The Sorthat Formation is a geologic formation on Bornholm, Denmark, and the Rønne Graben, Baltic Sea, from the Latest Pliensbachian to Late Toarcian. It holds plant fossils and invertebrate traces, overlain by fluvial and lacustrine deposits of the Aalenian-Bathonian Bagå Formation. Initially classified as part of the Bagå Formation until 2003, it spans the Latest Pliensbachian to Early Aalenian. It reflects a deltaic to marine setting with eastern river systems forming in the Toarcian. Early Pliensbachian volcanism from southern Sweden extended across the North Sea. The Central Skåne Volcanic Province and Egersund Basin contributed volcanic material, affecting tectonics. Early Jurassic porphyritic nephelinite lavas in the Egersund Basin, akin to those in the formation's clay pits, suggest fluvial sediment transport to the Grimmen Formation and Ciechocinek Formation. The Grimmen Formation is its sister unit.

Description

Sorthat Formation layers, mainly mudstones, claystones, and sandstone bands

Bornholm's Lower-Middle Jurassic includes the Rønne (HettangianSinemurian), Hasle (Early–Late Pliensbachian), Sorthat, and Bagå Formations. Coal-bearing clays and sands overlie the Hasle Formation, divided between the Sorthat and Bagå. The Sorthat Formation aligns with the Röddinge Formation, sharing a fluvial system, and correlates with the Ciechocinek Formation, Fjerritslev Formation, and Rya Formation. Originally termed Levka, Sorthat, and Bagå beds, its key exposure is the Korsodde section. Faulting and scarce marine fossils hinder stratigraphic clarity. Palynological data from the Levka-1 core and Korsodde section date it to Late Pliensbachian–Toarcian, possibly Early Aalenian. Megaspores indicate Toarcian–Aalenian strata. It features bioturbated sands, heteroliths, clays, coal veins, and dinoflagellates, suggesting brackish to marine settings, capped by Bagå deposits.

The Sorthat Formation's lithology varies. The Levka-1 well reveals fining-upward units, 3–14 m thick, with coarse sand, muddy, coal- and mica-rich sands, clays, and coal seams. Parallel lamination dominates, with minor cross-bedding. Abundant plant fragments and quartz, but no marine palynomorphs, indicate a coastal delta plain. This mirrors the Ciechocinek Formation's Toarcian–Bajocian deltaic shoreline. The North German Basin shows four sea-level fluctuations forming delta generations, with Toarcian regressive deltas depositing 40 m in Prignitz and Brandenburg. Palynomorphs tie to the Sorthat Formation.

The upper formation (~40 m) has bioturbated sands, heteroliths, syneresis cracks, pyrite nodules, and ichnofossils like Planolites and Teichichnus, reflecting nearshore lagoons and channels. The 93 m Korsodde section, with organic-rich sands, suggests fluvial channels tied to coastal lakes, with plant remains and ichnofossils like Diplocraterion. The top features fine, yellowish-brown sands and sandstones with bioturbated, wave-rippled beds.

At Korsodde, the environment includes the following:

Stratigraphy of the Korsodde section
UnitLithologyThickness (metres)Type of environmentFossil floraFossil fauna
Unit AYellow, weakly cemented muscovite quartz sandstone, medium- to fine-grained in the lower part, fine-grained in the upper part.0.45–2.3 mEstuarine channel fill (upper or marginal, less energetic part)None recoveredPlanolites isp. Rosselia isp. Bornichnus tortuosus Diplocraterion parallelum
Unit BIntercalations of muscovite quartz sandstones and dark mudstone drapes, with abundant heteroliths.2.3–3.41 mUpper tidal flat deposits surrounding an estuaryNone recoveredPlanolites 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 recoveredD. 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 estuaryRootsPlanolites isp. Palaeophycus isp. B. tortuosus D. parallelum
Unit EMostly fine-grained sediments with abundant organic matter.4.7–6.9 mLagoonal environment above a coal bedNeocalamites sp. stems Coal Plant cuticles Roots Root structures?Thalassinoides isp. Planolites isp. D. parallelum
Unit FMostly pale, fine-grained, ripple cross muddy sandstone and normal sandstone, separated by thin, pale sandy mudstones or thin mudstone drapes.6.9–9.9 mTidal flat deposits in an estuaryLignites 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 reportedRosselia isp. Palaeophycus isp. Cylindrichnus isp. Skolithos isp. D. parallelum
Unit I, JBioturbated muddy sandstone14.2–14.4 mShort-lived bay or lagoonNeocalamites sp. logsRosselia isp. Teichichnus isp. Teichichnus zigzag Planolites isp. Thalassinoides isp. Palaeophycus isp.

Biota

The Sorthat Formation hosts a rich PliensbachianToarcian flora, one of Europe's most complete for this period, with significant Jurassic palynological deposits. The flora, including gymnosperms, ferns (e.g., Dicksonia, Coniopteris, Osmundaceae), and thin-cutinised leaves like Podozamites and Equisetales, suggests a warm, humid climate conducive to diverse vegetation.

Environment

Late Pliensbachian–Early Toarcian Fennoscandinavia, with flora based on the Sorthat Formation and fauna from the Lehmhagen Member, Grimmen Formation and Drzewica Formation.
Paleogeography of the North German basin in the Toarcian, showing the Grimmen Formation and Sorthat Formation extent.

