
Sagenopteris[43]
  • Sagenopteris cuneata
  • Sagenopteris phillipsi
  • Sagenopteris rhoifolia
  • Sagenopteris nilssoniana
  • Sagenopteris undulata
  • Sagenopteris sp.
  • Vellengsby
  • Bagagraven clay pit
CommonIsolated pinnaeAffinities with Caytoniaceae in the Pteridospermatophyta. Related to seed ferns present in the Rhaetic flora of Sweden.
Vitreisporites
  • Vitreisporites bjuvensis
  • Vitreisporites pallidus
  • Levka-1 borehole
  • Korsodde section
Very rare and only in the lower layersPollenFrom the family Caytoniaceae in the Caytoniales. Caytoniaceae are a complex group of Mesozoic fossil floras that may be related to both Peltaspermales and Ginkgoaceae.

Cycadophyta

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Butefia[45]
  • Butefia ensiformis
  • Bagagraven clay pit
Rare
  • Leaflets
Affinities with Cycadales in the Cycadopsida. Originally described as Podozamites ensiformis.
Chasmatosporites
  • Chasmatosporites apertus
  • Chasmatosporites elegans
  • Chasmatosporites hians
  • Chasmatosporites major
  • Chasmatosporites minor
  • Levka-1 borehole
  • Korsodde section
Abundant lower to very abundant upper
  • Pollen
Affinities with the family Zamiaceae in the Cycadales. It is among the most abundant flora recovered on the upper section of the coeval Rya Formation, and was found to be similar to the pollen of the extant Encephalartos laevifolius.
Clavatipollenites
  • Clavatipollenites hughesii
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Abudant but limited to lower layers in Levka-1; abundant to very abundant in Sorthat beds and Korsodde
  • Pollen
Affinities with the family Cycadaceae in the Cycadales. The structure of the exine of Clavatipollenites hughesii from Jurassic deposits is fundamentally different from that of Cretaceous grains referred to the same species, confirming observations made previously on the basis of analysis under the light microscope and suggesting a possible derivation from cycadalean rather than angiospermous plants.[46]
Ctenis
  • Ctenis nathorsti
  • Hasle clay pit
RareLeafletsAffinities with Cycadales in the Cycadopsida.
Cycadales
  • Cycadales Indet.
  • Korsodde section
Rare
  • Coalified fragments
  • Cuticles
Affinities with Cycadidae in the Cycadopsida.

Bennettitales

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Cycadopites
  • Cycadopites nitidus
  • Cycadopites andrewsii
  • Korsodde section
Very abundant but present only in the uppermost sections
  • Pollen
Affinities with the family Cycadaceae and Bennettitaceae. It has been found associated with the Bennetite pollen cone Bennettistemon. It increases towards the Toarcian section.
Dictyozamites[47]
  • Dictyozamites johnsirupi
  • Bagagraven clay pit
Rare
  • Leaflets
Affinities with Williamsoniaceae in the Bennettitales.
Nilssonia
  • Nilssonia polymorpha
  • Nilssonia münsteri
  • Nilssonia acuminata
  • Vellengsby
  • Bagagraven clay pit
  • Nebbeodde
Very abundant
  • Leaflets
Affinities with Cycadeoidaceae in the Bennettitales. The most common and abundant bennetite on the formation.
Nilssoniopteris[48]
  • Nilssoniopteris tenuinervis
  • Nilssoniopteris glandulosa
  • Bagagraven clay pit
  • Hasle clay pit
Very abundant
  • Leaflets
Affinities with Cycadeoidaceae in the Bennettitales.
Otozamites
  • Otozamites bornholmiensis
  • Otozamites latior
  • Otozamites bartholini
  • Otozamites tenuissimus
  • Otozamites bunburyanus
  • Otozamites obtusus
  • Otozamites pusillus
  • Otozamites beani
  • Otozamites pterophylloides
  • Otozamites molinianus
  • Otozamites cf. reglei
  • Otozamites cf. mimetes
  • Vellengsby
  • Bagagraven clay pit
  • Stina-1 well
Dominant
  • Leaflets
Affinities with Williamsoniaceae in the Bennettitales. Insufficient and incomplete material prevents certain assignment of Otozamites cf. reglei and Otozamites cf. mimetes
Pterophyllum
  • Pterophyllum tenuicaule
  • Pterophyllum carnallianum
  • Pterophyllum cf. aequale
  • Pterophyllum cf. braunianum
  • Vellengsby
  • Bagagraven clay pit
Very abundant
  • Leaflets
Affinities with Williamsoniaceae in the Bennettitales.
Williamsonia
  • Williamsonia forchhammeri
  • Nebbeodde
Rare
  • Bennettitalean "flower"
Affinities with Williamsoniaceae in the Bennettitales.

