Djupadal Formation

The Djupadal Formation is a geologic formation in Skåne County, southern Sweden. It is Early Jurassic (probably Pliensbachian-Toarcian, or Late Toarcian) in age. It is part of the Central Skåne Volcanic Province, know by the discovery of basalt tuff layers, including Sandåkra, Korsaröd and Djupadal. An original analysis of the location of Korsaröd led to a Toarcian-Aalenian age,[3][4][5] but was dismissed in 2016, when a series of Palynogical samples recovered a Late Pliensbachian and probably Lower Toarcian age for the Korsaröd Outcrop.[6] The same year this result was also challenged by an in-depth study of the Lilla Hagstad neck that yield a Late Toarcian Age.[7] The formation was deposited in the Central Skane region, linked to the late early Jurassic volcanism. The Korsaröd member includes a volcanic-derived Lagerstatten with exceptional fern finds.[8] The data provided by fossilized wood rings showed that the location of Korsaröd hosted a middle-latitude Mediterranean-type biome in the late Early Jurassic, with low rainfall ratio, delayed to seasonal events. Superimposed on this climate were the effects of a local active Strombolian Volcanism and hydrothermal activity.[9]

Djupadal Formation
Stratigraphic range: Latest Pliensbachian- Latest Toarcian
~A Volcanic neck suggest 176.7 ± 0.5 Ma, Late Toarcian Age
About 10 m south of the shore of the Korsaröd lake exposed layers of the unit are known
TypeFormation
Unit ofCentral Skåne Volcanic Province
Sub-units(in part) Sapropel at Sandåkra
UnderliesCuaternary Sediments
OverliesHöör Sandstone, Brandsberga and Kolleberga erratics and (in part) Sapropel at Sandåkra
ThicknessUp to 60 m (200 ft)[1]
Lithology
PrimaryBasalt Tuff, Veined Gneiss[1]
OtherSandstone, Clay and Conglomerate
Location
Coordinates55.98°N 13.63°E / 55.98; 13.63
Approximate paleocoordinatesApprox. 35°N
RegionCentral Skåne County
Country Sweden
Extent1,000 km2 (390 sq mi)
Type section
Named forDjupadalsmölla, Ljungbyhed
Named byCarita Augustsson[2]
Year defined2001
Djupadal Formation is located in Sweden
Djupadal Formation
Djupadal Formation (Sweden)

Korsaröd Lagerstätten Location

Description

Djupadalsmölla, the original location has been known due to its volcanic Tuff and other volcanoclastic-derived facies.[10] It was originally described in 1826 as the local basalts where identified as coming from ancient volcanic eruptions.[11] The exposure of Anneklev, located near Höör was where the first volcanic neck was discovered, with other volcanic remnants mentioned on Jällabjär and Rallate along with the tuff at Djupadalsmölla.[12] These deposits were described as volcanic rocks composed mostly by tuff, that includes basaltic bombs (composed by Pyroxene and pseudomorphs from this, along with Olivine on adjacent pyroclastics), accidental lithics and occasional wood from conifers, in the form of small pieces to large logs.[13] At Djupadal the Rönne River has cut a 20 m deep valley mainly in the Precambrian basement, where Lower Jurassic strata (including volcanic tuff) form part of the southern valley side, and also occur in the valley northwestwards, indicating that the valley is partly an exhumed sub-Jurassic depression.[4] The modern strata of the Valley is over a Late Weichselian melt-water channel and it also contains eroded fluvioglacial deposits.[14] A nearby roadcut shows kaolinized basement beneath the Toarcian sediments and a nearby boring has penetrated 44 m of kaolinized Gneiss.[4] The Valley Bottom exposes small ridges and cupolas (in gneiss) and tors (mainly in Amphibolite).[4]

Geology

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

The southwestern margin of the Baltic Shield in southern Sweden and adjoining off-shore basins in the Baltic Sea is characterized by a 20–60 km wide fracture zone along which Mesoproterozoic basement rocks were downfaulted and deeply buried beneath sedimentary cover of the Danish Basin and Polish Trough.[15] The so-called Sorgenfrei–Tornquist Zone (STZ) forms part of the Tornquist Lineament, a trans-continental deep fracture system that extends for more than 1000 km from the North Sea to the Black Sea, demarcating the largely unexposed southwestern margin of the East European Craton.[16] Towards the northern zone there are 2 two Mesoproterozoic basement complexes: the uplifted Southwest Scandinavian Gneiss Complex of the Baltic Shield and the deeply buried Danish Massif. The southern branch of the lineament forms the Tornquist–Teisseyre Zone (TTZ), which separates the Precambrian lithosphere of the East European Craton from accreted Early Paleozoic peri-Gondwanan terranes of central Europe.[17] The STZ was result of continental rifting during the Permian-Carboniferous periods, with some zones continued until the Cretaceous, with an evolution similar to the North Sea system.[18] This zone is known by its complex horst and graben structures that resulted from Late CretaceousPaleogene inversion tectonics.[19] As result of the collapse of the Variscan Orogen 200 million years of intermittent lithospheric extension developed locally and in sister systems, developing Permian rift structures and basins, that were reactivated during the breakup of Pangea and opening of the Tethys & North Atlantic oceans during the Mesozoic.[20] The systemic rifting from Carboniferous to Mesozoic in the STZ was accompanied by low-volume mafic alkaline magmatism, with a 70 km wide NW-SE oriented Permo-Carboniferous Tholeiitic dyke swarm that runs across Scania and Bornholm island within the STZ.[21] In this context, the Central Skåne Volcanic Province was developed, being a 30 by 40 km large Mesozoic volcanic field that occurs at the southern tip of the Scandinavian Peninsula, composed by 100 volcanic plugs and necks form prominent steep sided hills in central Scania.[22] In this volcanic field either low-volume alkaline magmatism was repeatedly sourced from a common enriched Continental Lithospheric Mantle reservoir or was derived from edge-driven mantle convection by which newly inflowing asthenospheric material melted in structurally controlled positions.[23]

