Zagaje Formation


The Zagaje Formation is a Latest Triassic-Early Jurassic Epoch geologic formation located mostly in Poland with layers also exposed in north Germany. This unit is known for its diverse Ichnofossil assemblages, with traces of invertebrates along vertebrate footprints, as well plants, large coal accumulations, invertebrate remains and ichnofossils. The Zagaje Formation correlates with The lower part of the Höganäs Formation in Scania, as well the Munkerup Member and the Gassum Formation in Denmark.

Paleoenvironment

The Zagaje Formation is particularly visible in the Sołtyków region and is made mostly of Early Jurassic continental mudstone-sandstone deposits linked to the onset of "depositional sequence I". Its age is confirmed as mostly Early Hettangian through stratigraphic and paleontological analyses, including fossil flora and conchostraca findings. Sedimentological studies divide the Sołtyków profile into three parts: ephemeral reservoir deposits, floodplain and lacustrine sediments, and river channel deposits, highlighting dynamic depositional environments influenced by tectonic subsidence and varying hydrological conditions.
Climate wise, the area was located back in the Hettangian around 45°N paleolatitude in Laurasia within a rise of 5–10°C above present, where it experienced significant climatic and environmental change related with sea-level fluctuations, manifested locally with a notorious retrogradational fluvial-lacustrine sedimentation, with evidence of a humid climate interspersed with drier seasons. Some plant fossils like Hirmeriella mark points of aridity on what was mostly a humid swampy alluvial-lacustrine habitat.
The Zagaje Formation's deposits are known from both outcrops and borehole profiles that consist primarily of sandstones, mudstones, and interspersed coal and siderite layers. It represents a stratigraphic gap with the underlying Upper Triassic formations and is capped by a transgressive contact with the Skłoby Formation. This unit contains freshwater fauna and diverse trace fossils, including vertebrate tracks. The paleoenvironment reflects a dynamic alluvial plain shaped predominantly by high-sinuosity stream processes, transitioning from earlier braided and low-sinuosity stream systems. This evolution is attributed to climatic changes, rising base levels, and decreasing geomorphological gradients. Observations, both from exposures and borehole data, highlight the dominance of avulsion processes, with several depositional subsystems identified. Facies with organic remains are diverse: riverbed biofacies, derived from meandering channels characterized by fining-upward sequences composed of channel lag deposits, point-bar sands, and finer overlying sediments. These deposits exhibit lateral accretion bedding and significant fossil bivalves and large-sized floated plant remains consistent with subaqueous dune migration within the channels; levee deposits derived from successive floods with scarce root traces, while plant remnants are common; Paleosoils with sparse traces of plant roots and remains of floating plants of highly variable size, fragments of bivalve shells and vertebrate remains ; the biofacies of the ephemeral water reservoir with plant remains, mainly horsetails, and fossils of insects, ostracods, and conchostraca; The pedogenic soil biofacies with remains of plant roots with preserved organic matter and rhizomes and stems in a living position; Floodplain biofacies with traces of numerous plant roots and plant macroremains, and remains of sedge stems preserved in a living position; Lake-marsh biofacies, dark, laminated mudstones with plant roots and coal, with few fossil bivalves, a large amount of organic matter in the form of plant detritus, and layers of coal and numerous finds of miospores and megaspores. The local presence of charcoal fragments and high concentrations of PAHs, along with possible burnt plants, provides evidence for wildfires in the region, that likely occurred near the surface with charred wood fragments were subsequently incorporated into sediments by river transport.
The high presence of coprolites has allow also to stablish the tropic chain of the local biota, with a clear full ecosystemical substitution of the older Triassic archosaurs by Dinosaurs.

