Rya Formation
Stratigraphic range: Early Sinemurian-late Aalenian
~
TypeFormation
Sub-unitsDöshult, Pankarp, Katslösa & Rydebäck Members
UnderliesVilhelmsfält & Mariedal Formations
OverliesHöganäs Formation
ThicknessUp to 295 m (968 ft)
Lithology
PrimarySiltstone, claystone, sandstone, mudstone
OtherCoal
Location
Coordinates56°00′N 12°48′E / 56.0°N 12.8°E / 56.0; 12.8
Approximate paleocoordinates45°24′N 16°42′E / 45.4°N 16.7°E / 45.4; 16.7
RegionSkåne County
Øresund
Holstein
Country Sweden
 Denmark (offshore, subsurface)
 Germany (ex situ)
ExtentHöganäs & Øresund Basins
Type section
Named forRya, Katslösa
Rya Formation is located in Sweden
Rya Formation
Rya Formation (Sweden)

The Rya Formation (Swedish: Ryaformationen) is a geologic formation in Skåne County, southern Sweden. It is Early to early Middle Jurassic (early Sinemurian to late Aalenian) in age. The Rya Formation comprises siltstones, claystones, sandstones, mudstones and rare coal beds. The formation overlies the Höganäs Formation and is overlain by the Vilhelmsfält and Mariedal Formations.

The formation was deposited in the Höganäs and Øresund Basins that formed in the earliest Jurassic as part of the break-up of Pangea. The 295 metres (968 ft) thick formation comprises four members, from base to top the Döshult, Pankarp, Katslösa and Rydebäck Members. The depositional environment of the formation ranges from continental to open marine.

The Rya Formation has provided fossils of a number of sharks, ammonites, bivalves and ichnofossils. Coalified wood occurs as scattered pieces up to 6 centimetres (2.4 in) long and indeterminate belemnites, echinoids, serpulids, ostracods and nodosariid foraminifera were also recorded in the formation. Iron ooids containing erratic boulders, called Geschiebe in German, attributed to the Rya Formation were found in Holstein, northern Germany.

Description

The Rya Formation crops out in Colonus Shale Trough of western Scania.[1] The formation overlies the Höganäs Formation and is overlain by the Vilhelmsfält Formation in the Helsingborg area and by the Mariedal Formation in the area of Landskrona and Kävlinge.[2] The Rya Formation is subdivided, from base to top, into the Döshult, Pankarp, Katslösa and Rydebäck Members. The formation is found in the Ängelholm, Helsingborg, Landskrona and Kävlinge areas. In southwest Skåne, the Rya Formation is missing or only poorly developed.[3] In the Øresund Basin between Sweden and Denmark, the formation is truncated by the Basal Middle Jurassic unconformity.[4]

Erratic boulders, called Geschiebe in German, attributed to the Rya Formation and containing iron ooids were found in Holstein, northern Germany.[5]

Subdivision

Döshult Member

The early Sinemurian Döshult Member (Swedish Döshultsledet) comprises coarse-grained cross-layered sandstones and siltstones in the lower part, and is dominated by dark clays and marls rich in marine fossils in the upper part.[6] The member is up to 80 metres (260 ft) thick in the Ängelholm, Helsingborg and Landskrona areas. Presently, the basal part of this member is exposed at three localities in the Helsingborg area. These contain mineralogically and texturally mature, trough cross-bedded sandstones, commonly with herringbone structures showing north and south oriented paleocurrent directions. The occurrence of herringbone structures in well-sorted sand suggests high energy foreshore to subtidal marine depositional conditions for the lower part of the member. In an abandoned quarry in northwest Skåne (Gantofta brickpit in the Helsingborg area, outcrop very limited at present) the upper part of the Döshult Member commences with bioturbated marine nearshore sands, including burrows, as well as abundant marine invertebrate body fossils. This is followed by a bioturbated shelf mudstone with storm-deposited sand and silt intercalations (tempestites). A massive red mudstone with scarce marine body fossils and burrows follows, which is interpreted as having been deposited rapidly, in a low energy but oxidizing environment. The youngest part of the succession comprises siltstones and mudstones, with carbonate-rich beds, deposited in a shallow marine setting.[3] Coaly detritus, muscovite, very small shells and shell fragments, and framboidal pyrite nodules (0.1–0.3 mm in diameter) are characteristic constituents of this member.[7]

Pankarp Member

The dark mudstones of the Döshult Member are overlain by the Pankarp Member (Swedish Pankarpsledet) with a sharp conglomeratic boundary.[8] The late Sinemurian Pankarp Member has an estimated thickness of up to 70 metres (230 ft) in the subsurface of the Ängelholm, Helsingborg and Landskrona areas. In the Kävlinge area, the thickness is about 20 metres (66 ft) thick. In westernmost Skåne, the member has been observed in small diameter drill cores. There, the member is subdivided into a lower unit of variegated clays and shales, a middle, poorly sorted silty to sandy unit including a coal bed, and an upper monotonous mudstone unit which is silty and rich in organic matter at the base, and reddish–greenish at the top. It presents different coloration probably due to different degrees of oxidation of iron in the claystone.[6] In the uppermost part of one core, the Pankarp Member comprises lenticular bedded heteroliths with Planolites burrows.[3]

Katslösa Member
Map of the Katslösa and Rydeback members

The late Sinemurian to early Pliensbachian Katslösa Member (Swedish Katslösaledet) is mainly known from the subsurface in westernmost Skåne, and it has a thickness of 30 to 40 metres (98 to 131 ft) in the Ängelholm, Helsingborg and Landskrona areas. In the Kävlinge area, the thickness is about 75 metres (246 ft). Sedimentological interpretations are mainly based on the results of petrographical studies of museum collections. The Katslösa Member yields a rich marine microfauna and macrofauna, and it is dominated by homogeneous mudstone deposited in a marine low-energy environment. Is composed mostly by marine green, brown and dark gray claystones and sandstones.[6] Thin beds of matrix-rich quartz wackes are common. They are typically mineralogically mature but texturally highly immature with abundant angular sand grains. The matrix comprises organic matter, micrite, mica and clay minerals. In thin section, the sandstones show evidence of intense burrowing, which has obliterated depositional structures. Scattered berthierine ooids, as well as authigenic siderite crystals have been observed in the member.[9]