Stratigraphy

The Djupadalsmölla pyroclastic reaches almost 10 m high and 20 m wide, with pyroclastics appearing also through the west more than 100 m, along the valley of the Rönne River.[24] It is composed by a 3 m thick sequence of Jurassic rocks, starting overlying the kaolinized basement of Paleozoic Gneiss with 2 m of sandstone-claystone series ending with a single metre of green-brownish turfaceous rocks.[25] The strata is composed by mostly small Lapilli (around 30–50% of the content) and ash (-10 mm), with some samples being red in patches. Tuff concretions are recovered locally composed mostly of coarse ash, with rich amounts of Calcite and Wood pieces.[26] This composition indicates moderate explosivity on the genesis of the materials, relating the eruption products with short transport paths, as show little mechanical weathering, also corroborated by the thick layering and the low amount of basaltic bombs reported, while the correlation of wood and lapilli indicates a terrestrial deposition.[27] All together shows local Strombolian Volcanism, linked probably to a coeval rift, as recovered by the presence of more than 100 coeval volcanic necks in central Scania.[28] Dominating the Djupadal formation is moderately sorted lapilli tuff with abundant scoria, what indicates moderate explosivity, giving the eruption products short transport paths, preventing extensive mechanical weathering, that would create rounded fragments and large amounts of ash, that along thick layers and decimetre sized basaltic bombs are clear signs of closeness to the volcanic source.[27]

A more recent work has covered the only know Borehole that includes strata of the Central Skane Volcanic Province, the KBH2 (with a previous failed attempt, KBH1) at Djupadalsmölla, yielding more data about the stratotype location of the formation.[29] At this Borehole the Crystalline Basement is composed of invariably weathered red-whitish Gneiss, having a thickness of 6.32 m, with the lower part dominated by mafic minerals with coarse-grained quartz and feldspar. Facing the upper part, the gneiss becomes less clear, affected by weathering, becoming eroded and turning into white kaolinite.[30] The Jurassic formation itself is composed of shale, varying from light-dark gray in color, with a 30 cm level of silt/clay, with coal in the darker parts. The lower sandy shale part contains microfaults, being displaced towards the top by volcanic material, some fragments of this type being embedded in the shale in the uppermost part. The first section is composed of coarse silts and to a lesser extent sands in a clay matrix. It is followed by a level with finer grains without coarse elements, which evolves to a new level where the clay matrix returns, with high levels of Sulfur.[30] The properly volcanic strata have a thickness of 19.50 m, being marked by the appearance of lapilli tuff cemented by calcite or zeolite, with abundant xenoliths and accidental lithics of felsic type and a size of 10-20 cm. This level is subdivided into 4, starting from subunit 1, ranging from 25 to 48 cm, where it can be observed how the shale is replaced by volcanic material.[31] It is followed by subunit 2 of 48 to 150 cm, dominated by lapilli tuff whose initial point collects better preserved volcaniclastic grains and with a clear white cement filling the intergranular porosity and vesicles. About 80 cm, a nodule with intense white cement and almost black grains is produced, as well as the first large piece of carbonized wood is collected. Against the top of this sublevel appears a small interval of well-cemented sandstone.[32] Subunit 3 from 1.50 m to 14.75 m is a lapilli-tuff marked by a color change to a much darker texture and lower contrast with many small, lighter-colored pseudo-tuffs.[33] The cementum in the subunit is dark with the grains being lighter in color. The last subunit, 4, begins at +14.75 m where the darkening tone reaches an almost black hue in the interval of the first 2 meters, then returning to a greyish tone.[34]

On Karup the exposed layer is 1-1.5 m high and about 5 m wide, with an unclear distribution of sediments beyond quaternary sediments superimpose the pyroclastic material, while the substrate is not exposed.[35] This exposed layer represents one of the outcrop where the major abundance of charred pieces of wood and also silicified, non-charred pieces of wood, occur.[36]

Lilla Hagstad is composed by this is composed of Nepheline basalt with dark glass base in which crystal individuals of nepheline, Angite, olivine and magnetite are included. Also hosts lapillas from hazelnut to pea size are rounded, provided with a dense dark solidification zone and prove thereby as subaeric ejection products and not an origin from intrusive formations on cavities or open crevices. There are also quartz grains, often shattered, which become fragments cemented by the solidifying glass mass.[37]

On Koholma, a 0.5 m of green-brown trufaceous rocks, composed mostly by large clasts of crystalline rocks along lapilli and abundant plant remains, all identical to those seen on Djupadalsmölla, and also suggested to be derived form sliding flows from a nearby volcano.[38]

On Snälleröd (65 m thick, 44 m saprolite) samples taken, compared with the ones from the bottom of the road cut at Djupadal at 1.5 km NNE of this last one showed massive, soft lumps of a white, fine-grained material lacking any visually detectable grains of primary minerals, where only chemical data showed that this material is highly depleted in Calcium, Potassium and Sodium with significant kaolinization. The kaolinized rock has Kaolinite content exceeding 85%.[4] The samples taken at the NNE of Djudapal shows angular, gravel-sized material contains less altered granules of gneiss, remnant Feldspar and Quartz on a fraction dominated by Smectite, unlike the previous kaolinite-dominated ones, although this last one is also abundant.[4]

The Korsaröd Lagerstatten is located also on central Scania, and represents the best outcrop of the formation, leading to exquisitely preserved (with fossilized nuclei and chromosomes) specimens of ferns of the extant genus Osmundastrum.[39][40] This location was linked by Ulf Sivhed in 1984 with the Dajupadal Formation.[5] What was corroborated by recent studies.[41] It is also composed by volcaniclastic deposits, located at 380 m WNW of the nearest basaltic volcanic plug.[42] It is composed by mafic clasts agruped with agglomerates, oriented to this volcanic plug, coming probably from it or nearby ones.[42] Its clasts are angular and poorly sorted, recovered on a series of layers whose timing is uncertain, as there is no probe if represent discrete episodes separated by intervals of non-deposition or is result of variations due to a high-energy depositional setting.[42] Like in the Dajupadalsmölla type deposit, there is a great abundance of ash/mud content of the deposit filled with chaotic distributed wood fossil, what leds to the interpretation that this was a lahar deposit.[42] This location has been compared with modern Rotorua, New Zealand, considered an analogue for the type of environment represented in southern Sweden at this time.[43]