Biota

Indet. Invertebrates

Several unname Ichnofossils are recovered at Soltyków, including conical domichnia, bivalve straight to winding linear trails, smooth vertical and subvertical branching tunnels, knob-walled tunnels, mace-shaped or irregular ellipsoid chambers, etc.
GenusSpeciesLocationMaterialMade By

Cruziana
C. problematicacf.C. isp
  • Soltyków
Dwelling structures
CochlichnusC. isp.
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
Traces
  • Annelids
  • Insect larvae
  • Nematodes
ConichnusC. isp.
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
Subcylindrical burrows
  • Annelids
  • Nematodes
  • Crustaceans
DiplichnitesD. isp.
  • Soltyków
Hypichnial trackway
  • Insects
  • Myriapods
  • Arachnids
DiplocraterionD. parallelum
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
U-Shaped Burrows
  • Crustaceans
  • Annelids
  • Poronidans
  • Insects
  • Fish
ImbrichnusI. isp.
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
Burrows
  • Annelids
  • Bivalves
  • Crustaceans
  • Kouphichniumcf. K. isp.
    • Soltyków
    V-shaped hypichnial marks
    • Xiphosura
    • Malacostraca
    PalaeophycusP. isp.
    • Soltyków
    Straight or slightly curved burrows
    • Annelids
    • Bivalves
    • Crustaceans
    PlanolitesP. isp.
    • Soltyków
    Horizontal burrows
    • Annelids
    • Bivalves
    • Crustaceans
    RusophycusR. isp.
    • Soltyków
    Resting Traces
    • Resting traces of arthropods
    ScoliciaS. isp.
    • Gromadzice
    • Kontrewers
    • Odrowaz
    • Soltyków
    Moving Traces
    • Locomotion trace of gastropods
    ScoyeniaS. isp.
    • Soltyków
    Linear slender burrows
    • Beetles?
    SkolithosS. isp.
    • Gromadzice
    • Kontrewers
    • Odrowaz
    • Soltyków
    Sac/Bottle shaped burrows
    • Annelids
    • Crustaceans
    • Insects
    SpongeliomorphaS. carlsbergiS. isp.
    • Gromadzice
    • Kontrewers
    • Odrowaz
    • Soltyków
    Sac/Bottle shaped burrows
    • Annelids
    • Crustaceans
    • Insects

    Molluscs

    Indeterminate gastropod egg capsules are known, similar to the ones recovered in the extant Neritina. 4 unnamed morphotypes of freshwater bivalves of the family Unionidae are known.
    GenusSpeciesLocationMaterialNotesImages
    AnodontaA. liasokeuperina
    • Soltyków
    Isolated ShellsA freshwater mussel, member of the family Unionidae
    CalceoformitesC. uchmani
  • Soltyków
  • Clog-shaped protrusionsBivalve estabilization traces
    CardiniaC. follini C. inglensisC. cf.kullensis
  • Gromadzice
  • Odrowaz
  • Podole
  • Soltyków
  • Isolated ShellsA Carditidae Bivalve. Indicator of oligohaline settings and found also on the younger Skłoby Formation
    LockeiaL. siliquariaL. amygdaloidesL. czarnockii
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
  • Dwelling tracesResting traces of Bivalves
    PtychoplasmaP. conica
  • Soltyków
  • Locomotion traceGastropod Locomotion traces
    ScalichnusS. phialeS. isp.
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
  • Sac/Bottle shaped burrowsEscape structure of mud-dwelling bivalves
    ScoliciaS. isp.
  • Gromadzice
  • Kontrewers
  • Odrowaz
  • Soltyków
  • TracesLocomotion and feeding trace of gastropods
    UnioU. minutus
  • Soltyków
  • Isolated ShellsA freshwater mussel, member of the family Unionidae
    ViviparusV. spp.
  • Soltyków
  • Isolated ShellsA freshwater snail, member of the family Viviparidae