Rydebäck Member

The late Pliensbachian to late Aalenian Rydebäck Member (Swedish Rydebäcksledet) is up to 70 metres (230 ft) thick in the Ängelholm, Helsingborg and Landskrona areas. It is only known from subsurface material in westernmost Skåne, and sedimentological analysis is based on observations from two wells (Rydebäck–Fortuna-1 and -4). These layers were deposited in small bands during a sea regression, and consist of marine gray, black, green and reddish brown sand and siltstones.[6] The member comprises a uniform succession of muddy arenites with a rich marine microfauna (mostly foraminifera), and represents deposition in an offshore low-energy environment. The sediments are strongly burrowed, which has caused an effective mixing of sand and mud, resulting in the forming of quartz wackes. The sand is quartz-rich, and grains are typically well rounded. Berthierine ooids are common constituents of the sediment.[9]

Depositional environments

Early Jurassic paleogeography

Tethys Sea transgression entailed formation of fossil-bearing marine deposits in Skane, also associated with an increased tectonic activity.[10] Deposition of the Rya Formation began with nearshore coarse clastics, and continued with offshore mudstones with tempestites (the Döshult Member), followed by offshore muddy sediments with a brief non-marine interval (the Pankarp Member), and ended with deposition of open marine low-energy deposits (Katslösa and Rydebäck Members). The marine Rya Formation shows an overall fining-upwards trend, and an up-section bathymetric deepening of the depositional environment. The depositional environment in western Skåne was either physically protected from the storm energy due to basin topography, or deposition in Skåne took place below storm wavebase. Berthierine ooids occur scattered in the Katslösa Member and are increasingly abundant up-section in the Rydebäck Member. There is a possibility that iron ooid formation was promoted by precipitation of iron and silica from volcanic fluids rising up through the substrate, as has been reported from modern marine sediments offshore Indonesia. This hypothesis has emerged with the publication of age data for the volcanic rocks in Skåne, which now appear to be comparable in age to the prominent iron ooid-bearing deposits, i.e. the Rydebäck Member and the Röddinge Formation.[9]

Age

Based on foraminifers, ammonites and ostracods, the Döshult Member is dated to the early Sinemurian, the Pankarp Member to the late Sinemurian, the Katslösa Member to the late Sinemurian to early Pliensbachian and the Rydebäck Member to the late Pliensbachian to late Aalenian.[3]

The formation is time-equivalent with the Röddinge Formation of the Vomb Trough,[2] the Djupadal Formation in central Skane and the Sorthat Formation of Denmark, with which it shares the Spheripollenites–Leptolepidites and Callialasporites–Perinopollenites Zones.[11][12] The formation also correlates with the Fjerritslev Formation of the Danish Basin,[13] and the Gassum Formation of the Øresund Basin.[14] The storm-dominated, hummocky cross-stratified Hasle Formation on Bornholm is contemporaneous with the muddy Katslösa Member of the Rya Formation.[9]

Basin history

The Sorgenfrei-Tornquist Zone (STZ) running through southern Sweden and eastern Denmark is indicated in lightblue
Basement

The basins where the Rya Formation was deposited form part of the Sorgenfrei-Tornquist Zone (STZ) of the Trans-European Suture Zone, the boundary between Baltica to the northeast and Peri-Gondwana to the southwest. The orogeny was active in the Late Ordovician, or approximately 445 million years ago.

At the Carboniferous-Permian boundary around 300 Ma, the area was influenced by the Skagerrak-Centered Large Igneous Province, another large igneous province stretching across the North Sea, the eponymous Skagerrak between Denmark and Sweden and to the northwest up to northern England and Scotland.

Extent of the CAMP
Break-up of Pangea

The basins of southern Sweden and eastern Denmark were formed during the latest Triassic and earliest Jurassic. During this time the Central Atlantic magmatic province (CAMP), with an estimated 11,000,000 square kilometres (4,200,000 sq mi) the largest igneous province in Earth's history, was formed to the present southwest of the Danish-Swedish realm. In the Skåne area, the Central Skåne Volcanic Province was active during the time of deposition of the Rya Formation, commencing around the Sinemurian-Pliensbachian boundary. The earliest magmatism was partly emplaced into and across pre-existing extensional basin structures.[15] The first and the main volcanic phase of this volcanic province occurred in the Early Jurassic (late Sinemurian to Toarcian) at 191–178 Ma.[16] Analysis of the volcanic rocks produced by this Jurassic volcanism suggests a continental Strombolian-type eruptive character close to the oceans of the Early Jurassic.[17] No correlative pyroclastic beds have yet been identified in sedimentary basins surrounding central Skåne.[18]

Toarcian

During deposition of the Rydebäck Member, the Toarcian turnover happened. This event at the Pliensbachian-Toarcian boundary characterized by widespread anoxic conditions globally, led to the extinction of various groups of flora and fauna. Taxa inhabiting the upper water column were unaffected by anoxia and included ammonites and belemnites. Epifaunal taxa adapted to low-oxygen conditions, such as the buchiids, posidoniids and inoceramids, flourished in the post-extinction environment during the survival interval.[19]

Paleogeography of northwestern Europe during the Early Jurassic with Agaleus shark fossil finds. Elevated land areas are shown in grey.