The Sorthat Formation, deposited in the Rønne Graben and on Bornholm's coasts, reflects a deltaic to marine environment, with the Grimmen Formation as its sister unit. A modern analog is New Zealand's Northland humid coastal forests, where peat-forming woodlands thrive. The Stina-1 well shows sand, clay, and coal, indicating an emerged graben. High kaolinite and reworked Carboniferous palynomorphs suggest erosion of a Carboniferous regolith. A Late Pliensbachian regression allowed coal deposition, halted by an Early Toarcian transgression. The Levka-1 well and Korsodde section show lagoons, channels, and floodplains, with rapid subsidence in the Rønne Graben during the Toarcian. Shoreface deposits with bioturbation mark a deepening trend, correlating with the Fjerritslev Formation. Depositional settings include the Levka Beds, interpreted as fluvial channels, floodplains, and peat swamps. Marine palynomorphs (e.g., Nannoceratopsis) indicate lagoonal settings. The Sorthat Beds represent delta plain deposits with pyrite nodules and Arenicolites traces. The Korsodde Section was a fluvial channel sands with coal and rootlets, transitioning to lagoons and palaeosols, with dinoflagellates (e.g., Mendicodinium). Outside emerged Bornholm, offshore Rønne and Kolobrzeg grabens include mostly fluvial-coastal layers, fed by regional currents.

The Sorthat Formation, with the Grimmen Formation as its sister unit, records significant vegetation changes during the Early Toarcian, linked to carbon cycle perturbations and the Toarcian oceanic anoxic event, driven by large-scale volcanism. A modern analog is New Zealand's Northland humid coastal forests, supporting peat-forming woodlands.

The Sorthat Fm was a cold, humid y forested setting, whose best modern analogs can be found in New Zealand
Stratigraphic map of Bornholm

In the Late Pliensbachian, pollen assemblages, dominated by Cupressaceae (e.g., Perinopollenites) and cycads (Cycadopites), indicate a warm, humid Mediterranean climate. During the Toarcian event, spore-rich layers reflect a shift to ferns and lycophytes, suggesting increased humidity. Post-event, Hirmeriellaceae pollen (e.g., Corollina, Spheripollenites) dominates, indicating a drier, warmer climate. Woody vegetation shows changes in carbon isotopes, with pollen from Sciadopityaceae, Miroviaceae (Cerebropollenites), cycads (Chasmatosporites), and Corollina. Macroflora, mainly from the Hasle clay pit and Korsodde section, includes 68 species, 50% ferns. The Late Pliensbachian flora, rich in ferns and sphenophytes, grew in marshy floodplains along a meandering river, with Bennettites as shrubs and conifers (e.g., Pagiophyllum) and ginkgoaleans as trees, reflecting a warm, seasonal climate. In the Toarcian, Hirmeriellaceae conifers (95% of pollen), seed ferns, Bennettites, and Czekanowskiales dominate, with Pagiophyllum and Corollina torosus indicating high temperatures, aridity, and seasonal wildfires. Wood fragments, both macroscopic and microscopic (0.25–1 mm), are common in Korsodde's nearshore deposits.

Coal seams, mainly from Levka-1 and Korsodde, formed in anoxic, nutrient-rich swamps. Peat accumulated rapidly (~1 mm/yr), akin to Central Kalimantan, in a warm, humid climate. High huminite content (e.g., Eu-ulminite, densinite) indicates anoxic conditions. Wildfires, evidenced by charcoal, increased during the Toarcian oceanic anoxic event, reflecting drier conditions. Coals can be found at Levka-1 (112 m of coal, sand, and clay with abundant coalified wood), representing fluvial channels and floodplains with lagoons. At Korsodde six coal seams from a coastal lagoon setting can be found, with huminite-rich coal and dinoflagellates like Mendicodinium reticulatum.

Fungi

Color key Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon MorphotaxonNotes Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
TaxonReclassified taxonTaxon falsely reported as presentDubious taxon or junior synonymIchnotaxonOotaxonMorphotaxon
GenusSpeciesStratigraphic positionMaterialNotesImages
Fungi"Morphotype A" "Morphotype B" "Morphotype C"Korsodde sectionFungal spores Hyphae?Fungal spores of uncertain classification. The three uppermost samples of the Korssode section are poor in diversity, but fungal spores are common in at least one sample; these have not been recorded from the samples below.Extant Geastrum campestre specimen, found linked with plant matter. Spores recovered on the Sorthat Formation may be derived from similar fungi.

Phytoplankton

In the Lower Jurassic of Bornholm there were several successions of nearshore peat formations with dinoflagellates. 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.