Ginkgoales

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Baiera
  • Baiera czekanowskiana
  • Baiera pulchella
  • Baiera sp.
  • Korsodde section
  • Bagagraven clay pit
  • Vellengsby
Common
  • Leaf compressions
  • Cuticles
Affinities with Karkeniaceae in the Ginkgoales. Unlike other plant specimens from the location, it is more characteristic of Middle Jurassic flora.
Czekanowskia
  • Czekanowskia hartzii
  • Czekanowskia cf. setacea
  • Korsodde section
  • Hasle clay pit
  • Vellengsby
Common
  • Leaf compressions
  • Coalified fragments
  • Cuticles
Affinities with Czekanowskiales in the Ginkgoales. This genus is related to flora from the Rhaetian–Hettangian boundary of Jameson Land, but also present in Romania.
Hartzia
  • Hartzia tenuis
  • Hartzia sp.
  • Korsodde section
Rare
  • Leaf compressions
  • Coalified fragments
Affinities with Czekanowskiales in the Ginkgoales. Linked to the Lower Liassic flora of Greenland.
Ginkgoites
  • Ginkgoites troedssonii
  • Ginkgoites sibirica
  • Ginkgoites obovata
  • Korsodde section
  • Bagagraven clay pit
  • Hasle clay pit
  • Nebbeodde
Common
  • Leaf compressions
  • Coalified fragments
  • Cuticles
Affinities with Ginkgoaceae in the Ginkgoales. Seven species assigned to either Ginkgo or Ginkgoites have been reported from Latest Triassic to middle Jurassic strata of southern Sweden.
Ginkgoales
  • Ginkgoales indet.
  • Korsodde section
Rare
  • Coalified fragments
  • Cuticles
Affinities with Ginkgoales in the Ginkgoopsida.
Monosulcites
  • Monosulcites minimus
  • Monosulcites punctatus
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Abundant but limited to the middle layers
  • Pollen
Affinities with the family Karkeniaceae and Ginkgoaceae in the Ginkgoales. Had been considered pollen of Chloranthaceae but is likely from Ginkgoales, which can have similar features
Solenites
  • Solenites murrayana
  • Korsodde section
Rare
  • Leaf compressions
  • Coalified fragments
Affinities with Czekanowskiales in the Ginkgoales. This species was described on the basis of individuals collected in Greenland from the Triassic–Jurassic boundary.