    Insects

    Radial chambers around large tunnels have been recovered, they may be arthropod burrows or traces of roots. Large nest structures with septa, similar to nesting behaviour of insects like Cicadas are known.
    GenusSpeciesLocationMaterialNotesImages
    ArtematopoditesA. ssp.
    • Odrowaz
    MPK 5/36, 39, 40A Coleopteran, member of the family Permosynidae
    BlattodeaIndeterminate
  • Odrowaz
  • MPK 5/54Indeterminate Blattodean remains
    BlattulidaeIndeterminate
  • Odrowaz
  • MPK 5/1Indeterminate Cockroach remains
    CaraboideaIndeterminate
  • Odrowaz
  • MPK 5/12, 15Indeterminate Beetle remains
    ColeopteraIndeterminate
  • Odrowaz
  • Isolated WingsIndeterminate Beetle remains
    Helminthoidichnitescf. H. isp.
  • Sołtyków
  • Gnawing tracessurficial gnawing traces made by insects
    HydrobiitesH. sp.
  • Odrowaz
  • MPK 5/10, 13, 17, 22, 25, 33A Coleopteran, member of the family Permosynidae
    LinckichnusL. terebrans
  • Sołtyków
  • Boring TracesDetritivorous habitation dwellings or oviposition structures of insects in dead wood
    MemptusM. sp.
  • Odrowaz
  • MPK 5/44A Coleopteran, Incertade sedis
    NotocupesN. sp.
  • Odrowaz
  • MPK 5/6A Coleopteran, Incertade sedis
    OdrowazicorisO. polonicus
  • Odrowaz
  • MPK 5/2A Hemipteran, member of the family Belostomatidae
    PolysitumP.? sp.
  • Odrowaz
  • MPK 5/14, 29A Coleopteran, Incertade sedis
    PhoroschizidaeIndeterminate
  • Odrowaz
  • MPK 5/4,5, 8, 20, 35Indeterminate Beetle remains
    XylonichnusCf.X. isp.
  • Sołtyków
  • Boring TracesBorings in the wood made probably by insect larvae

    Fish

    Unidentified Actinopterygian fish scales and teeth were collected from clayish, organic-rich lake deposits, while some coprolites have been referred to Hybodontiform sharks.
    GenusSpeciesLocationMaterialNotesImages
    SemionotusS. cf. bergeriCzarniecka GóraSingle specimenA Semionotiform bony fish of the family Semionotidae
    PaleoniscidaeIndeterminateSołtykówScales & TeethIndeterminate Palaeonisciformes specimens

    Theropods

    Some elliptical "post-egg" structures egshells & eggs with embryo remains have been referred to theropods, yet may also belong to Ornithischians.'
    GenusSpeciesLocationMaterialNotesImages
    AnchisauripusA. ispp. Cf.A. isp.SołtykówFootprintsAdscribed to smal slender primitive predatory dinosaurs, related with genera such as Coelophysis
    EubrontesE. isp.Cf.E. isp.SołtykówFootprintsEubrontes is related to the Genus Dilophosaurus, representing a basal Neotheropods
    GrallatorG. ispp.Cf.G. isp.SołtykówFootprintsSimilar pes with Coelophysidae-alike dinosaurs, related with neotheropods such as Dracoraptor.
    KayentapusK. soltykovensisK. ispp.Cf.K. isp.SołtykówFootprintsAssumed to come from Genera similar to Sarcosaurus
    MegalosauripusCf.M. isp.SołtykówFootprintsLarge bodied taxa, maybe related with Sinosaurus. Among the largest early Jurassic theropod tracks worldwide.
    Plesiorniscf. P. isp.SołtykówFootprintsTheropod Tracks from small sized taxa with convergent features with latter Avians
    StenonyxCf.S. isp.SołtykówFootprintsSmall Theropod tracks, likely from juveniles of larger taxa
    Theropoda'Indeterminate
    • Hucisko
    • Sołtyków
  • Teeth
  • Isolated Bones
  • Bromalites
  • Some coprolites, referred to Theropods include plant material, probably ingested accidentally by drinking water. Others include large bone remains or fish scales.' Teeth corroborate the presence of large taxa in the area.'