Economic geology

A study on the geothermal potential of reservoirs in the Øresund Basin published in 2018 by Erlström et al. gave results of the formation together with the Gassum and Höganäs Formations, giving the following characteristics of the three Early Jurassic formations:[20]

  • Net sand thickness - 60 to 100 metres (200 to 330 ft)
  • Porosity - 18 to 34%
  • Permeability - 50 to 1500 mD
  • Cl− concentration - 120 to 190 gram/liter
  • Productivity index - 7.0 m3/hr/bar

A study published in the same year analyzing the CO2 storage potential of the Rya and Höganäs Formations concluded a storage capacity of 543 megatons of carbon dioxide.[21]

The organic content of the Jurassic strata in Skåne is typically dominated by gas-prone kerogen (type III), which is below, or at the onset of, thermal maturity.[18]

Paleoenvironment

The sedimentological evolution of the Jurassic in southwestern Skåne, specially the Rya Formation, has provided depth diverse data about its different environments, specially on those samples recovered on the Höllviken-2 core and sidewall cores from the FFC-1 well.[22] The unit was linked on all its sedimentological history with the main Fennoscandinavian land. The local succession in the Höllviken Halfgraben and Barsebäck Platform has measured continue during most of the Triassic and Early Jurassic times.[23] Inte western part of the Höllviken Halfgraben during the Hettangian–Pliensbachian succession there is evidence of higher subsidence rate along the Öresund Fault, indicating tectonically controlled deposition and the presence of a submarine high at the west of this fault.[24] In the east is the Skarup Platform, interpreted as a Jurassic high based on its incomplete or missing Rhaetian–Lower Jurassic strata, along redeposited spores and pollen in the Höllviken Halfgraben and increased sand output, that point to this zone as the major freshwater terrestrial source.[25][26] This deposits shows several nearshore environments, from offshore marine environment probably below wave base on the sinemurian level, as proven by the presence of well cemented fine sand and 2 m thick clay dominated heterolites, that change to more marine influenced flaser bedded heterolites with some strongly bioturbated horizons and convolute beds in the Pliensbachian, and end in the Pliensbachian–Lower Toarcian with a sandy seashore setting with high energy lenticular beds.[27] Some sections are suggested to be tidal flat zones and/or distal bar deltaic deposits.[28] Coeval with the Rydebäck member, at Anholt (Fjerritslev Formation) where studied several levels that point to a connected fluvial system.[29] Concretely the Upper Pliensbachian-Toarcian boundary marks the beginning of a major regression, which continued through the Toarcian and Aalenian.[29] The discovery of the foraminifera Eoguttuliiia liassica, Bony Fishes and Scolecodonts, points that the Pliens-Toar boundary has a shallow water environment, possibly of reduced salinity.[30] The Pollen and spores, identical of those found on the Rydebäck member (Vilhelmsfält Bore No. 1 and Karindal bore no. 1), increase, and the dinoflagellate cysts decrease, suggesting more proximal shoreline.[30] This layers, on both Anholt and the Rydebäck member, are considered to represent a regression from marine inner shelf environments to near lagoonal or deltaic conditions during the late Pliensbachian and early Toarcian, but with light marine influence, as show the presence of Dactylioceras on the Rydebäck member.[30] The sequence points concretely than in both units The Late Pliensbachian sediments at were deposited in a storm-influenced Inner Shelf setting, that changed on the Lower Toarcian into a more restricted, marginal-marine conditions with delta progradation.[31] Then in the late Toarcian increased brackish conditions, to end on a short marine encroachment during the Early Aalenian.[31]

The study of the Jurassic sediments has allowed to know that the basement rocks of southern Sweden were deeply weathered in Late Triassic-Cretaceous times, with formed saprolites on the sub-mesozoic basement and high amount of Kaolinite on the stronger weathered profiles, opposed to smectite on the less weathered ones.[32] Thus, fluvial currents released smectite-rich weathering material to the Late Triassic–Jurassic receiving basins.[32] The members of the Rya Formation recover a transition from deltaic to paralic coast and shallow marine, dominated by kaolinite, along with peaks of illite and smectite.[33] The record of this minerals showed that at the time of deposition of the Rya Formation, mid-latitude warmth and pronounced humidity to drier pseudomediterranean climates allowed on the denuded bedrock in the Fennoscandian Shield, composed of Gneisses or Granites, at places intersected by Dolerite dykes, fracturation and active erosion/weathering.[34] The coeval Djupadal Formation volcanic ash falls at the east Skarup Platform-Cenntral Skane Volcanic Province, may have diluted the marine sediments, with raised smectite content. The pronounced mineralogical maturity of most Swedish post-Norian Mesozoic arenites confirms widespread feldspar destruction in the weathering profiles of the Paleozoic crystalline basement at the north, as a consequence of the increased humidity.[35] A fish tooth recovered from the lower Toarcian of the North West German Basin presents a radiogenic seawater value of −6 ε-units, which is quite counter-intuitive to the idea of massive unradiogenic crustal-derived inputs from Laurasia. Clastic fractions found on the same layer suggest brief radiogenic Nd influxes from the Skåne flood basalts erupted at this time.[36]

Pliensbachian

Lower-late Pliensbachian Layers at Katslösa are similar in faunal composition. The Katslösa Member composition suggest deposition on a Low Energy Marginal marine environment with absence of changes in the salinity (As proven by the presence of Echinoderms and other low salt intolerant fauna), with active bottoms filled by traces of diverse invertebrates and abundance of Bivalves.[9] The Upper Pliensbachian section that mark the appearance of the Rydebäck Member continues the stable salinity environment with increased presence of Echinoderms, but lack of a diverse Bivalve fauna as in the older section. This unit was deposited likely on a marine setting more proximal to the shore and low-energy, more strongly burrowed than older layers. The presence of Selachian fauna can be more likely derived from a more suitable depositional setting for this remains than a biota turnover.[37] Palynology in this section is dominated by spores in the Karindal bore no. 1, suggesting a humid climate on nearby emerged lands.[26]

Toarcian

Toarcian layers of the formation where influenced by the ongoing vulcanism located in the Central Skåne Volcanic Province (Djupadal Formation), as, marine water influence is observed in the main outcrop of the last unit, where enriched Zeolite by Barium is suggested to derive from oceanic water, that may have circulated as hydrothermal flows in the lapilli tuff and facilitating diagenetic changes.[38] Is also known by palynological analysis on the Bonnarp Cone, where saltwater/brackish acritarchs like Leiosphaera and Leiofusa or Dinoflajellates like Nannoceratopsis where recovered.[39] This section also recovers the increased influence of Terrestrial weathering measured in the layers after the Toarcian AOE, as seen in the Vilhelmsfalt borehole, where plant remains increase their presence, suggesting a regression of the coast influenced by the increased volcanic-derived materials.[40] The Palynology in this section is dominated by the Pollen of Chasmatosporites (Cycads), with at least six species recovered, playing more than 50% of the total palynological samples, implicating a clear dominant role of the producer of this Pollen and suggesting along the increased amount of Cheirolepidiaceae pollen and general decrease of spores a shif towards a more arid climate on nearby settings.[41]