Coniferophyta

GenusSpeciesStratigraphic positionAbundanceMaterialNotesImages
Agathoxylon
  • Agathoxylon württembergica
  • Bagagraven clay pit
  • Isolated logs
  • Isolated branches
  • Isolated indeterminate woody material
  • Coalified fragments
AbundantAffinities with Hirmeriellaceae or Araucariaceae in the Pinales. Originally Araucarioxylon württembergica. This genus is usually associated with leaf-bearing twigs referred to as Pagiophyllum, abundant in the Sorthat Formation.
Araucariacites
  • Araucariacites australis
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very rare and only in upper layers
  • Pollen
Affinities with Araucariaceae in the Pinales.
Bartholinodendron
  • Bartholinodendron punctulatum
  • Bagagraven clay pit
  • Hasle clay pit
Rare
  • Fragmentary axis compressions with preserved leaves
Affinities with Taxaceae in the Pinales. Was first identified in Bornholm. Is similar to the cretaceous Taxus huolingolensis and extant Taxus in leaf gross morphology and has papillate abaxial cuticles, probably being close to this genus.[49]
Brachyphyllum
  • Brachyphyllum mamillare
  • Brachyphyllum sp.
  • Vellengsby
  • Korsodde section
Abundant
  • Fragmentary axis compressions with preserved leaves
  • Coalified fragments
  • Cuticles
Affinities with Araucariaceae or Hirmeriellaceae in the Pinales. Is related to the Hettangian axis found in Scania, Sweden
Callialasporites
  • Callialasporites dampieri
  • Callialasporites turbatus
  • Callialasporites microvelatus
  • Callialasporites segmentatus
  • Sorthat beds
  • Korsodde section
Very rare and only in a few layers
  • Pollen
Affinities with the family Araucariaceae in the Pinales. Conifer pollen from medium to large arboreal plants.
Cerebropollenites
  • Cerebropollenites macroverrucosus
  • Cerebropollenites thiergartii
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Abundant to very abundant
  • Pollen
Affinities with both Sciadopityaceae and Miroviaceae in the Pinopsida. This pollen's resemblance to extant Sciadopitys suggest that Miroviaceae may be an extinct lineage of Sciadopityaceae-like plants.[50]
Coniferales
  • Coniferales indet.
  • Korsodde section
Very abundant
  • Coalified fragments
  • Cuticles
Affinities with Coniferales in the Coniferopsida.
Corollina
  • Corollina torosus
  • Corollina meyeriana
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Very abundant, but with intercalations of layers of total absence in Levka-1; very abundant and almost dominant in some samples in Sorthat beds and KorsoddePollenAffinities with the Hirmeriellaceae in the Pinopsida.
Cyparissidium
  • Cyparissidium blackii
  • Korsodde section
Very Rare
  • Coalified fragment
Affinities with Cupressoideae in the Cupressales. It matches with the Middle Jurassic Cyparissidium blackii from Yorkshire, England.
Dactylethrophyllum
  • Dactylethrophyllum ramonensis
  • Korsodde section
Dominant, up to 95%
  • Cuticles
Affinities with Hirmeriellaceae in the Pinales. It is related to other representatives of the genus of the Toarcian of Italy and Lower Jurassic of Israel. Spheripollenites co-occurs with cuticles of Dactylethrophyllum ramonensis, and the species S. psilatus may be produced by the conifer genus Dactylethrophyllum.
Elatocladus[51]
  • Elatocladus subzamioides
  • Bagagraven clay pit
Rare
  • Fragmentary axis compressions with preserved leaves
Affinities with Thujaceae in the Cupressales. It was originally described as Taxites? subzamioides, later merged with Elatocladus.
Exesipollenites
  • Exesipollenites tumulus
  • Korsodde section
Very rare and limited to upper layersPollenAffinities with the family Cupressaceae in the Pinopsida. Pollen that resembles that of extant genera such as the genus Actinostrobus and Austrocedrus, probably derived from dry environments.
Hirmeriella
  • Hirmeriella münsteri
  • Bagagraven clay pit
Rare
  • Ovuliferous Cones
Affinities with Hirmeriellaceae in the Pinales. The main genus of the Hirmeriellaceae, found in dry environments and probably fire tolerant.
Quadraeculina
  • Quadraeculina anellaeformis
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Rare to very rare and limited to the lower middle and uppermost layers in Levka-1; peak of abundance in middle layers in Sorthat beds and KorsoddePollenAffinities originally suggested with the family Podocarpaceae in the Pinopsida. Quadraeculina is not comparable to pollen of any modern gymnosperm family.
Lindleycladus[52]
  • Lindleycladus lanceolatus
  • Hasle clay pit
  • Bagagraven clay pit
Rare
  • Isolated leaves
Affinities with Krassiloviaceae in the Voltziales.
Marskea[53]
  • Marskea jurassica
  • Bagagraven clay pit
Rare
  • Fragmentary axis compressions with preserved leaves