    Plants

    In Palynology, the Zagaje Formation belongs to the Nathorstisporites ''hopliticus assemblage, indicating a spike in marshland and lacustrine settings. The Sołtyków outcrop is dominated by Classopollis, Aratrisporites, Concavisporites and Cyathidites.
    GenusSpeciesStratigraphic positionMaterialNotesImages
    AciphyllumA. triangulatum
    • Sołtyków
    CuticlesAffinities with Pinaceae inside Pinales. The oldest record of a Pinus-like needle in the fossil record
    BrachyphyllumB. sp.
  • Sołtyków
  • CuticlesAffinities with Cheirolepidiaceae or Araucariaceae inside Pinales
    Caytonia
  • C. sp.
  • Odrowąż
  • Sołtyków
  • Reproductive structureAffinities with Caytoniaceae in the Caytoniales
    CzekanowskiaC. sp.
  • Hucisko
  • Branched ShootsAffinities with the Czekanowskiales inside Ginkgoopsida. This Genus is related with relatively drier-cooler conditions.
    DesmiophyllumD. harrisii
  • Sołtyków
  • CuticlesA possible Conifer leaf, recent finds of it associated with the cone genera Sphaerostrobus and Ourostrobus points to a coniferophyte affinity, maybe as a member of Palissyaceae.
    DictyophyllumD. sp.
  • Odrowąż
  • Sołtyków
  • PinnaeAffinities with Dipteridaceae inside Gleicheniales.
    Goepertella
  • G. microloba
  • Odrowąż
  • Sołtyków
  • PinnaeAffinities with Dipteridaceae inside Gleicheniales
    HirmeriellaH. muensteri
  • Hucisko
  • Odrowąż
  • Branched Shoots and reproductive conesAffinities with the Cheirolepidiaceae inside Pinales.
    KomlopterisK. distinctiva
  • Odrowąż
  • Sołtyków
  • CuticlesAffinities Corystospermaceae inside Corystospermales.
    MatoniaM. braunii
  • Niekłań PGI-1
  • PinnaeAffinities with Matoniaceae inside Gleicheniales
    NeocalamitesN. lehmannianus
  • Sołtyków
  • StemsAffinities with Calamitaceae inside Equisetopsida. A common horsetail on the Liassic of Europe.
    NilssoniaN. sp.
  • Sołtyków
  • CuticlesAffinities with Cycadeoidaceae in the Bennettitales or alternatively a member of Nilssoniales
    OdrolepisO. liassica
  • Odrowąż
  • Sołtyków
  • Complete PlantsAffinities with Lycopodiales
    OtozamitesO. brevifolius
  • Odrowąż
  • Sołtyków
  • LeafletsAffinities with Williamsoniaceae in the Bennettitales.
    Pachypteris'P. lanceolataP. papillosa
  • Odrowąż
  • Sołtyków
  • PinnaeAffinities Corystospermaceae inside Corystospermales.
    ParacycasP. minuta
  • Odrowąż
  • Sołtyków
  • LeafletsAffinities with Cycadales in the Cycadopsida.
    PiroconitesP. kuespertii
  • Odrowąż
  • Sołtyków
  • Reproductive structureAffinities with Gnetales, maybe with Welwitschiaceae
    PhlebopterisP. angustiloba
  • Odrowąż
  • Sołtyków
  • CuticlesAffinities with Matoniaceae in the Gleicheniales.
    Podozamites'P. cf. schenkii P. sp. Cf. P. sp.
  • Sołtyków
  • Branched shootsAffinities with Krassiloviaceae inside Voltziales
    PseudotorelliaCf.P. sp.
  • Sołtyków
  • CuticlesAffinities with the Pseudotorelliaceae inside Ginkgoopsida.
    PterophyllumP. sp.Cf.P. sp.
  • Odrowąż
  • Sołtyków
  • LeafletsAffinities with Williamsoniaceae in the Bennettitales.
    Ptilozamites
  • P. cycadea
  • Sołtyków
  • CuticlesAffinities Corystospermaceae inside Corystospermales.
    SagenopterisS. nilssoniana
  • Odrowąż
  • Sołtyków
  • LeavesAffinities with Caytoniaceae in the Caytoniales
    SchmeissneriaS. microstachys
  • Odrowąż
  • Sołtyków
  • Reproductive structureAffinities with Ginkgoopsida or with Angiosperm-convergent Gimnosperms
    SwedenborgiaS. sp.
  • Sołtyków
  • Branched ShootsAffinities with Krassiloviaceae inside Voltziales.
    ThaumatopterisT. brauniana
  • Hucisko
  • Odrowąż
  • PinnaeAffinities with Dipteridaceae inside Gleicheniales
    ToditesT. princeps''
  • Hucisko
  • Odrowąż
  • PinnaeAffinities with Osmundaceae in the Osmundales.