GenusSpeciesStratigraphic positionMaterialNotesImages
BaltisphaeridiumB. infulatumKorsodde sectionCystsAn algal acritarch, probably related to freshwater red algae, similar to extant Florideophyceae (for example, Hildenbrandia) or Batrachospermales (Batrachospermum) and Thoreales.Extant Hildenbrandia; Baltisphaeridium may be derived from a similar genus
BotryococcusB. sp.Korsodde section Sorthat beds Levka-1 boreholeMiosporesType 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.Extant specimen
ChomotriletesC. minorKorsodde section Levka-1 borehole Sorthat bedsMiosporesAffinities with the family Zygnemataceae. A genus derived from freshwater filamentous or unicellular, uniseriate (unbranched) green algae.
CrassosphaeraC. coccinia C. hexagonalisKorsodde sectionMiosporesAffinities with the family Pycnococcaceae.
CymatiosphaeraC. sp.Korsodde section Sorthat bedsMiosporesAffinities with the family Pterospermopsidaceae.
HaplophragmoidesH. tryssa H. platus H. sp.Sorthat bedsCystsA foraminifer, member of the family Lituoloidea in the Lituolida.
KorystocystaK. sp.Korsodde sectionCystsA dinoflagellate, member of the Cribroperidinioideae.
LecaniellaL. foveataKorsodde section Levka-1 borehole Sorthat bedsMiosporesAffinities with the family Zygnemataceae.
LeiosphaeridaL. spp.Korsodde section Levka-1 borehole Sorthat bedsMiosporesAffinities with the family Prasinophyceae.
LuehndeaL. spinosaKorsodde sectionCystsA dinoflagellate, member of the Luehndeoideae. It establishes the Luehndea spinosa zone; the age of this zone is late Pliensbachian to early Toarcian.
MancodiniumM. semitabulatumKorsodde sectionCystsA dinoflagellate, type genus of the Mancodinioideae.
MendicodiniumM. groenlandicum M. reticulatumKorsodde section Sorthat bedsCystsA dinoflagellate, member of the family Gonyaulacales.
MicrhystridiumM. fragile M. intromittum M. lymensis M. spp.Korsodde section Levka-1 boreholeCystsAn acritarch, familia incertae sedis
NannoceratopsisN. senex N. gracilis N. ridingui N. triangulata N. triceras N. dictyanbonis N. sp.Korsodde section Levka-1 borehole Sorthat bedsCystsA 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.
OvoiditesO. sp. A O. sp. B O. sp. C O. spp.Korsodde section Levka-1 boreholeMiosporesAffinities with the family Zygnemataceae. A genus derived from freshwater filamentous or unicellular, uniseriate (unbranched) green algae.Extant Spirogyra; Ovoidites may be derived from a similar genus
PterospermellaP. spp.Korsodde sectionMiosporesAffinities with the family Pterospermataceae.
RotundusR. granulatusSorthat bedsMiosporesAn algal palynomorph unique to the setting and probably related to freshwater red algae; similar to extant Batrachospermales.Extant Batrachospermum
SpirillinaS. sp.Sorthat bedsMiosporesA foraminifer, type genus of the Spirillinidae in the Spirillinida.
StriatellaS. jurassica S. parva S. seebergensis S. scanicaKorsodde section Levka-1 borehole Sorthat bedsMiosporesBrown algae, type genus of the family Striatellaceae in the Striatellales. These brown algae diatoms are associated with either brackish or marginal marine environments.
TasmanitesT. sp.Korsodde section Levka-1 borehole Sorthat bedsMiosporesAffinities with the family Pyramimonadaceae. Found on shoreface and shoreface–offshore transition zone deposits.
TetraporinaT. compressaKorsodde section Levka-1 borehole Sorthat bedsMiosporesAffinities with the family Zygnemataceae.
VeryhachiumV. spp.Levka-1 boreholeCystsA dinoflagellate, member of the Dinophyceae.

Bryophyta

GenusSpeciesStratigraphic positionMaterialNotesImages
CingutriletesC. infrapunctus C. oculusKorsodde sectionSporesIncertae sedis; affinities with Bryophyta. This spore is found in Jurassic sediments associated with the polar regions. The Sorthat Formation is among its southernmost locations.
ForaminisporisF. jurassicusKorsodde sectionSporesAffinities with the family Notothyladaceae in the Anthocerotopsida. Hornwort spores.Extant Notothylas specimens; Foraminisporis probably come from similar genera.
PolycingulatisporitesP. circulus P. liassicus P. triangularisLevka-1 borehole Korsodde sectionSporesAffinities with the family Notothyladaceae in the Anthocerotopsida. Hornwort spores.
SculptisporisS. aulosenensisSorthat beds Korsodde sectionSporesAffinities with the family Sphagnaceae in the Sphagnopsida.
StaplinisporitesS. caminusKorsodde sectionSporesAffinities with the family Encalyptaceae in the Bryopsida. Branching moss spores, indicating high water-depleting environments.Extant Encalypta specimens; Staplinisporites probably come from similar genera
StereisporitesS. antiquasporites S. stereoidesLevka-1 borehole Sorthat beds Korsodde sectionSporesAffinities with the family Sphagnaceae in the Sphagnopsida. "Peat moss" spores, related to genera such as Sphagnum that can store large amounts of water.Extant Sphagnum specimens; Stereisporites, Sculptisporis and Rogalskaisporites probably come from similar genera
TaurocusporitesT. verrucatusKorsodde sectionSporesAffinities with the family Sphagnaceae in the Sphagnopsida.
RogalskaisporitesR. cicatricosusLevka-1 borehole Sorthat beds Korsodde sectionSporesAffinities with the family Sphagnaceae in the Sphagnopsida.