Fossil content

The formation has provided fossils of typically marine fauna. With the exception of a continental coal bed, the formation is marine in character. Shark teeth were reported from the Rydebäck Member.[42]

Annelida

Echinodermata

  •  ?Acrosaleniidae Indeterminate[37]
  • Balanocrinus subteroides[45]
  • Hispidocrinus scalaris[45]
  • Isocrinus ranae[37]
  • Pentacrinus basaltiformis[37]

Ammonites

Belemnites

Brachiopods

Bivalves

Gastropods


Scaphopods

Branchiopoda

Ostracods

  • Astacolus denticulina carinata[47]
  • Brizalina liassica amalthea[47]
  • Citharina inaequistriata[47]
  • Cristacythere crassireticulata[55]
  • Marginulina spinata spinata[47]
  • Nanacythere simplex[47]
  • Ogmoconchella danica[55]
  • Ogmoconchella mouhersensis[55]
  • Ogmoconchella adenticulata[47]
  • Ogmoconcha amalthei amalthei[55]
  • Pleurifera harpa[55]

Mites

Ichnofossils

Fish

Apart from a few teeth of the hybodont Hybodus reticulatus, the shark fauna from the Rya Formation is exclusively neoselachian.[146]

Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
 Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
Genus/Family Species Location Member Material Notes/Affinities Images

Archaeotolithus[147][148]
  • Archaeotolithus bornholmiensis

Katslösa, Bed 30

Katslösa Member
  • 20 Otoliths

Interpreted as sagittal otoliths of a Palaeonisciformes Indet.[149] Originally assigned to the extant family Sciaenidae, then from members of Palaeoniscidae and finally some recent research suggest affinities with Pholidophoriformes. Bony Fish Otoliths Similar to ones recovered from the Hasle Formation.[150]

Acrodus[147]
  • Acrodus sp.

Katslösa, Bed 42
  • Small Fragment of Tooth Crown

A marine Shark, member of the Acrodontidae.

Actinopterygii[151]
  • Actinopterygii Indeterminate

Road between the hamlets of Rya and Katslosa

Rydebäck Member
  • Scales
  • Ganoid Scales

Incertae Sedis Bony Fish Scales

Agaleus[151]
  • Agaleus dorsetensis
  • Four incomplete teeth

A Marine Shark of the Family Agaleidae

Chimaeriformes[151]
  • Chimaeriformes Indeterminate
  • Fragmentary Remains

Incertae Sedis fragmentary remains of Chimaeras

Hybodus[151]
  • Hybodus reticulatus
  • Fin Spine
  • Multiple Teeth

A marine Shark, member of the Hybodontiformes. Related to Hybodus hauffianus and other genera from the south of Germany. Only non-Neoselachian recovered from the location.

Hexanchidae[151]
  • Hexanchidae indet.
  • Single incomplete tooth

Marine Cow Sharks, Incertae Sedis Inside Hexanchidae. Isolated teeth appear to be similar to Hexanchus arzoeensis.

Paraorthacodus[152]
  • Paraorthacodus sp.
  • Three broken teeth

Marine Sharks of the Family Palaeospinacidae. One of the earliest records of the genus Paraorthacodus, together with others from France.

Palidiplospinax[153]
  • Palidiplospinax occultidens
  • Palidiplospinax enniskilleni
  • Palidiplospinax? sp.
  • Teeth
  • One complete scale

Marine Sharks of the Family Palaeospinacidae. The complete scale described above most closely resembles scales from Synechodus pinnai.

"Synechodus"[151]
  • "Synechodus" sp.
  • One complete tooth

Marine Sharks of the Family Palaeospinacidae. The tooth described may represent a new species.

Sphenodus[154]
  • Sphenodus sp.
  • Two complete teeth

Marine Sharks of the Family Orthacodontidae.

Plant Remains

Coalified wood occurs as scattered pieces up to 6 centimetres (2.4 in) long and indeterminate belemnites, echinoids, serpulids, ostracods and nodosariid foraminifera were also recorded in the formation.[7] Ammonites where found on layers with plant fossils. Fragments of Dactylioceras were found at a depth of 170 m in the Vilhelmsfalt borehole, together with plant remains and beautifully preserved Pelecypods.[155] The plant material, probably derived from watercourses emptying in the neighborhood, consists mostly of small fragments that seem to have been intimately sedimented with the muddy material that flocculated on meeting the seawater.[155] Other specimen, Arnioceras sp. indet. occurs on what appears to be a transitional environment, probably a back-beach with coalified fragments of plants, some of a large size. Probably due to the shell being washed to land due to a storm.[155]

Palynology
Genus Species Stratigraphic position Member Material Notes Images

Botryococcus[25][26][156]
  • Botryococcus braunii
  • Botryococcus calcareus
  • Botryococcus nsp. A
  • Botryococcus nsp. B
  • Vilhelmsfält Bore No. 1
  • Karindal bore no. 1

Rydebäck Member
  • Miospores

Type Genus of the Botryococcaceae inside Trebouxiales. A colonial green microalga related with freshwater and brackish ponds and lakes around the world, where it often can be found in large floating masses. This genus indicates towards the Toarcian section an increasing amount of terrestrial components, as well decreasing salinit in the nearby emerged Skarup Platform.[157]
Extant Specimen

Schizosporis[158]
  • Schizosporis sp.

A Green Algae, member of Zygnemataceae

Tasmanites[26]
  • Tasmanites sp.

A Green Algae, member of Pyramimonadaceae

Campenia[159]
  • Campenia gigas

A Green Algae, member of Prasinophyceae

Nannoceratopsis[25][160]
  • Nannoceratopsis gracilis
  • Nannoceratopsis senex
  • Cysts

A Dinoflajellate, member of the family Nannoceratopsiaceae. It is a genus related with Marine deposits.

Veryhachium[161]
  • Veryhachium sp.