Eneskogen, Bonnarp and Säte volcanic necks are the main coeval of the Formation.[44] While Bonnarp (5–6 m height and covers roughly 5,000 square meters, covered by Jurassic sediments) is calculated to have at least 185.4+4.6 Ma (Middle Pliensbachian), Säte (Comprise two basalt pipes, each roughly 6–10 m high and some 10,000 square meters in area) yielded 180.0+0.7 Ma and Eneskogen (A large hill covered by quaternary sediments. Some few boulders and basalt pillars were exposed) 182.1+0.6 Ma, both Lower Toarcian in age.[45] It recovers one of the tree major Mesozoic volcanic events on Skåne.[46] Säte & Bonnarp volcanic strata is composed by Basanite, being the first Glassy Facies, and the second microcrystalline, while Eneskogen also microcrystalline, is dominated by Melanephelinite.[47] Bonarp has a very special character, which differed considerably from the normal cone of a Stratovolcano.[48] At Bonarp, between the crystalline basement and the basalt tuff, a sedimentary layer sequence intervenes, alternating sandy and clayey layers.[48] The layers linked to the volcano consists of sand and clay stones with thin coal seams, with bottom redistributed disturbed layers of kaolin.[48] The overall appearance of the Bonarp volcano suggests that local tectonic movements were more or less complete when the eruption began.[48] The pre-basaltic weathering process was also a deep kaolinization of the crystalline basement (Knutshög, Djupadal). At Bonarp the basement is also deeply decomposed.[49]

The Brandsberga and Kolleberga erratics represent a series of Upper Pliensbachian sandstones, linked with the Bonnarp Volcano and deposited on a marginal marine to coastal setting, with a faunal composition dominated by Bivalves, as well as Belemnnites and Bony Fish remains, yet the bivalves were the only identifiable to genus level.[50] This unit has traces of volcanoclastic elements on it, yet is dominated by sandstones, the same that are found in the associated Dinoflajellate-rich layers of the Bonnarp Volcano.[51] The unit is also the only part that recovers marine fauna, mostly molluscs.[52] The erratics suggest the presence of undiscovered solid rock in the immediate vicinity, and, like on the main volcanic layers, they're abundant on large angular and rounded pieces of half-weathered Archaean sediments, probably either or result of exposed nearby layers of this period or result of the basement erosion due to coeval volcanism.[53] The deposit was part of a regression trend where an embayment was formed in western central Skane, likely derived from the earlier Lower Pliensbachian major transgression. The presence of Kaolinite as well indeterminate Plant remains suggest the flow of terrestrial matter through rivers from the east.[54]

Lithology

The Basalts where originally classified by Eichstädt in 1882 on its own scheme, divided in groups such as Feldspar basalts, Nepheline basalts, Leucite basalts and Glass basalts.[55] The Basalt magmas must have been formed in the mantle and were hastily transported towards the surface during eruptions.[56] The local basanites contain spinel-bearing and spinel-free Peridotite xenoliths.[57] The ultramafic and mafic xenoliths document two types of petrogenetic origin: the group A, composed of Dunite, Harzburgite and Spllherzolite, whose origin is linked with the upper mantle; B, Pyroxenite, Gabbronorites, Anorthosite and mafic xenoliths that represent likely intrusive bodies, which might be located at the crust–mantle or upper crust.[58] Beyond the Basanites, the overall lithology includes Iddingsite, Augite, Serpentinite, Magnetite, Orthoclase, Plagioclase, Olivine, Chlorite, Apatite, Titanite, Feldspar and indeterminate Glass-like particles.[59] Mostly of the Volcanic necks host moderate to high quantities of olivine.[60]

Petrologically Borehole KBH2 contains volcaniclastic grains with angular to subrounded swelling clay minerals formed from degradation of the original ash and lapilli glass.[61] Subunits 2-3 show a clear volcaniclastic texture like a lapilli-tuff supported by clasts, calcite cement filling all the spaces between grains, together with Kaolinite clay. A first section is cemented only by iron-rich calcite, followed by a second with various types of cement, being Calcite, Chlorite and Zeolites, including some of the latter with Armotome. In between are patches that combine Fe-calcite with Ca-rich siderite.[61]

At Djupadalsmölla the layers are composed by a more or less transformed tuff, that is composed of volcanic ash, sand and lapilli, and small portions of completely or almost completely disintegrated basalt. The color of this rocks changes between blue-green, green-yellow, brown-yellow or brown, depending on the weathering stage in which it is located, with the greenish blue being the rocks in the healthiest state, while brownish color tones seem to indicate a more advanced decomposition.[62] The rocks have abundant clearly distinguished rounded grains that range from the size of a pea to the size of a hazelnut. This grains compose most of the rocks, cemented together and cohesive, mostly forming Limestone cement.[63] The cavities are generally completely filled with secondary products, mostly calcareous limestone, in addition to Zeolites and Viridite. The cement between the grains consists mainly of limnic limestone and sandstones.[64] This rocks were originally either a feldspar basalt or Limburgite ("glass" basalt), in which both kinds of Augit and olivine are always included, albeit in varying amounts.[65] In the Limburgites, feldspar, augit and olivine, syllic microliters (Ferrite) as well sparsely sprinkled grains of Magnetite have been reported in varying amounts.[66] The "Glass" sections appear to have been strongly grained in most cases, but these grains did not fully correspond to the globulitic grains, which are otherwise usually found in the glass of basalts, likely made of Olivine or Augite, as well small Serpentine particles.[63] The basalt tuff therefore came to form a porous glass, in which no other crystallized substances can be found than the olivine crystals already formed in the crater. The cement, which fills the space between the sideromelan grains and seals them together, consists for the most part of limestone, which is easily recognized under the microscope and is also clearly visible when the preparations are treated with acid. The cement is both sparse and forms a rich matrix for the sideromelan grains.[67] Other layers are composed with red Gneiss, which is similar to that found in the area, and a sandstone, which is similar to that found at Höör. This Gneiss is recovered in the Volcanic Bombs (lapilli) along Amphibolite.[68] These rocks are the most common around the volcanic sediments, with other layers composed of mica Diorite, limestone, clay slate and muddy, shale sandstone.[13] In this layers wood is common, included from irregular pieces to completely preserved branch pieces.[24]