Affinities with Taxaceae in the Pinales. Originally described as Taxus jurassica.
Pagiophyllum[54]
  • Pagiophyllum kurrii
  • Pagiophyllum peregrinum
  • Pagiophyllum johnstrupi
  • Pagiophyllum falcatum
  • Pagiophyllum sp.
  • Korsodde section
  • Bagagraven clay pit
  • Hasle clay pit
Very abundant
  • Fragmentary axis compressions with preserved leaves
  • Coalified fragments
  • Cuticles
Affinities with Araucariaceae or Hirmeriellaceae in the Pinales. P. kurrii (originally P. steenstrupi) is preferred as this species is characterised by relatively broad leaves inserted at high angles to the stem. P. peregrinum has been found on the Hettangian Rønne Formation associated with hirmeriellidaceous wood of Simplicioxylon. On the Toarcian levels, is the most common plant cuticle recovered locally.
Paleopicea
  • Paleopicea glaesaria
  • Korsodde section
Very rare and limited to the middle layers
  • Pollen
Affinities with the family Pinaceae in the Pinopsida. Conifer pollen from medium to large arboreal plants.
Palissya
  • Palissya sphenolepis
  • Palissya sternbergi
  • Vellengsby
  • Nebbeodde
Rare
  • Ovuliferous cones
Affinities with Palissyaceae in the Palissyales. Descriptions of Palissya come mostly from coeval deposits in the Northern Hemisphere, based on a very few specimens from Sweden, Germany and America.
Perinopollenites
  • Perinopollenites elatoides
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Intercalations of very abudant presence in some layers with others of total absence in Levka-1; very abudant in all layers in Sorthat beds and KorsoddePollenAffinities with the family Cupressaceae in the Pinopsida.
Pinuspollenites
  • Pinuspollenites minimus
  • Pinuspollenites pinoides
  • Levka-1 borehole
  • Sorthat beds
  • Korsodde section
Abundant to very abundant with intercalations of absencePollenAffinities with the family Pinaceae in the Pinopsida.
Pityophyllum
  • Pityophyllum angustifolium
  • Bagagraven clay pit
  • Hasle clay pit
  • Vellengsby
Rare
  • Leaf compressions
  • Cuticles
Affinities with Schizolepisaceae in the Pinaceae. This genus is found associated with Schizolepis on many places, making diverse authors to put both on Pinaceae.
Pityocladus[55]
  • Pityocladus longifolius
  • Bagagraven clay pit
  • Hasle clay pit
  • Vellengsby
Rare
  • Leaf compressions
  • Cuticles
Affinities with Schizolepisaceae in the Pinaceae.
Podozamites
  • Podozamites angustifolius
  • Podozamites cuspiformis
  • Podozamites agardhianus
  • Podozamites schenkii
  • Podozamites gramineus
  • Podozamites sp.
  • Hasle clay pit
  • Bagagraven clay pit
  • Korsodde section
  • Vellengsby
  • Stina-1 well
Abundant
  • Isolated leaves
  • Cuticles
Affinities with Krassiloviaceae in the Voltziales. The local Podozamites show a great range of growth, reflecting tropical to subtropical conditions.
Schizolepidopsis[56]
  • Schizolepidopsis follini
  • Vellengsby
  • Bagagraven clay pit
Rare
  • Ovulate strobili
Affinities with Schizolepisaceae in the Pinaceae. Placed in the Pinaceae on the basis of separated scales and bract scales.
Sewardiodendron[57]
  • Sewardiodendron steenstrupii
  • Bagagraven clay pit
  • Hasle clay pit
Rare
  • Fragmentary axis compressions with preserved leaves
Affinities with Cunninghamioideae in the Cupressales. Cunninghamia-like conifers belonging to half-evergreen trees.
Simplicioxylon[58]
  • Simplicioxylon rotnaensis
  • Bagagraven clay pit
  • Stina-1 well
Abundant
  • Isolated logs
  • Isolated branches
  • Isolated Indeterminate woody material
  • Coalified fragments
Affinities with Hirmeriellaceae in the Pinales. Originally identified as Brachyoxylon rotnaensis, now thought to be a synonym of Simplicioxylon.[59] Wood from these conifers is also found in the HettangianSinemurian Rønne Formation and the Toarcian Úrkút Manganese Ore Formation.
Spheripollenites[60]
  • Spheripollenites psilatus
  • Spheripollenites subgranulatus
  • Spheripollenites subscabratus
  • Korsodde section
  • Levka-1 borehole
  • Sorthat beds
Very rare in lower layers to dominant (95% of total) in upper onesPollenAffinities with the Hirmeriellaceae in the Pinopsida. Spheripollenites psilatus composes up to 95% of the Lower Toarcian section and is correlated with Toarcian carbon cycle anomalies including the oceanic anoxic event, suggesting dry climates.
Stachyotaxus
  • Stachyotaxus septentrionalis
  • Hasle clay pit
Very rare
  • Isolated leaves
Affinities with Palissyaceae in the Palissyales.
Torreya
  • Torreya moelleri
  • Bagagraven clay pit
Rare
  • Fragmentary axis compressions with preserved leaves
Affinities with Taxaceae in the Pinales. Known only from Bornholm and belongs to an extant genus. This species is related to the Middle Jurassic floras of Yorkshire.