Lycophyta

GenusSpeciesStratigraphic positionMaterialNotesImages
AnapiculatisporitesA. spiniger A. sp.Levka-1 borehole Korsodde sectionSporesAffinities 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.Extant Selaginella, typical example of Selaginellaceae. Genera like Anapiculatisporites or Densoisporites probably come from a similar or a related Plant
CadargasporitesC. granulatusKorsodde sectionSporesAffinities with the Selaginellaceae in the Lycopsida.
CamarozonosporitesC. golzowensis C. rudisKorsodde sectionSporesAffinities with the family Lycopodiaceae in the Lycopodiopsida.
KraeuselisporitesK. reissingeriKorsodde sectionSporesAffinities with the Selaginellaceae in the Lycopsida.
NeoraistrickiaN. gristhorpensis N. sp.Levka-1 borehole Sorthat beds Korsode sectionSporesAffinities with the Selaginellaceae in the Lycopsida.
RetitriletesR. austroclavatidites R. clavatoides R. semimurus R. spp.Korsodde section Levka-1 borehole Sorthat bedsSporesAffinities with the family Lycopodiaceae in the Lycopodiopsida.
SelaginellitesS. falcatusHasle clay pitFine 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.
SestrosporitesS. pseudoalveolatusSorthat beds Korsodde sectionSporesAffinities with the family Lycopodiaceae in the Lycopodiopsida. Lycopod spores, related to herbaceous to arbustive flora common in humid environments.Extant Lycopodium specimens. Genera like Sestrosporites, Camarozonosporites, Retitriletes, Lycopodiumsporites and Semiretisporis probably come from a similar plant
UvaesporitesU. argenteaeformis U. microverrucatus U. puzzleiLevka-1 borehole Sorthat beds Korsodde sectionSporesAffinities with the Selaginellaceae in the Lycopsida.

Equisetales

GenusSpeciesStratigraphic positionMaterialNotesImages
CalamosporaC. tenerLevka-1 borehole Sorthat beds 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.Reconstruction of the genus Calamites, found associated with Calamospora
EquisetitesE. munsteri E. lyelli E. sp.Bagagraven clay pit Nebbeodde Stina-1 wellStemsAffinities 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.Equisetites specimens from Germany
NeocalamitesN. hoerensis N. sp.Korsodde section Bagagraven clay pitThree incomplete axes Isolated Incomplete fragmentsAffinities 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.Neocalamites specimen from Austria
PhyllothecaP. cf. equisetiformisHasle clay pitLeaf whorlsAffinities with Equisetidae in the Equisetales.