Siliciclastic interbeds

The Siliclastic interbeds of the Djupadal Formation where originally assigned to the underliying fluvial Höör Sandstone, of Hettangian-Pliensbachian age. This interbeds outcrop in several concrete locations: Between Vitseröd and Hultarp areas, in south Hultarp and N. and S. Hultseröd, where sandstone blocks are so rare that there is no reason to assume solid sandstone,as is limited to 2 concrete blocks.[69] At Dagstorp the layers compose an arkose, Quartzite and pebbles, with partly very large blocks, which gave rise to local quarrying. The outcrops is 350 in NO-SW and 250 in perpendicular to it, located on the slope from the large basalt cup in the north and surrounded almost entirely by bedrock. The sandstones spread to the southern side of the Dagstorpssjön lake, while the eastern boundary consists of a gneiss slab with a steep precipice to the east, which is partly Kaolin-weathered, being overlaid in the west by a rich block field of sandstone. It in this location where the interbeds are best preserved, standing firm, sheltered of bedrock and basalt. The ichnofossil Monocraterion isp. and the bivalve Cardinia follini occur in the sandstone pointing to low oligohaline faunas, yet it cannot be precisely assigned to either the uppermost Höör layers (Vittseröd Member) or the lowermost Djupadal.[70] This faunal agruppation occurs together with abundant foliar remains and are equivalent to the Brandsberga erratics, which was closer to the mouth of the bay developed in the region, as it proves the find on its sandstone of more clear marine fauna.[71] Farthest to the south-east, in the innermost part of the original bay, in the area around Höör train station, plant remains in abundance where embedded in the sandstone, with only a few mussels found.[72] At 1 km west of Hallaröd there is a series of layers with blocks of quartzite gray, hard sandstone, followed 200 m N by a low ridge with crumbs, shards and blocks of weathered gneiss, probably the basis for now destroyed sandstones.[73] The nearby sandstones of the Djupadal deposits are covered in ripple marks and clearly indicative of fluvial or lagoonal deposition, as well along Dagstorp ones, the sandstones that best evidence being deposited at the same time the volcanic eruptions take place.[74]

Age and Correlations

Tralau (1973) measured the age for the local deposits, establishing that where typical Middle Late Triassic to Lias strata, with absent Middle Jurassic sediments, with the exception of the volcanic event ejecting the tuffs in Korsaröd, stating that they took place in the Middle Toarcian.[75] Radiometric ages obtained by using K–Ar techniques scatter in a wide range between 171 and 179 Ma.[76] Following works on the 80s and 90s recovered also this original datation, putting this and the Dajupadalsmölla outcrop on the Toarcian-Aalenian boundary, as example of latest lower Jurassic volcanism on the region. In 2006-2009 a depth study of the Volcanic Plugs led to stablish a range 191–178 Ma based on 40Ar/39Ar whole-rock ages for samples derived from eight basanite–nephelinite plugs, with included late Jurassic and Lower Cretaceous necks.[77][78] Palynological studies on the 2014 changed the perspective of the age of the location, proposing a more fitting Late Pliensbachian based on palynology.[79] In 2016 an in depth palynological study of the Korsaröd section led to stablish a Pliensbachian–early Toarcian(?) age, based on the high presence of the genus Perinopollenites elatoides (Pinales) and Eucommiidites troedsonii (Erdtmanithecales).[80] Yet other more recent works support that the outcrop is Late Toarcian in age, with a more recent work recovering a high-precision 40Ar/39Ar anorthoclase feldspar age of 176.7 ± 0.5 Ma (2-sigma), Late Toarcian in the Lilla Hagstad Volcano, and not supporting the 190–110 Ma age range, and establishing the region as short lived intraplate magmatism volcanic field.[81] This confirmed Tralau Original palynology results and the Paleomagnetic Studies done in 1993, that found 177-171 Ma as the most probable age.[82] A recent sample of the KBH2 at Djupadalsmölla has found again solid evidence of a Latest Pliensbachian age, namely by the ocurrence of scarce Callialasporites turbatus, the presence of the marine palynomorphs Mancodinium semitabulatum and Nannoceratopsis gracilis and the total absence of the genera Parvocysta and Phallocysta.[83]

The formation overlies the Höör Sandstone, and is time-equivalent with the Rydebäck and Katslösa members of the Rya Formation on NW Skane, the Röddinge Formation of the Vomb Trough and the Sorthat Formation of Denmark, with which it shares the abundance of Fern-derived material.[84] The formation also correlates with the Fjerritslev Formation of the Danish Basin, and the Gassum Formation of the Øresund Basin.[85]

The volcanic material was translated to the Continental margin by large fluvial channels, that ended on the sea deposits of the Green Series of Grimmen and Dobbertin, with the three-dimensional clay of this unit probably originated as the weathering product of this.[86] The Volcanic activity very likely eroded the underlying Hettangian-Sinemurian layers of the Höör Sandstone, deposited on the Fennoscandian coast as result of the weathering of the Precambrian-Paleozoic. This is seen as, after the increased amount of clays with abundant volcanic materials, sands were repeatedly poured into the North German Basin from Skåne, as result of the erosion of the Höör sandstone.[87]

Basin history

Geology of Sweden, showing the Precambrian-Paleozoic Basement and the Mesozoic limited to Skane
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.[88] The main volcanic phase of this volcanic province occurred in the Early Jurassic (Pliensbachian to Toarcian) at 184–176 Ma.[89] 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.[90] No correlative pyroclastic beds have yet been identified in sedimentary basins surrounding central Skåne.[91]

Toarcian

During deposition of the Rydebäck Member of the coeval Rya Formation, 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.[92]

Environment

Paleogeography of northwestern Europe during the Early Jurassic with Agaleus shark fossil finds, know from the (in part) coeval Rya Formation. Elevated land areas are shown in grey.

At Djupadalsmölla the presence of wood, together with the moderately sorted lapilli tuff, indicate a terrestrial depositional environment, probably influenced by freshwater deposits.[27] It has been suggested to be deposited on a fluvial setting that was influenced by a debris flow, mixing plants and sediments on a downhill transport, probably from the nearby Äskekull Volcano, one and a half kilometers south.[38] This was proven by the fact there are sandstone layers with Ripple marks in the western part of the locality close to the pyroclastics.[10] It has been suggested this sandstone underlies the volcanic sedimentary rock, yet it has abundant Silicon dioxide from the pyroclastics, implicating the transport of large amount of this last one early after deposition, precluding major sediment compaction. This process requires a coeval age relationship between the sandstone and the lapilli tuff in Djupadalsmölla.[28] The same category includes the lamellar deposits in the quartz grains of a dark sandstone, which Nathorst found as debris near Dagstorp Lake (Dagstorper sandstone), interpreted as a Microcline. Similar debris has also been found at Ikersberg near the Höör Train station.[93] At Lilla Hagstad tuff changes into a similar sandstone-like that is, as its geological occurrence shows, formed in connection with the deposition of a genuinely volcanic tuff, and is therefore itself a tuff with predominantly allothigenic quartz and feldspar grains. These sandstones were also formed as tuff sandstones that were deposited during the initial phase of eruptions.[94] The Dagstorp sandstone is filled with green stone fragments, considered basalt and Diabase, coming either from the vicinity of their current occurrence or further from the east, proving that tuff formations in connection with the basal eruptions originally had a not inconsiderable spread.[95] This sandstones where latter moved by rivers and deposited in both the Fennoscandian Border Zone and the Danish Basin.[96] Pyroclastic rain may have deposited material on slopes, with landslides to low areas as a result. Deposition in water normally creates more well-sorted deposits than those studied. However, zeolite has been enriched in Barium, which is often enriched in seawater, which probably circulated as hydrothermal flows in the lapilli tuft creating diagenetic changes, including deposition of Calcite and Zeolite.[97]