A Dinoflajellate, Dinophyceae familia incertae sedis

Foraminisporis[159][162]
  • Foraminisporis jurassicus
  • Spores

Affinities with the family Notothyladaceae inside Anthocerotopsida. Hornwort spores.
Example of extant Notothylas specimens, Foraminisporis come probably from similar genera

Neochomotriletes[163]
  • Neochomotriletes triangularis

Taurocusporites[25]
  • Taurocusporites verrucatus

Polycingulatisporites[164]
  • Polycingulatisporites circulus

Staplinisporites[159]
  • Staplinisporites caminus

Affinities with the family Encalyptaceae inside Bryopsida. Branching Moss Spores, related with high water-depleting environments
Example of extant Encalypta specimens, Staplinisporites come probably from similar genera

Stereisporites[165][166]
  • Stereisporites aulosenensis
  • Stereisporites cicatricosus
  • Sulcatisporites pinoides

Affinities with the family Sphagnaceae inside Sphagnopsida. "Peat moss" spores, related to genera such as Sphagnum that can store water large amount of water.
Example of extant Sphagnum specimens, Stereisporites, Sculptisporis and Rogalskaisporites come probably from similar genera

Lycopodiumsporites[165][167]
  • Lycopodiumsporites clavatoides
  • Lycopodiumsporites pseudoreticulatus
  • Lycopodiumsporites reticulumsporites
  • Lycopodiumsporites semimuris
  • Lycopodiumsporites vilhelmii
  • Lycopodiumsporites sp.
  • Lycopodiumsporites sp sp. nov.
  • Lycopodiumsporites sp.1 sp. nov.
  • Lycopodiumsporites sp.2 sp. nov.

Affinities with the family Lycopodiaceae inside Lycopodiopsida. Lycopod spores, related with herbaceous to arbustive flora common on humid environments
Extant Lycopodium specimens. Genera like Lycopodiumsporites probably come from a similar plant

Foveosporites[163][166]
  • Foveosporites moretonensis
  • Fovesporites foveoreticulatus

Lycospora[161]
  • Lycospora salebrosacea

Semiretisporis[163][161]
  • Semiretisporis cf. gothae

Sestrosporites[168][166]
  • Sestrosporites pseudoalveolatus

Ceratosporites[159]
  • Ceratosporites spinosus

Affinities with the Selaginellaceae inside Lycopsida. Herbaceous Lycophyte flora, similar to Ferns, ralated with Humid Settings. This Family of Spores are also the most diverse on the Formation.
Extant Selaginella, typical example of Selaginellaceae. Genera like Ceratosporites and Densoisporites probably come from a similar or a related Plant

Densoisporites[168][166]
  • Densoisporites crassus
  • Densoisporites erdtmanii
  • Densoisporites scanicus
  • Densoisporites velatus

Heliosporites[169]
  • Heliosporites altmarkensis

Cepulina[165][166]
  • Cepulina truncata

Neoraistrickia[170]
  • Neoraistrickia truncata
  • Neoraistrickia sp sp. nov.

Uvaesporites[171]
  • Uvaesporites argenteaeformis
  • Uvaesporites puzzlei
  • Uvaesporites sp.

Calamospora[172]
  • Calamospora mesozoica

Affinities with the Calamitaceae inside Equisetales. Horsetails, herbaceous flora related to high humid environments, flooding tolerant plants.
Recosntruction of the Genus Calamites, found associated with Calamospora

Conbaculatisporites[163][164]
  • Conbaculatisporites mesozoicus

Incertae Sedis affinities with the Pteridophyta. Uncertain Pteridophyte origin

Tigrisporites[166]
  • Tigrisporites microrugulatus

Converrucosisporites[170]
  • Converrucosisporites sp sp. nov.

Verrucosisporites[161]
  • Verrucosisporites obscurilaesuratus

Cingutriletes[166]
  • Cingutriletes sp.

Laevigatisporites[161]
  • Laevigatisporites sp.

Convolutispora[164]
  • Convolutispora sp.

Ischyosporites[168][166][173]
  • Ischyosporites sp. Guy-.,1986 sp. nov.
  • Ischyosporites variegatus

Simozonotriletes[171]
  • Simozonotriletes arcuatus

Platyptera[174]
  • Platyptera sp.1 Guy-.,1986 sp. nov.
  • Platyptera sp.2 sp. nov.

Trachysporites[174][169]
  • Trachysporites fuscus
  • Trachysporites asper
  • Trachysporites sp sp. nov.

Leptolepidites[163][161]
  • Leptolepidites bossus
  • Leptolepidites equatibossus
  • Leptolepidites macroverrucosus
  • Leptolepidites major
  • Leptolepidites paverus
  • Leptolepidites rotundus
  • Leptolepidites sp.1 sp. nov.
  • Leptolepidites sp.2 sp. nov.

Affinities with the family Dennstaedtiaceae inside Polypodiales. Forest Fern Spores
Example of extant Dennstaedtia specimens, Leptolepidites come probably from similar genera

Lophotriletes[168][161]
  • Lophotriletes verrucosus

Affinities with the family Botryopteridaceae inside Polypodiales. Forest Fern Spores

Apiculatisporites[162]
  • Apiculatisporites parvispinosus

Affinities with the family Zygopteridaceae inside Zygopteridales.

Lycopodiacidites[168][166]
  • Lycopodiacidites rugulatus

Affinities with the Ophioglossaceae inside Filicales. Fern spores from lower herbaceous flora
Example of extant Helminthostachys specimens, Lycopodiacidites come probably from similar genera or maybe a species from the genus

Contignisporites[25][164][175]
  • Contignisporites dunrobinensis
  • Contignisporites problematicus

Affinities with the Pteridaceae inside Polypodiopsida. Forest Ferns from humid ground locations
Example of extant Pityrogramma specimens, Contignisporites and Manumia come probably from similar genera or maybe a species from the genus

Auritulinasporites[159]
  • Auritulinasporites scanicus
  • Auritulinasporites triclavis

Affinities with the Gleicheniales inside Polypodiopsida. Fern spores from lower herbaceous flora
Example of extant Gleichenia specimens, Gleicheniidites and Auritulinasporites come probably from similar genera or maybe a species from the genus

Gleicheniidites[25][160]
  • Gleicheniidites senonicus
  • Gleicheniidites umbonatus

Dictyophyllidites[166]
  • Dictyophyllidites harrisii

Affinities with the Dipteridaceae inside Gleicheniales. Fern spores from lower herbaceous flora
Example of extant Dipteris specimens, Dictyophyllidites come probably from similar genera or maybe a species from the genus

Marattisporites[170][167]
  • Marattisporites scabratus

Affinities with the Marattiaceae inside Polypodiopsida. Fern spores from lower herbaceous flora
Example of extant Marattia specimens, Marattisporites come probably from similar genera

Matonisporites[176][169]
  • Matonisporites crassiangulatus
  • Matonisporites sp.