Pteridophyta

GenusSpeciesStratigraphic positionMaterialNotesImages
AnnulisporaA. folliculosaKorsodde sectionSporesAffinities 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.Extant Saccoloma specimens; Annulispora probably comes from similar genera or maybe a species in the genus
BaculatisporitesB. comaumensis B. primarius B. wellmanii B. sp.Korsodde section Levka-1 borehole Sorthat bedsSporesAffinities with the family Osmundaceae in the Polypodiopsida. Near fluvial current ferns, related to the modern Osmunda regalis.Extant Osmunda specimens; Baculatisporites and Todisporites probably come from similar genera or maybe a species from the genus
CladophlebisC. nebbensis C. roesserti C. svedbergii C. hirtaVellengsby Bagagraven clay pit Hasle clay pit NebbeoddeIsolated pinnaeAffinities with Osmundaceae in the Osmundales. Related to species commonly reported from the Triassic–Jurassic of southern Sweden.Cladophlebis nebbensis specimen from Japan
CladothecaC. undansBagagraven clay pitFertile pinna fragmentsAffinities 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.
CibotiumsporaC. jurienensisSorthat beds Korsodde sectionSporesAffinities with the family Cyatheaceae in the Cyatheales. Arboreal fern spores.
ClathropterisC. meniscioides C. platyphyllaBagagraven clay pit Hasle clay pit NebbeoddeIsolated pinnaeAffinities with Dipteridaceae in the Polypodiales.Clathropteris meniscioides specimen from an unknown location
ConbaculatisporitesC. mesozoicus C. spinosusLevka-1 borehole Sorthat beds Korsodde sectionSporesIncertae sedis; affinities with the Pteridophyta
ConiopterisC. hymenophylloides C. acutidensBagagraven clay pitIncomplete 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).Coniopteris specimen from Hungary
DeltoidosporaD. minor D. toralis D. spp.Korsodde section Levka beds Sorthat bedsSporesIncertae sedis; affinities with the Pteridophyta
DicksoniaD. pingelii D. paucilobaBagagraven clay pit Hasle clay pitLeafletsAffinities 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.Extant Dicksonia
DictyophyllumD. acutilobium D. munsteri D. barthollini D. cf. nilssonii D. cf. spectabileVellengsby Bagagraven clay pit Hasle clay pitIsolated pinnaeAffinities with Dipteridaceae in the Polypodiales. Dictyophyllum is a common dipteridacean genus of the mid-Mesozoic.Dictyophyllum nilssonii specimen from an unknown location
EboraciaE. lobifolia E. sp.Bagagraven clay pit Hasle clay pit VellengsbyIsolated pinnaeAffinities 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.
GleicheniiditesG. senonicusLevka-1 borehole Korsodde sectionSporesAffinities with the Gleicheniales in the Polypodiopsida. Fern spores from low herbaceous flora.Extant Gleichenia specimens; Gleicheniidites and Iraqispora probably come from similar genera or maybe a species in the genus
GutbieraG. angustilobaVellengsby NebbeoddeIsolated pinnaeAffinities with Matoniaceae in the Gleicheniales.
HausmanniaH. crenata H. dichotoma H. dentata H. lasciniata H. acutidensBagagraven clay pit Hasle clay pitIsolated pinnaeAffinities with Dipteridaceae in the Polypodiales. Specimens from the same species have been found in the Hettangian Höör Sandstone at southern Sweden.
IntrapunctisporisI. toralisSorthat bedsSporesIncertae sedis; affinities with the Pteridophyta
IraqisporaI. labrataKorsodde sectionSporesAffinities with the Gleicheniales in the Polypodiopsida. Fern spores from low herbaceous flora.
IschyosporitesI. crateris I. variegatusLevka-1 borehole Korsodde sectionSporesIncertae sedis; affinities with the Pteridophyta
KlukisporitesK. lacunusKorsodde sectionSporesAffinities with the family Lygodiaceae in the Polypodiopsida. Climbing fern spores.Extant Lygodium; Lygodioisporites probably comes from similar genera or maybe a species from the genus
LaevigatosporitesL. mesozoicusKorsodde section Levka-1 boreholeSporesIncertae sedis; affinities with the Pteridophyta
LeptolepiditesL. bossus L. macroverrucosus L. major L. sp.Korsodde sectionSporesAffinities with the family Dennstaedtiaceae in the Polypodiales. Forest fern spores.Extant Dennstaedtia specimens; Leptolepidites probably comes from similar genera
LycopodiaciditesL. infragranulatus L. infragranulatusLevka-1 borehole Sorthat beds Korsodde sectionSporesAffinities with the Ophioglossaceae in the Filicales. Fern spores from lower herbaceous flora.Extant Helminthostachys specimens; Lycopodiacidites probably comes from similar genera or maybe a species from the genus
ManumiaM. delcourtiiLevka-1 borehole Sorthat beds Kosodde sectionSporesAffinities with the Pteridaceae in the Polypodiopsida. Forest ferns from humid ground locations.Extant Pityrogramma specimens; Contignisporites and Manumia probably come from similar genera or maybe a species in the genus
MarattiaM. munsteriVellengsbyIsolated pinnaeAffinities with Marattiaceae in the Marattiopsida.Extant Marattia specimen
MarattisporitesM. scabratusLevka-1 borehole Sorthat beds Korsodde sectionSporesAffinities with the Marattiaceae in the Polypodiopsida. Fern spores from low herbaceous flora.Extant Marattia specimens; Marattisporites probably comes from similar genera
PhlebopterisP. schouwii P. elegans P. mirovensis P. woodwardii P. affinis P. polypodioidesVellengsby Bagagraven clay pit Hasle clay pit NebbeoddeIsolated pinnaeAffinities with Matoniaceae in the Gleicheniales.Phlebopteris specimen from an unknown location
SkarbysporitesS. crassexiniusSorthat beds Korsodde sectionSporesIncertae sedis; affinities with the Pteridophyta
SpiropterisS. sp.Bagagraven clay pitSingle impressionIncertae ordinis in the Pteridophyta. Spiropteris is the name given to the fossil of a coiled, unopened fern leaf.
TigrisporitesT. halleinis T. microrugulatusSorthat beds Korsodde sectionSporesIncertae sedis; affinities with the Pteridophyta
ThaumatopterisT. braunianaVellengsby Bagagraven clay pitIsolated pinnaeAffinities with Dipteridaceae in the Polypodiales.Thaumatopteris specimen from Germany
TodisporitesT. major T. minorLevka-1 borehole Sorthat bedsSporesAffinities with the family Osmundaceae in the Polypodiopsida.
TripartinaT. variabilisLevka-1 borehole Sorthat beds Korsodde sectionSporesAffinities with the genus Dicksoniaceae in the Polypodiopsida. Tree fern spores.Extant Lophosoria specimens; Tripartina and Undulatisporites probably come from similar genera
VerrucosisporitesV. obscurilaesuratusKorsodde sectionSporesIncertae sedis; affinities with the Pteridophyta
VesicasporaV. fuscusKorsodde sectionSporesAffinities with the Callistophytaceae in the Callistophytales. Spores from large arboreal to arbustive ferns.
ZebrasporitesZ. interscriptusKorsodde sectionSporesAffinities with the family Cyatheaceae in the Cyatheales. Arboreal fern spores.Extant Cyathea; Zebrasporites and Cibotiumspora probably come from similar genera