The Borehole KBH2 also at Djupadalsmölla presents a notorious weathering of the crystalline basement, an aspect that could not have occurred in the Late Triassic due to the local dry conditions, but rather took place at some point in the 25 Ma between the Rhaetian and the Late Pliensbachian.[98] Above the basement, the appearance of a dark colored shale level indicates an aquatic environment, which shows fluctuations due to weak currents due to changes in organic-rich clay/silt and fine-grained sand. This aquatic environment presents traces of dinoflajelates, which implies an environment with marine influence, an aspect contrasted by the presence of algae that are only found in calm to brackish lake environments. The presence of both groups tends to suggest a restricted area, such as a protected lagoon or bay. The composition of the vegetation based on palynology suggests a humid and moderately warm climate, combining palynomorphs from cold (Perinopollenites elatoides and Chasmatosporites spp., etc.) and warmer (Monosulcites spp. and Deltoidospora toralis, etc.) environments.[99] In general, it suggests a thick forest cover dominated by conifers, ginkgoes, and seed ferns, while the understory/ground had different kinds of ferns, all in low and humid lands with a somewhat limited topography where the Cupressaceae, providers of Perinopollenites elatoides, dominated. Unusually dark pollen grains can indicate a burning forest fire.[100] The arrival of Volcanism in the area is marked in the profile by the presence of volcanic clasts embedded in the upper part of the shale unit, indicating that it was not yet consolidated when they were deposited, confirming that they are of the same age and coinciding with volcanoes of the same age of Eneskogen and Jällabjär 1 and 3 km away. There is no evidence of repeated eruptions and charred wood abounds confirming transport and deposition of the material through forested areas. The local water body must have been quite small/shallow and have a low energy level, as sorting and rounding of transported grains is not observed. After the volcanic deposition, an early lithification is observed, with the presence of Ba-rich zeolites, evidence of a later marine invasion that affected diagenesis, possibly the transgression collected in the Brandsberga erratics.[101]

In the last 10.000 years, in the Chaîne des Puys (Massif Central region, Puy de Pariou and Puy Mercœur in the images) volcanoes where formed in a similar "sudden" way of the volcanic field that generated the layers recovered at Djupadalsmölla and Karup outcrops of the formation.[102] Over this landscape environments similar to actual Rotorua's Waimangu Volcanic Valley and Whakarewarewa Thermal Village where developed.[103]

In Korsaröd there where found frashwater algae that suggest also a river influence.[104] The data provided by the fossilized wood rings showed that the location of Korsaröd hosted a middle-latitude Mediterranean-type biome in the late Early Jurassic, with low rainfall. Superimposed on this climate were the effects of a local active Strombolian Volcanism and hydrothermal activity. This location has been compared with modern Rotorua, New Zealand, considered an analogue for the type of environment represented in southern Sweden at this time. The locality was populated mostly by Cupressaceae trees (including specimens up to 5 m in circunference), known thanks to the great abundance of the wood genus Protophyllocladoxylon and the high presence of the genus Perinopollenites elatoides (also Cupressaceae) followed by Eucommiidites troedsonii (Erdtmanithecales).[80] Volcanogenic deposits are dominated by cypress family pollen with an understorey component rich in putative Erdtmanithecales, both representing vegetation of disturbed habitats. The abundance of Protophyllocladoxylon sp. is also related with a sporadic intraseasonal and multi-year episodes of growth disruption, probably due to the volcanic action, with rapid permineralization of woody remains, suggesting that the vegetation grew in a hydrothermal region, with major challenges for roots to cope with warm, mineral-laden fluids percolating through the soil.[105] Pollen, spores, wood and charcoal locally indicate a complex forest community subject to episodic fires and other forms of disturbance in an active volcanic landscape under a moderately seasonal climate.[106] Osmundastrum pulchellum, the best preserved fossil identified in this unit (whose rhizome hosts epiphytes, micro-herbivores, parasites, saprotrophs and fine organic remains) were a prominent understorey element in this vegetation and were probably involved in various competitive interactions with neighboring plant species, such as Lycophytes, whose roots have been recovered inside the Rhizome.[106] The ferns where part of a fern and conifer rich vegetation occupying a topographic depression in the landscape (moist gully) that was engulfed by one or more lahar deposits.[106]

Sandåkra Lake System

The Sandåkra Lake System, which appeared before the emergence of the local volcanics but was influenced by in the upper sections, was likely born of an event where the hinterland flood a geological rift breach, washing sediments and volcanic materials into it from far sources, in a similar way to modern Lake Rotokakahi of New Zealand

The Jurassic layers at the north of the main volcanic outcrops include a unit known as the Sapropel of Sandåkra (south of Finjasjön). This unit is composed by a powerful layer of up to 150 m with sandstones, clays, Oil Shales, Breccias, etc., being clearly younger than the Höör Sandstone and resting directly on the Paleozoic bedrock.[107] This subunit is known mostly by boreholes, and was shown to be in part coeval with the volcanic eruptions of the end Lower Jurassic, as samples recovered from the main Sandåkra bore where identical in abundance of volcanic minerals and hosted the same type of palynomorphs seen in Djupadal and Korsaröd.[108] The borehole of Sandåkra includes a 70 m thick Shale/mudstone/Hialoclastite layer, indicating a water body of that depth was developed locally. This water body was interpreted as a freshwater lake on the basis of the absence of marine palynomorphs.[109] This lake likely developed either on a tectonic breach originated from the same rifting system that give rise to the local vulcanism or in a topographical depression, being either of the two options filled latter with sediments coming from several freshwater flows of different density from the inner hinterland, with the volcanic minerals, only present in the uppermost sections, coming from a source far from the shoreline at the south.[109] The enormous abundance of palynomorphs suggest the presence of static waters creating hydrodynamic traps, as well a stagnant lake system, like the Deer Island Lake of Michigan, allowing to a stagnation of the sediments and the development of anoxic conditions at the bottom, as proven by the Shale abundance.[110] Ephemeral streams feed the system, while the shores where mostly composed of sandstone paleosoils.[111] This unit developed in the lower part in a similar way to the Toarcian Sichuan Lake of the Ziliujing Formation, as well host similar to the shales of modern Kastoria Lake of Greece or the organic sapropel of the Sinove Lake of Ukraine. Towards the upper part, the evolution of the Sandåkra Lake was almost identical to the "Carapace Lake" of the Toarcian Mawson Formation, both heavily influenced by local volcanism, with either hydrothermal leaks or tuff-derived material washed by rivers and streams. An environment similar to modern Waimangu Volcanic Valley likely developed locally when the Djupadal Formation deposited.[112]

Fossils

Pseudofungi

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 Species Location Material Notes Images

Peronosporomycetes[106]

Peronosporomycetes Indeterminate

  • Korsaröd
  • Oogonia
  • Ellipsoidal spores
  • Reticulum?