Affinities with the Matoniaceae inside Polypodiopsida. Fern spores from lower herbaceous flora
Example of extant Matonia specimens, Matonisporites come probably from similar genera

Osmundacidites[172][177]
  • Osmundacidites wellmanii

Affinities with the family Osmundaceae inside Polypodiopsida. Near Fluvial currents ferns, reted to the modern Osmunda Regalis.
Example of extant Osmunda specimens, Osmundacidites and Baculatisporites come probably from similar genera or maybe a species from the genus

Baculatisporites[25][162]
  • Baculatisporites comaumensis

Todisporites[178][177]
  • Todisporites granulatus
  • Todisporites major
  • Todisporites minor

Cibotiumspora[25][162]
  • Cibotiumspora jurienensis

Affinities with the family Cyatheaceae inside Cyatheales. Arboreal Fern Spores
Example of extant Cyathea, Zebrasporites and Cibotiumspora come probably from similar genera

Zebrasporites[179][180]
  • Zebrasporites interscriptus

Enzolasporites[163][161]
  • Enzolasporites manifestus
  • Enzolasporites vigens

Cyathidites[25][160]
  • Cyathidites australis
  • Cyathidites concavus
  • Cyathidites minor

Brachysaccus[159][26]
  • Brachysaccus microsaccus
  • Pollen

Incertae sedis Affinities with the Gymnospermophyta

Alisporites[159][26]
  • Alisporites robustus

Affinities with the families Umkomasiaceae, Peltaspermaceae and Corystospermaceae inside Peltaspermales. Possible seed Fern Pollen

Protopinus[171][169]
  • Protopinus scanicus
  • Protopinus sp. sp nov

Affinities with the families Caytoniaceae inside Caytoniales. Possible seed Fern Pollen

Vitreisporites[181][180]
  • Vitreisporites pallidus

Aratisporites[159]
  • Aratisporites palettae

Eucommiidites[163][160][182]
  • Eucommiidites troedssonii
  • Eucommiidites major

Type Pollen of the Erdtmanithecales, that can be related with the Gnetales.

Chasmatosporites[25][162][183]
  • Chasmatosporites apertus
  • Chasmatosporites elegans
  • Chasmatosporites major
  • Chasmatosporites minor
  • Chasmatosporites rimatus

Affinities with the family Cycadaceae inside Cycadales. Is among the most abundant flora recovered on the upper section of the formation, and was found to be similar to the pollen of the extant Encephalartos laevifolius. On the Toarcian-Aalenian section, at least six species of Chasmatosporites are recovered, playing more than 50% of the total palynological samples, implicating a clear dominant role of the producer of this Pollen.[184] C. minor is present more on sandy brackish facies, while C. major is found more on brackish-marine facies.[184]
Extant Encephalartos laevifolius. Chasmatosporites maybe come from a related plant

Ginkgocycadophytus[165][167]
  • Ginkgocycadophytus nitidus
  • Ginkgocycadophytus sp.

Affinities with the Ginkgoaceae inside Ginkgoales.

Entylissa[25][160]
  • Entylissa pyriformis
  • Entylissa reticulata
  • Entylissa tecta

Affinities with the Palissyaceae inside Pinopsida.

Classopollis[159][162][185]
  • Classopollis classoides

Affinities with the Cheirolepidiaceae inside Pinopsida.

Spheripollenites[178]
  • Spheripollenites subgranulatus

Cerebropollenites[25][160][186]
  • Cerebropollenites mesozoicus

Affinities with the family Pinaceae inside Pinopsida. Conifer pollen from medium to large arboreal plants
Extant Picea. Cerebropollenites and Pinuspollenites maybe come from a related plant

Ovalipollis[174][169]
  • Ovalipollis ovalis

Pinuspollenites[181][177]
  • Pinuspollenites globosaccus
  • Pinuspollenites minimus

Pityosporites[181][177]
  • Pityosporites nigraeformis
  • Pityosporites scaurus

Parvisaccites[181][177]
  • Parvisaccites enigmatus

Affinities with the Podocarpaceae inside Pinopsida. Conifer pollen from medium to large arboreal plants
Extant Podocarpus. Podocarpidites and Parvisaccites maybe come from a related plant

Podocarpidites[181][177]
  • Podocarpidites ellipticus
  • Podocarpidites sp.

Exesipollenites[25][160]
  • Exesipollenites tumulus

Affinities with the family Cupressaceae inside Pinopsida. Pollen that resembles 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

Perinopollenites[181][177]
  • Perinopollenites elatoides

Callialasporites[159][26]
  • Callialasporites dampieri
  • Callialasporites turbatus

Affinities with the family Araucariaceae inside Pinales. Conifer Pollen from medium to large Arboreal Plants
Extant Araucaria. Araucariacites and Callialasporites maybe come from a related plant

Araucariacites[159][26][187]
  • Araucariacites australis
Plant Macrofossils
Genus Species Stratigraphic position Member Material Notes Images

Coniopteris[188]
  • Coniopteris hymenophylloides
  • West of Heiligendamm

Katslösa Member
  • Single Leaflet

A fern of the family Polypodiidae inside Polypodiopsida. Similar to Sphenopteris species of Permian rocks of the same region
Specimen

Ginkgoites[189]
  • Ginkgoites acosmia
  • Single Leaf Impression

A member of the family Ginkgoaceae inside Ginkgoales. The shape and the clearly visible bifurcation of the veins agrees with the referral to the genus Ginkgoites