"Peltaspermales"/Indet. Spermatophytes

GenusSpeciesStratigraphic positionMaterialNotesImages
AlisporitesA. grandis A. radialis A. robustus A. thomasii A. microsaccus A. diaphanusLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities 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.
CarpolithesC. cinctus C. nebbensis C. nummulariusVellengsby Bagagraven clay pit NebbeoddePlant propagulesPlant propagules that may be from Pteridospermatophyta, Vladimariales, Bennettitales or Pinales. Fruits or seeds of uncertain placement.Extant Cephalotaxus fruits. Some Carpolithes are similar conifer-derived propagules.
CtenozamitesC. leckenbyiBagagraven clay pitIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta.
CycadopterisC. heterophylla C. braunianaBagagraven clay pitIsolated pinnaeAffinities with Corystospermaceae in the Pteridospermatophyta.Cycadopteris specimen from an unknown location
FeildeniaF. cuspiformisHasle clay pitLeaf compressionsAffinities with Umaltolepidaceae in the Vladimariales. These belong to a group parallel to Gingkoaceans and derived probably from Umkomasiaceae.
KekryphalosporaK. distinctaSorthat beds Levka-1 borehole Korsodde sectionPollenAffinities with the families Peltaspermaceae, Corystospermaceae or Umkomasiaceae in the Peltaspermales.
KomlopterisK. nordenskioeldiiVellengsby Korsodde sectionIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta.
PachypterisP. laceolata P. papillosaVellengsby Korsodde sectionIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta. Less common than other arboreal plants.
PtilozamitesP. falcatus P. cycadeaBagagraven clay pitIsolated pinnaeAffinities with Umkomasiaceae in the Pteridospermatophyta.Ptilozamites specimen from Italy
PteridospermaeIndeterminateKorsodde sectionCuticlesAffinities with Pteridospermae in the Pteridospermatophyta.
SagenopterisS. cuneata S. phillipsi S. rhoifolia S. nilssoniana S. undulata S. sp.Vellengsby Bagagraven clay pitIsolated pinnaeAffinities with Caytoniaceae in the Pteridospermatophyta. Related to seed ferns present in the Rhaetic flora of Sweden.Sagenopteris specimen from an unknown location
VitreisporitesV. bjuvensis V. pallidusLevka-1 borehole Korsodde sectionPollenFrom the family Caytoniaceae in the Caytoniales. Caytoniaceae are a complex group of Mesozoic fossil floras that may be related to both Peltaspermales and Ginkgoaceae.

Erdtmanithecales

GenusSpeciesStratigraphic positionMaterialNotesImages
EucommiiditesE. major E. troedssonii E. sp.Levka-1 borehole Sorthat beds Korsodde sectionPollenType pollen of the Erdtmanithecales, related to the Gnetales. Thick tectum, infratectum of small granules, indistinct or absent foot layer. Originally thought to come from angiosperms, then suggested to come from arbustive Bennettites. It was recently found to come from Eucommiitheca, a member of the enigmatic Erdtmanithecales, reinterpreted as an unusual gymnosperm grain with a single distal colpus flanked by two subsidiary lateral colps. Is very similar to the pollen of the extant Ephedra and Welwitschia (mainly on the basis of the granular structure of the exine).

Cycadophyta

GenusSpeciesStratigraphic positionMaterialNotesImages
ButefiaB. ensiformisBagagraven clay pitLeafletsAffinities with Cycadales in the Cycadopsida. Originally described as Podozamites ensiformis.
ChasmatosporitesC. apertus C. elegans C. hians C. major C. minorLevka-1 borehole Korsodde sectionPollenAffinities 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.Extant Encephalartos laevifolius. Chasmatosporites may come from a related plant
ClavatipollenitesC. hughesiiLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities 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.Extant Cycas platyphylla. Clavatipollenites may come from a related plant
CtenisC. nathorstiHasle clay pitLeafletsAffinities with Cycadales in the Cycadopsida.Ctenis specimen from Sweden

Bennettitales

GenusSpeciesStratigraphic positionMaterialNotesImages
CycadopitesC. nitidus C. andrewsiiKorsodde sectionPollenAffinities with the family Cycadaceae and Bennettitaceae. It has been found associated with the Bennetite pollen cone Bennettistemon. It increases towards the Toarcian section.
DictyozamitesD. johnsirupiBagagraven clay pitLeafletsAffinities with Williamsoniaceae in the Bennettitales.
NilssoniaN. polymorpha N. münsteri N. acuminataVellengsby Bagagraven clay pit NebbeoddeLeafletsAffinities with Cycadeoidaceae in the Bennettitales. The most common and abundant bennetite on the formation.Nilssonia specimen from an unknown location
NilssoniopterisN. tenuinervis N. glandulosaBagagraven clay pit Hasle clay pitLeafletsAffinities with Cycadeoidaceae in the Bennettitales.
OtozamitesO. bornholmiensis O. latior O. bartholini O. tenuissimus O. bunburyanus O. obtusus O. pusillus O. beani O. pterophylloides O. molinianus O. cf. reglei O. cf. mimetesVellengsby Bagagraven clay pit Stina-1 wellLeafletsAffinities with Williamsoniaceae in the Bennettitales. Insufficient and incomplete material prevents certain assignment of Otozamites cf. reglei and Otozamites cf. mimetesOtozamites specimen from an unknown location
PterophyllumP. tenuicaule P. carnallianum P. cf. aequale P. cf. braunianumVellengsby Bagagraven clay pitLeafletsAffinities with Williamsoniaceae in the Bennettitales.Pterophyllum specimen from Austria
WilliamsoniaW. forchhammeriNebbeoddeBennettitalean "flower"Affinities with Williamsoniaceae in the Bennettitales.Williamsonia "flower" fossil from England