A parasite/saprotroph Pseudofungal Protist, incertae sedis inside Peronosporomycetes. Was recovered from the petiole of the holotype of Osmundastrum puchellum and interpreted as a peronosporomycete with parasitic or saprotrophic relation with this part of the plant. If the identification of the oogonia of peronosporomycetes is correct, then this implies regularly moist conditions for the growth of Osmundastrum pulchellum and this is consistent with the general habitat preferences of extant Osmundastrum.[106]

Extant Phytophthora, a Pseudofungal protist with sparsely bullate ornament, like the ones found Osmundastrum pulchellum

Fungi

Genus Species Location Material Notes Images

Fungi[106]

Fungi Indeterminate

  • Korsaröd
  • Spores
  • Hyphae

A Fungus, incertae sedis inside Fungi. From the petiole and the root of Osmundastrum puchellum where recovered thread-like structures, identified as derived from a pathogenic or saprotrophic fungus invading necrotic tissues of the host plant. The fungus' interaction with the plant was probably mycorrhizal.[106]

Extant Microglossum, a saprotrophic fungus, whose Hypae are similar to the ones found Osmundastrum pulchellum

Acritarchs

Genus Species Location General ecology Abundance Material Notes Images

Leiofusa[113]

  • Leiofusa jurassica
  • Bonnarp Volcano

Brackish-marine

Rare

  • Acritarchs

An Acritarch, that can be both from Green/Red Algal origin.

Leiosphaera[113]

  • Leiosphaera sp. A (Gross)
  • Leiosphaera sp. B (Thin)
  • Bonnarp Volcano

Brackish-marine

Rare

  • Acritarchs

An Acritarch, that can be both from Green/Red Algal origin.

Micrhystridium[113]

  • Micrhystridium inconspicuum
  • Micrhystridium stellatum
  • Micrhystridium cf. fragile
  • Bonnarp Volcano

Brackish-marine

Rare to moderately common

  • Acritarchs

An Acritarch, that can be both from Green/Red Algal and Sphagnopsida origin.

Tetraporina[114]

  • Tetraporina tetragona
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Miospores

An Algae acritarch

Veryhachium[114]

  • Veryhachium spp.
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Miospores

An Algae acritarch

Dinoflajellates

Genus Species Location General ecology Abundance Material Notes Images

Cymatiosphaera[113]

  • Cymatiosphaera sp.
  • Bonnarp Volcano

Brackish-marine

Rare

  • Cysts

A Marine/Brackish Dinoflajellate, member of the family Pterospermopsidaceae inside Gonyaulacales.

Dapcodinium[114]

  • Dapcodinium priscum
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Common

  • Cysts

A Marine/Brackish Dinoflajellate, considered a primitive member of the order Peridiniales. A possible Fennoscandian Endemism

Mancodinium[114]

  • Mancodinium semitabulatum
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Common

  • Cysts

A Marine/Brackish Dinoflajellate, member of the family Mancodinioideae.

Maturodinium[114]

  • Maturodinium inornatum
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Cysts

A Marine/Brackish Dinoflajellate, considered a primitive member of the order Peridiniales.

Mendicodinium[114]

  • Mendicodinium groenlandicum
  • Mendicodinium reticulatum
  • Mendicodinium varigranosum
  • Mendicodinium vermiculatum
  • Mendicodinium sp. cf. morgenrothum
  • Mendicodinium spp.
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Common

  • Cysts

A Marine/Brackish Dinoflajellate, member of the order Gonyaulacales.

Nannoceratopsis[113][114]

  • Nannoceratopsis gracilis
  • Nannoceratopsis cf. pellucida
  • Nannoceratopsis spp.
  • KBH2 Borehole at Djupadalsmölla
  • Bonnarp Volcano

Brackish-marine

Common

  • Cysts

A Marine/Brackish Dinoflajellate, type member of the family Nannoceratopsiaceae.

Chlorophyta

Genus Species Location General ecology Abundance Material Notes Images

Botryococcus[115][104][114]

  • Botryococcus braunii
  • Botryococcus sp.
  • KBH2 Borehole at Djupadalsmölla
  • Sandåkra Borehole
  • Korsaröd

Freshwater

  • Almost dominant (+70%) in some samples in Sandåkra
  • Goes from a 2%, being the most abundant algae on some samples in Korsaröd to lack any presence on others
  • Miospores

A Freshwater Algae, type member of Botryococcaceae inside Chlorophyta

Cymatiosphaera[114]

  • Cymatiosphaera spp.
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Miospores

A Brackish/Marine Algae, member of Pterospermopsidaceae.

Lecaniella[104][114]

  • Lecaniella korsdoddensis
  • Lecaniella spp.
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Freshwater

A proportion of near a 3%

  • Miospores

A Freshwater Algae, member of Zygnemataceae inside Charophyceae. On some samples is the only recovered algae.

Leiosphaerida[114]

  • Leiosphaerida spp.
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Miospores

A Brackish/Marine Algae, member of Prasinophyceae.

Ovoidites[114]

  • Ovoidites spp.
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Miospores

A Brackish/Marine Algae, member of Zygnemataceae.

Pediastrum[104]

Pediastrum sp.

  • Korsaröd

Freshwater

Rare

  • Miospores

A Freshwater Algae, member of Hydrodictyaceae inside Chlorophyceae. The less abundant algae sampled locally.

Extant Pediastrum

Tasmanites[114]

  • Tasmanites spp.
  • KBH2 Borehole at Djupadalsmölla

Brackish-marine

Rare

  • Miospores

A Brackish/Marine Algae, member of Pyramimonadaceae.

Bryophyta

Genus Species Location General ecology Abundance Material Notes Images

Annulispora[116]

  • Annulispora folliculosa
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Sphagnopsida.