Ichnofossils

GenusSpeciesLocationMaterialTypeOriginNotesImages
Arenicolites
  • Arenicolites isp.
  • Levka section
  • Sorthat beds
Dwelling traces
  • Polychaetes (Spionida and Carpitellida)
  • Suspension-feeding amphipodan crustaceans
  • Deposit-feeding Sipuncula
Marine, brackish or freshwater unbranched U-shaped burrows having a subvertical orientation, with or without lining and passive fill. Are common on modern coastal environments.
Bornichnus
  • Bornichnus tortuosus
  • Korsodde section
Tubular traces Agrichnia
  • Polychaetes
Burrow-like ichnofossils. Exclusive to the Sorthat Formation, Bornichnus differs from Palaeophycus Hall in its tangled, contorted morphology and abundant branching. Small open burrows produced probably by farming worm-like animals (probably Polychaeta). Similar complicated burrow systems are produced by the polychaete Capitomastus cf. aciculatus.
Chondrites
  • Chondrites isp.
  • Korsodde section
Tubular fodinichniaFodinichnia
  • Annelids (Polychaeta
Burrow-like ichnofossils. Interpreted as the feeding burrow of a sediment-ingesting animal. A more recent study has found that Scoloplos armiger and Heteromastus filiformis, occurring in the German Wadden Sea in the lower parts of tidal flats, make burrows that are homonymous with numerous trace fossils of the ichnogenus.[62]
Cylindrichnus
  • Cylindrichnus isp.
  • Korsodde section
Burrowing and track ichnofossils Domichnia
  • Polychaetes
Burrow-like ichnofossils, found only in the uppermost part of the section; probably represents Polychaeta burrows.
Diplocraterion
  • Diplocraterion parallelum
  • Levka section
  • Baga beds
  • Sorthat beds
  • Korsodde section
U-shaped burrowsDomichnia Burrow-like ichnofossils. Most show only protrusive spreit, like the local ones, produced under predominantly erosive conditions where the organism was constantly burrowing deeper into the substrate as sediment was eroded from the top. Most Diplocraterion show only protrusive spreiten, like the local ones produced under predominantly erosive conditions where the organism was constantly burrowing deeper into the substrate as sediment was eroded from the top.
Palaeophycus
  • Palaeophycus isp.
  • Korsodde section
Cylindrical, predominantly horizontal to inclined burrows Domichnia
  • Polychaetes
  • Semiaquatic insects (Orthoptera and Hemiptera)
  • Semiaquatic and non-aquatic beetles.
Burrow-like ichnofossils. They occur in different size classes, 3, 5 and 10 mm in diameter.
Planolites[63]
  • Planolites isp.
  • Baga beds
  • Korsodde section
Cylindrical burrows Pascichnia
  • Polychaetes
Burrow-like ichnofossils referred to vermiform deposit-feeders. Sometimes considered a junior synonym of Palaeophycus.[64]
Rosselia[65]
  • Rosselia erecta
  • Korsodde section
Trace fossilSequestrichnia
  • Shrimp
  • Other aquatic arthropods
Burrow-like ichnofossils. Vertical or oblique complex trace fossil composed of a bunch of spindle-shaped structures and associated tubes, typical of a restricted environment (?estuarine/lagoonal). The local Rosselia is similar to the ichnogenus Parahentzschelinia surlyki from the lower Jurassic of Greenland, which may be a junior synonym. This trace fossil is interpreted as made by a small deposit-feeding animal, living in a tube communicating with the sea floor. These traces are linked with shrimps or other aquatic arthropods, since the tunnels possess scratch patterns.
Skolithos
  • Skolithos isp. A
  • Skolithos isp. B
  • Korsodde section
Cylindrical to subcylindrical burrows Domichnia Burrow-like ichnofossils made by organisms advancing along the bottom surface. Very narrow, vertical or subvertical, slightly winding unlined shafts filled with mud. Interpreted as dwelling structures of vermiform animals; specifically, the domichnion of a suspension-feeding worm or phoronidan, with certain Skolithos representing entrance shafts to more complicated burrows.
Teichichnus[66]
  • Teichichnus zigzag
  • Teichichnus isp.
  • Baga beds
  • Korsodde section
Dwelling tracesFodinichnia
  • Polychaetes
  • Dwelling Echiurans
  • Dwelling Holothurians.
Burrow-like ichnofossils. The level where this ichnogenus is more abundant is also composed of abundant fragments of spreite lamination, derived from the intersection with the ichnofossil. They are believed to be fodinichnia, with the organism adopting the habit of retracing the same route through varying heights of the sediment, which would allow it to avoid going over the same area. Believed to derive from annelid worms.
Thalassinoides
  • Thalassinoides isp.
  • Korsodde section
Tubular fodinichniaFodinichnia Burrow-like ichnofossils. Large burrow-systems consisting of smooth-walled, essentially cylindrical components. Found in association with Teichichnus.

See also

Notes and References

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