Ginkgoales

GenusSpeciesStratigraphic positionMaterialNotesImages
BaieraB. czekanowskiana B. pulchella B. sp.Korsodde section Bagagraven clay pit VellengsbyLeaf compressions; CuticlesAffinities with Karkeniaceae in the Ginkgoales. Unlike other plant specimens from the location, it is more characteristic of Middle Jurassic flora.Baiera specimen from an unknown location
CzekanowskiaC. hartzii C. cf. setaceaKorsodde section Hasle clay pit VellengsbyLeaf compressions; CuticlesAffinities with Czekanowskiales in the Ginkgoales. This genus is related to flora from the Rhaetian–Hettangian boundary of Jameson Land, but also present in Romania.
HartziaH. tenuis H. sp.Korsodde sectionLeaf compressions; CuticlesAffinities with Czekanowskiales in the Ginkgoales. Linked to the Lower Liassic flora of Greenland.
GinkgoitesG. troedssonii G. sibirica G. obovataKorsodde section Bagagraven clay pit Hasle clay pit NebbeoddeLeaf compressions; CuticlesAffinities 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.Ginkgoites sibirica reconstruction
MonosulcitesM. minimus M. punctatusLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities with the family Karkeniaceae and Ginkgoaceae in the Ginkgoales. Had been considered pollen of Chloranthaceae but is likely from Ginkgoales, which can have similar featuresExtant Ginkgo, the only surviving member of the Ginkgoaceae. Monosulcites pollen is similar to the pollen of this extant species.
SolenitesS. murrayanaKorsodde sectionLeaf compressions; CuticlesAffinities with Czekanowskiales in the Ginkgoales. This species was described on the basis of individuals collected in Greenland from the Triassic–Jurassic boundary.

Coniferophyta

GenusSpeciesStratigraphic positionMaterialNotesImages
AgathoxylonA. württembergicaBagagraven clay pitFossil WoodAffinities 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.Agathoxylon
AraucariacitesA. australisLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities with Araucariaceae in the Pinales.
BartholinodendronB. punctulatumBagagraven clay pit Hasle clay pitFragmentary axis compressions with preserved leavesAffinities 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.
BrachyphyllumB. mamillare B. sp.Vellengsby Korsodde sectionFragmentary axis compressions with preserved leaves; Coalified fragments; CuticlesAffinities with Araucariaceae or Hirmeriellaceae in the Pinales. Is related to the Hettangian axis found in Scania, SwedenBrachyphyllum specimen from an unknown location
CallialasporitesC. dampieri C. turbatus C. microvelatus C. segmentatusSorthat beds Korsodde sectionPollenAffinities with the family Araucariaceae in the Pinales. Conifer pollen from medium to large arboreal plants.Extant Araucaria. Callialasporites may come from a related plant
CerebropollenitesC. macroverrucosus C. thiergartiiLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities 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.Extant Sciadopitys. Cerebropollenites likely come from a related plant
CorollinaC. torosus C. meyerianaLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities with the Hirmeriellaceae in the Pinopsida.
CyparissidiumC. blackiiKorsodde sectionCoalified fragments; CuticlesAffinities with Cupressoideae in the Cupressales. It matches with the Middle Jurassic Cyparissidium blackii from Yorkshire, England.
DactylethrophyllumD. ramonensisKorsodde sectionCoalified fragments; CuticlesAffinities 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.
ElatocladusE. subzamioidesBagagraven clay pitFragmentary axis compressions with preserved leavesAffinities with Thujaceae in the Cupressales. It was originally described as Taxites? subzamioides, later merged with Elatocladus.Elatocladus specimen from Italy
ExesipollenitesE. tumulusKorsodde sectionPollenAffinities 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.Extant Austrocedrus. Exesipollenites and Perinopollenites maybe come from a related plant
HirmeriellaH. münsteriBagagraven clay pitOvuliferous ConesAffinities with Hirmeriellaceae in the Pinales. The main genus of the Hirmeriellaceae, found in dry environments and probably fire tolerant.
QuadraeculinaQ. anellaeformisLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities originally suggested with the family Podocarpaceae in the Pinopsida. Quadraeculina is not comparable to pollen of any modern gymnosperm family.
LindleycladusL. lanceolatusHasle clay pit Bagagraven clay pitLeaf compressions; CuticlesAffinities with Krassiloviaceae in the Voltziales.
MarskeaM. jurassicaBagagraven clay pitFragmentary axis compressions with preserved leavesAffinities with Taxaceae in the Pinales. Originally described as Taxus jurassica.
PagiophyllumP. kurrii P. peregrinum P. johnstrupi P. falcatum P. sp.Korsodde section Bagagraven clay pit Hasle clay pitFragmentary axis compressions with preserved leaves; Coalified fragments; CuticlesAffinities 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.Pagiophyllum specimen from an unknown location
PaleopiceaP. glaesariaKorsodde sectionPollenAffinities with the family Pinaceae in the Pinopsida. Conifer pollen from medium to large arboreal plants.Extant Picea. Paleopicea and Pinuspollenites may come from a related plant
PalissyaP. sphenolepis P. sternbergiVellengsby NebbeoddeOvuliferous ConesAffinities 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.
PerinopollenitesP. elatoidesLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities with the family Cupressaceae in the Pinopsida.
PinuspollenitesP. minimus P. pinoidesLevka-1 borehole Sorthat beds Korsodde sectionPollenAffinities with the family Pinaceae in the Pinopsida.
PityophyllumP. angustifoliumBagagraven clay pit Hasle clay pit VellengsbyLeaf compressions; CuticlesAffinities with Schizolepisaceae in the Pinaceae. This genus is found associated with Schizolepis on many places, making diverse authors to put both on Pinaceae.
PityocladusP. longifoliusBagagraven clay pit Hasle clay pit VellengsbyLeaf compressions; CuticlesAffinities with Schizolepisaceae in the Pinaceae.
PodozamitesP. angustifolius P. cuspiformis P. agardhianus P. schenkii P. gramineus P. sp.Hasle clay pit Bagagraven clay pit Korsodde section Vellengsby Stina-1 wellFragmentary axis compressions with preserved leaves; Coalified fragments; CuticlesAffinities with Krassiloviaceae in the Voltziales. The local Podozamites show a great range of growth, reflecting tropical to subtropical conditions.Podozamites reconstruction
SchizolepidopsisS. folliniVellengsby Bagagraven clay pitOvulate strobiliAffinities with Schizolepisaceae in the Pinaceae. Placed in the Pinaceae on the basis of separated scales and bract scales.
SewardiodendronS. steenstrupiiBagagraven clay pit Hasle clay pitFragmentary axis compressions with preserved leavesAffinities with Cunninghamioideae in the Cupressales. Cunninghamia-like conifers belonging to half-evergreen trees.
SimplicioxylonS. rotnaensisBagagraven clay pit Stina-1 wellFossil WoodAffinities with Hirmeriellaceae in the Pinales. Originally identified as Brachyoxylon rotnaensis, now thought to be a synonym of Simplicioxylon. Wood from these conifers is also found in the HettangianSinemurian Rønne Formation and the Toarcian Úrkút Manganese Ore Formation.
SpheripollenitesS. psilatus S. subgranulatus S. subscabratusKorsodde section Levka-1 borehole Sorthat bedsPollenAffinities 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.
StachyotaxusS. septentrionalisHasle clay pitFragmentary axis compressions with preserved leavesAffinities with Palissyaceae in the Palissyales.Stachyotaxus specimens from Germany
TorreyaT. moelleriBagagraven clay pitFragmentary axis compressions with preserved leavesAffinities 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.Extant Torreya specimen