Polycingulatisporites[117]

Polycingulatisporites sp.

  • Korsaröd

Hygrophilous

Less than 1.1% of the total samples

  • Miospores

A Miospore, related with the family Notothyladaceae inside Anthocerotopsida. Very scarce

Rogalskaisporites[116]

  • Rogalskaisporites cicatricosus
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Sphagnopsida.

Stereisporites[79][117][116]

  • Stereisporites antiquasporis
  • Stereisporites aulosenensis
  • Stereisporites seebergensis
  • Stereisporites psilatus
  • Stereisporites spp.
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous

Less than 1.5% of the total samples

  • Miospores

A Miospore, member of Sphagnaceae inside Sphagnopsida. The major recovered Bryophyte spore

Extant Sphagnum, typical example of Sphagnaceae. Stereisporites probably come from a similar or a related Plant

Lycophyta

Genus Species Location General ecology Abundance Material Notes Images

Aequitriradites[118]

  • Aequitriradites salebrosaceus
  • Aequitriradites sp. nov.
  • Sandåkra Borehole

Hygrophilous

Very rare

  • Miospores

A Miospore, affinities with Selaginellaceae or Lycopodiaceae inside Lycopsida.

Aratrisporites[116]

  • Aratrisporites minimus
  • Aratrisporites spp.
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Lycopodiopsida.

Densoisporites[119]

Densoisporites crassus

  • Korsaröd

Hygrophilous

Rare

  • Miospores

A Miospore, affinities with Pleuromeiaceae, Selaginellaceae and Lycopodiaceae inside Lycopodiopsida.

Kraeuselisporites[116]

  • Kraeuselisporites reissingeri
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Lycopodiopsida.

Lycospora[116]

  • Lycospora salebrosacea
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Lycopodiopsida.

Neoraistrickia[117]

Neoraistrickia sp.

  • Korsaröd

Hygrophilous

Near a 3.4% on some samples

  • Miospores

A Miospore, affinities with Selaginellaceae or Lycopodiaceae inside Lycopsida. The most abundant Lycopsid spore recovered locally.

Extant Selaginella, typical example of Selaginellaceae. Neoraistrickia probably come from a similar or a related Plant

Retitriletes[117][116]

  • Retitriletes austroclavatidites
  • Retitriletes clavatoides
  • Retitriletes semimuris
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous

Less than 1%

  • Miospores

A Miospore, affinities with Lycopodiaceae inside Lycopsida. Diverse, but less abundant

Extant Austrolycopodium, typical example of Lycopodiaceae. Retitriletes probably come from a similar or a related Plant

Selaginellites?[120]

  • Selaginellites? sp.
  • Korsaröd

Hygrophilous

Rare, limited to the Roots of Osmundastrum pulchellum

  • Epiphytic lycopsid roots

A small herbaceous epiphytic lycopsid, incertae sedis inside Lycopsida. External exotic roots are preserved within detritus-filled cavities between the petiole bases of Osmundastrum pulchellum, with wall thickenings similar to the vasculature evident in ancient and modern herbaceous lycopsids.[121]

Equisetopsida

Genus Species Location General ecology Abundance Material Notes Images

Calamospora[117][116]

  • Calamospora tener
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous

A 4.3% on a single sample

  • Miospores

A Miospore, affinities with Equisetaceae inside Equisetales. Horsetails, herbaceous flora related to high humid environments, flooding tolerant plants.

Extant Equisetum cone, the typical example of the Equisetopsida. Calamospora spores are pretty similar to the extant ones

Filicopsida

Genus Species Location General ecology Abundance Material Notes Images

Acanthotriletes[118]

  • Acanthotriletes ovalis
  • Acanthotriletes trigonis
  • Acanthotriletes varius
  • Sandåkra Borehole

Reworked

Very scarce

  • Miospores

Affinities with the Botryopteridaceae inside Filicopsida. Reworked from primitive ferns found in Devonian and Carboniferous rocks of Europe

Apiculatasporites[122]

Apiculatasporites varians

  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A Miospore, affinities with Zygopteridaceae inside Filicopsida.

Auritulinasporites[123]

  • Auritulinasporites intrastriatus
  • Auritulinasporites scanicus
  • Auritulinasporites triclavis
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A Miospore, affinities with Matoniaceae inside Filicopsida.

Extant Matonia, typical example of the Matoniaceae. Auritulinasporites may have come from a similar genus

Baculatisporites[124][116]

  • Baculatisporites comaumensis
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous/Xerophilous?

Very scarce

  • Miospores

A Miospore, affinities with Osmundaceae or Hymenophyllaceae inside Filicopsida.

Extant Hymenophyllum, typical example of the Hymenophyllaceae. Baculatisporites may have come from a similar genus

Bracteolinasporites[125]

  • Bracteolinasporites clavatus
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A Miospore, affinities with Osmundaceae inside Filicopsida.

Cibotiumspora[117][126][116]

Cibotiumspora jurienensis

  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous/Xerophilous?

Present on various samples with ratios from 1-1.5%

  • Miospores

A Miospore, related with Cyatheaceae and Dicksoniaceae inside Filicopsida. Recovered on the petiole and the root of Osmundastrum puchellum.[126]

Conbaculatisporites[117][116]

Conbaculatisporites mesozoicus

  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous?

Scarce, with less than 2.3% on the samples

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Concavisporites[127][113]

  • Concavisporites crassexinius
  • Concavisporites sp.
  • Bonnarp Volcano
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A Miospore, related with Cibotiaceae, Gleicheniaceae, Matoniaceae and Dipteridaceae inside Filicopsida

Contignisporites[128]

Contignisporites problematicus

  • Korsaröd

Hygrophilous/Xerophilous?

Rare

  • Miospores

A Miospore, related with Cibotiaceae inside Filicopsida.

Extant Cibotium, typical example of the Cibotiaceae. Contignisporites may have come from a similar genus

Corrugatisporites[129]

  • Corrugatisporites klukiformis
  • Corrugatisporites scanicus
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A Miospore, related to Schizaeaceae inside Filicopsida. Miospores of the fern Klukia exilis

Cosmosporites[130]

  • Cosmosporites elegans
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Converrucosisporites[116]

  • Converrucosisporites spp.
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A Miospore, affinities with Osmundaceae inside Filicopsida.

Cyathidites[117][126][113][116]

  • Cyathidites australis
  • Cyathidites minor
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Bonnarp Volcano

Hygrophilous/Xerophilous?