Marchantiolites[190]
  • Marchantiolites cf. porosus
  • Almost complete thallus of a liverwort

A member of the family Marchantiaceae inside Hepatophyta. Suggested initially to be an angiosperm leaf. Like extant Marchantia polymorpha, mostly blackish stripes appear along the midrib of the thallus.
Extant Marchantia polymorpha

See also

References

  1. ↑ Ahlberg et al., 2003, p.528
  2. 1 2 Ahlberg et al., 2003, p.530
  3. 1 2 3 4 Ahlberg et al., 2003, p.533
  4. ↑ Erlström et al., 2018, p.129
  5. ↑ Hinz-Schallreuter & Schallreuter, 2009, p.2
  6. 1 2 3 4 Andersson & Hybertsen, 2010-8-17
  7. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Frandsen & Surlyk, 2003, p.547
  8. ↑ Frandsen & Surlyk, 2003, p.550
  9. 1 2 3 4 5 Ahlberg et al., 2003, p.534
  10. ↑ Greiff, 2019-8
  11. ↑ Nielsen et al., 2003, p.588
  12. ↑ Nielsen et al., 2003, p.590
  13. ↑ Frandsen & Surlyk, 2003, p.543
  14. ↑ Erlström et al., 2018, p.138
  15. ↑ Bryan & Ferrari, 2013, p.1058
  16. ↑ Bergelin, 2009
  17. ↑ Augustsson, 2009
  18. 1 2 Ahlberg et al., 2003, p.539
  19. ↑ Harries & Little, 1999
  20. ↑ Erlström et al., 2018, p.139
  21. ↑ Sjöberg, 2018, p.90
  22. ↑ Bou Daher (2012)-p10
  23. ↑ Bou Daher (2012)-p7
  24. ↑ Bou Daher (2012)-p37
  25. 1 2 3 4 5 6 7 8 9 10 11 12 13 Guy-Ohlson, 1986-p3
  26. 1 2 3 4 5 6 7 8 Guy-Ohlson, 1990-p218
  27. ↑ Bou Daher (2012)-p14
  28. ↑ Bou Daher (2012)-p15
  29. 1 2 Seidenkrantz & Koppelhus (1993)-p201
  30. 1 2 3 Seidenkrantz & Koppelhus (1993)-p210
  31. 1 2 Nagy, J. (2003)-p28
  32. 1 2 Ahlberg, Olsson & Šimkevičius, 2003-p15
  33. ↑ Ahlberg, Olsson & Šimkevičius, 2003-p17
  34. ↑ Ahlberg, Olsson & Šimkevičius, 2003-p19
  35. ↑ Ahlberg, Olsson & Šimkevičius, 2003-p21
  36. ↑ Dera, Prunier, Smith, Haggart, Popov, Guzhov, Rogovf, Delsate, Detlev,Pucéatj, Charbonnierk & Germain (2015)-p1610
  37. 1 2 3 4 Hunter, A.W. & Rees, J.(2010)-p10-24
  38. ↑ Augustsson (2001)-p28
  39. ↑ Bölau & Kockel-Brosius (1965)-p55
  40. ↑ Dera, Prunier, Smith, Haggart, Popov, Guzhov, Rogovf, Delsate, Detlev,Pucéatj, Charbonnierk & Germain (2015)-p1611
  41. ↑ Guy-Ohlson, 1988-p11
  42. ↑ Rya Formation at Fossilworks.org
  43. ↑ Weitschat & Gründel (2002)-p39
  44. 1 2 3 Troedsson (1951)-p146-147
  45. 1 2 Troedsson (1951)-p15
  46. ↑ Reyment (1959)-p118
  47. 1 2 3 4 5 6 7 8 9 10 11 Sivhed, U. 1984-p19
  48. ↑ Doguzhaeva, Mutvei & Weitschat (2003)-p83
  49. ↑ Reyment (1959)-p129
  50. ↑ Reyment (1959)-p130
  51. 1 2 3 4 5 6 Frandsen & Surlyk, 2003, p.545
  52. ↑ Reyment (1959)-p110
  53. ↑ Reyment (1959)-p134
  54. ↑ Reyment (1959)-p142
  55. 1 2 3 4 5 6 7 8 9 Sivhed, U. (1984)- p.251
  56. ↑ Lehmann (1961)-p42-43
  57. ↑ Reyment (1959)-p132
  58. ↑ Reyment (1959)-p112
  59. ↑ Reyment (1959)-p136
  60. ↑ Reyment (1959)-p114
  61. ↑ Doguzhaeva, Mutvei & Weitschat (2003)-p86
  62. ↑ Reyment (1959)-p139
  63. ↑ Reyment (1959)-p137
  64. ↑ Doguzhaeva, Mutvei & Weitschat (2003)-p80
  65. ↑ Doguzhaeva, Mutvei & Weitschat (2003)-p81
  66. 1 2 3 4 5 Troedsson (1951)-p243-245
  67. ↑ Doguzhaeva, Mutvei & Weitschat (2003)-p79
  68. 1 2 3 4 Troedsson (1951)-p148-150
  69. ↑ Troedsson (1951)-p224
  70. 1 2 Troedsson (1951)-p185
  71. 1 2 Troedsson (1951)-p192
  72. 1 2 Troedsson (1951)-p193
  73. ↑ Troedsson (1951)-p167
  74. 1 2 Troedsson (1951)-p168
  75. ↑ Troedsson (1951)-p169
  76. ↑ Troedsson (1951)-p170
  77. 1 2 Troedsson (1951)-p171
  78. ↑ Troedsson (1951)-p159
  79. ↑ Troedsson (1951)-p160
  80. ↑ Troedsson (1951)-p161
  81. ↑ Troedsson (1951)-p162
  82. ↑ Troedsson (1951)-p163
  83. ↑ Troedsson (1951)-p211
  84. ↑ Troedsson (1951)-p212