Amber

TypeLocationMaterialNotes
AmberSorthat bedsAmber fragmentsB. Eske Koch corroborated the presence of amber drops in the Sorthat Formation. This record represents one of the few worldwide from Jurassic layers. This amber was quoted as derived from Coniferales indet.

Ichnofossils

GenusSpeciesLocationMaterialOriginImages
ArenicolitesA. isp.Levka section Sorthat bedsDwelling tracesPolychaetes (Spionida and Carpitellida) Suspension-feeding amphipodan crustaceans Deposit-feeding SipunculaArenicolites specimen from Mongolia
BornichnusB. tortuosusKorsodde sectionTubular tracesPolychaetes
ChondritesC. isp.Korsodde sectionTubular fodinichniaAnnelids (Polychaeta SipunculaIllustration of Chondrites bollensis
CylindrichnusC. isp.Korsodde sectionBurrowing and track ichnofossilsPolychaetes
DiplocraterionD. parallelumLevka section Baga beds Sorthat beds Korsodde sectionU-shaped burrowsPolychaeta annelids (Axiothella, Abarenicola and Scolecolepis) Sipunculans (Sipunculus) Enteropneustans (Balanoglossus) Echiurans (Urechis).Diplocraterion parallelum diagram
PalaeophycusP. isp.Korsodde sectionCylindrical, predominantly horizontal to inclined burrowsPolychaetes Semiaquatic insects (Orthoptera and Hemiptera) Semiaquatic and non-aquatic beetles.Palaeophycus fossil from the United States
PlanolitesP. isp.Baga beds Korsodde sectionCylindrical burrowsPolychaetesPlanolites fossil from Junget Beach, Salling, Denmark
RosseliaR. erectaKorsodde sectionTrace fossilShrimp Other aquatic arthropods
SkolithosS. isp. A S. isp. BKorsodde sectionCylindrical to subcylindrical burrowsPolychaetes PhoronidansSkolithos ichnofosil reconstruction, with possible fauna associated
TeichichnusT. zigzag T. isp.Baga beds Korsodde sectionDwelling tracesPolychaetes Dwelling Echiurans Dwelling Holothurians.Teichichnus fossil from the United States
ThalassinoidesT. isp.Korsodde sectionTubular fodinichniaThalassinidea Several crustaceans (Anomura, Decapoda) Annelids (Polychaeta) Sipuncula DipnoiThalassinoides burrowing structures, with modern related fauna, showing the ecological convergence and the variety of animals that left this Ichnogenus.

See also