  85. 1 2 Troedsson (1951)-p213
  86. ↑ Troedsson (1951)-p214
  87. 1 2 Troedsson (1951)-p220
  88. 1 2 Troedsson (1951)-p216
  89. ↑ Troedsson (1951)-p217
  90. ↑ Troedsson (1951)-p218
  91. 1 2 Troedsson (1951)-p181
  92. ↑ Troedsson (1951)-p184
  93. ↑ Troedsson (1951)-p203
  94. ↑ Troedsson (1951)-p204
  95. ↑ Troedsson (1951)-p205
  96. ↑ Troedsson (1951)-p206
  97. ↑ Troedsson (1951)-p195
  98. 1 2 Troedsson (1951)-p154
  99. ↑ Troedsson (1951)-p157
  100. ↑ Troedsson (1951)-p158
  101. ↑ Troedsson (1951)-p188
  102. ↑ Troedsson (1951)-p189
  103. ↑ Troedsson (1951)-p191
  104. 1 2 Troedsson (1951)-p207
  105. ↑ Troedsson (1951)-p225
  106. 1 2 Troedsson (1951)-p208
  107. 1 2 Troedsson (1951)-p210
  108. ↑ Troedsson (1951)-p226
  109. ↑ Troedsson (1951)-p227
  110. 1 2 Troedsson (1951)-p228
  111. ↑ Troedsson (1951)-p231
  112. 1 2 Troedsson (1951)-p232
  113. 1 2 Troedsson (1951)-p234
  114. ↑ Troedsson (1951)-p233
  115. 1 2 Troedsson (1951)-p151
  116. ↑ Troedsson (1951)-p152
  117. 1 2 Troedsson (1951)-p200
  118. ↑ Troedsson (1951)-p149
  119. ↑ Troedsson (1951)-p150
  120. ↑ Troedsson (1951)-p187
  121. ↑ Troedsson (1951)-p194
  122. ↑ Troedsson (1951)-p221
  123. 1 2 3 Troedsson (1951)-p178
  124. ↑ Troedsson (1951)-p179
  125. ↑ Troedsson (1951)-p219
  126. 1 2 3 Troedsson (1951)-p173
  127. ↑ Troedsson (1951)-p153
  128. ↑ Troedsson (1951)-p175
  129. 1 2 Troedsson (1951)-p176
  130. ↑ Troedsson (1951)-p177
  131. ↑ Troedsson (1951)-p223
  132. ↑ Troedsson (1951)-p165
  133. ↑ Troedsson (1951)-p166
  134. ↑ Troedsson (1951)-p172
  135. 1 2 3 4 5 6 7 Troedsson (1951)-p238-240
  136. ↑ Weitschat & Gründel (2002)-p40
  137. ↑ Engeser & Riedel (1992)-p49-50
  138. 1 2 Troedsson (1951)-p230-245
  139. ↑ Engeser & Riedel (1992)-p46
  140. ↑ Engeser & Riedel (1992)-p43
  141. ↑ Engeser & Riedel (1992)-p47
  142. ↑ Engeser & Riedel (1992)-p48
  143. ↑ Peng, Slater, McLoughlin & Vajda, (2023)-p4-20
  144. ↑ Sivhed & Wallwork (1978)-p65-66
  145. 1 2 3 4 5 6 Frandsen & Surlyk, 2003, p.546
  146. ↑ Rees, 2000, p.411
  147. 1 2 Troedsson (1951)-p246
  148. ↑ Schwarzhans (2018)-p79
  149. ↑ Schwarzhans (2018)-p112
  150. ↑ Schwarzhans (2018)-p83
  151. 1 2 3 4 5 6 Rees, 2000, p.412
  152. ↑ Rees, 2000, p.416
  153. ↑ Rees, 2000, p.417-18
  154. ↑ Rees, 2000, p.415
  155. 1 2 3 Reyment (1959)-p152
  156. ↑ Guy-Ohlson, 1996-p250
  157. ↑ Guy-Ohlson, 1996-p252
  158. ↑ Guy-Ohlson, 1990-p217
  159. 1 2 3 4 5 6 7 8 9 10 11 Guy-Ohlson, 1986-p2
  160. 1 2 3 4 5 6 7 Guy-Ohlson, 1990-p220
  161. 1 2 3 4 5 6 7 8 Guy-Ohlson, 1990-p222
  162. 1 2 3 4 5 6 Guy-Ohlson, 1990-p219
  163. 1 2 3 4 5 6 7 Guy-Ohlson, 1986-p4
  164. 1 2 3 4 Guy-Ohlson, 1990-p221
  165. 1 2 3 4 Guy-Ohlson, 1986-p5
  166. 1 2 3 4 5 6 7 8 9 10 Guy-Ohlson, 1990-p223
  167. 1 2 3 Guy-Ohlson, 1990-p224
  168. 1 2 3 4 5 Guy-Ohlson, 1986-p6
  169. 1 2 3 4 5 Guy-Ohlson, 1990-p225
  170. 1 2 3 Guy-Ohlson, 1986-p7
  171. 1 2 3 Guy-Ohlson, 1986-p10
  172. 1 2 Guy-Ohlson, 1986-p9
  173. ↑ Guy-Ohlson, 1988-p807
  174. 1 2 3 Guy-Ohlson, 1986-p11
  175. ↑ Guy-Ohlson, 1988-p808
  176. ↑ Guy-Ohlson, 1986-p8
  177. 1 2 3 4 5 6 7 Guy-Ohlson, 1990-p226
  178. 1 2 Guy-Ohlson, 1986-p12
  179. ↑ Guy-Ohlson, 1986-p16
  180. 1 2 Guy-Ohlson, 1990-p227
  181. 1 2 3 4 5 6 Guy-Ohlson, 1986-p15
  182. ↑ Guy-Ohlson, 1988-p809
  183. ↑ Guy-Ohlson, 1988-p810
  184. 1 2 Guy-Ohlson, 1988-p6-12
  185. ↑ Guy-Ohlson, 1988-p812
  186. ↑ Guy-Ohlson, 1988-p813
  187. ↑ Guy-Ohlson, 1988-p815
  188. ↑ Roselt (1968)-p65
  189. ↑ Roselt (1968)-p66
  190. ↑ Roselt (1968)-p68

Bibliography

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