It is the most common independent palynologycal residue recovered on Korsaröd, with Cyathidites spp. occupying a 21%

  • Miospores

A Miospore, related with Cyatheaceae or Adiantaceae inside Filicopsida. Recovered on the petiole and the root of Osmundastrum puchellum.[126] Cyathidites minor almost certainly belong to well known Mesozoic species Coniopteris hymenophylloides and to other fossil cyatheaceous or dicksoniaceous ferns such as Eboracia lobifolia and Dicksonia mariopteri.

Extant Cyathea, typical example of the Cyatheaceae. Cyathidites may have come from a similar genus

Cyclinasporites[131]

  • Cyclinasporites minutus
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Deltoidospora[117][126][116]

Deltoidospora toralis

  • Deltoidospora australis
  • Deltoidospora minor
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous/Xerophilous?

Moderately common

  • Miospores

A Miospore, related with Cyatheaceae, Dicksoniaceae, Gleicheniaceae and Schizaeaceae inside Filicopsida. Recovered on the petiole and the root of Osmundastrum puchellum.[126]

Gleicheniidites[113][116]

  • Gleicheniidites conspiciendus
  • Gleicheniidites senonicus
  • Gleicheniidites sp.
  • KBH2 Borehole at Djupadalsmölla
  • Bonnarp Volcano

Hygrophilous/Xerophilous?

Very scarce

  • Miospores

A Miospore, related with Gleicheniaceae inside Filicopsida.

Laevigatosporites[116]

  • Laevigatosporites spp
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A Miospore, affinities with Marattiales inside Filicopsida.

Lycopodiacidites[132][116]

Lycopodiacidites rugulatus

  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous?

Very scarce

  • Miospores

A Miospore, affinities with Ophioglossaceae inside Filicopsida.

Extant Ophioglossum, typical example of the Ophioglossaceae. Lycopodiacidites may have come from a similar genus

Marattisporites[79][117][133][116]

Marattisporites scabratus

  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole

Hygrophilous/Xerophilous?

3-14% on the samples

  • Miospores

A Miospore, affinities with Marattiaceae inside Filicopsida. The second most abundant spore recovered on the location

Extant Marattia, typical example of the Marattiaceae. Camptotriletes may have come from a similar genus

Matonisporites[116]

  • Matonisporites phlebopteroides
  • Matonisporites spp.
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A Miospore, affinities with Matoniaceae inside Filicopsida.

Osmundacidites[79][117][113][116]

Osmundacidites wellmanii

  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Bonnarp Volcano

Hygrophilous/Xerophilous?

Abundant, 10% on some samples

  • Miospores

A Miospore, affinities with Osmundaceae inside Filicopsida.

Extant Osmunda, typical example of the Osmundaceae. Osmundacidites may have come from a similar genus

Osmundastrum[134][135]

  • Korsaröd

Hygrophilous/Xerophilous?

Limited to a single Rhizome

  • Rizhome
  • Xylem cylinder
  • Leaf traces
  • Roots
  • Nuclei & Chromosomes
  • Subcellular Detail
  • Biotic Interactions
  • Preserved Ongoing Mitosis

A small (50 cm tall) Royal Fern, member of Osmundaceae inside Filicopsida. The most known fossil of the location, thanks to its exceptional fossilized Rhizome, that has preserved Nuclei and Chromosomes, a fine subcellular detail has rarely been documented in fossils.[136] Its Rooted in DNA content was used to extrapolate relative genome, finding relationships with extant Osmundastrum cinnamomeum, and confirmed a monophyletic Osmunda.[137]

Punctatasporites[138][116]

  • Punctatasporites flavus
  • Punctatisporites globosus
  • KBH2 Borehole at Djupadalsmölla
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Skarbysporites[116]

  • Skarbysporites crassexina
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Simplicesporites[139]

  • Simplicesporites granulosus
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Striatella[79][117][116]

Striatella seebergensis

  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous?

Very scarce, with less than 1.5% on all the samples

  • Miospores

A Miospore, affinities with Polypodiaceae inside Filicopsida.

Tigrisporites[116]

  • Tigrisporites scurrandus
  • KBH2 Borehole at Djupadalsmölla

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Trachysporites[133][116]

  • Trachysporites asper
  • Trachysporites fuscus
  • Trachysporites microclavatus
  • Trachysporites punctulosus
  • Trachysporites sparsus
  • Trachysporites tuberosus
  • Trachysporites spp.
  • KBH2 Borehole at Djupadalsmölla
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Todisporites[132][104]

  • Todisporites major
  • Todisporites minor
  • Korsaröd

Hygrophilous/Xerophilous?

Very scarce, with less than 1–3.1% on all the samples

  • Miospores

A Miospore, affinities with Osmundaceae inside Filicopsida.

Extant Osmunda, typical example of the Osmundaceae. Todisporites may have come from a similar genus

Undulatisporites[140]

  • Undulatisporites arquatus
  • Sandåkra Borehole

Hygrophilous

Very scarce

  • Miospores

A miospore, incertae sedis inside Filicopsida.

Zebrasporites[141][116]

  • Zebrasporites interscriptus
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Hygrophilous/Xerophilous?

Rare

  • Miospores

A Miospore, related with Cyatheaceae inside Filicopsida. Arboreal Fern Miospores.

Extant Cyathea, typical example of the Cyatheaceae. Zebrasporites may have come from a similar genus

Peltaspermales

Genus Species Location General ecology Abundance Material Notes Images

Alisporites[142][143][104][144]

  • Alisporites grandis
  • Alisporites thomasii
  • Alisporites robustus
  • Alisporites microsaccus
  • KBH2 Borehole at Djupadalsmölla
  • Sandåkra Borehole
  • Korsaröd

Xerophilous

A. grandis reaches near a 10% on some samples

  • Pollen

A Pollen Grain, affinities with Umkomasiaceae, Peltaspermaceae, Corystospermaceae and Caytoniaceae inside Pteridospermae.

Chordasporites[144]

  • Chordasporites spp.
  • KBH2 Borehole at Djupadalsmölla

Xerophilous

Very scarce

  • Pollen

A Pollen Grain, affinities with Corystospermaceae inside Pteridospermae.

Caytonipollenites[145]

  • Caytonipollenites sp.
  • Sandåkra Borehole

Xerophilous

Very scarce

  • Pollen

A Pollen Grain, affinities with Caytoniaceae inside Pteridospermae.

Crassipollenites[146]

  • Crassipollenites classopolloides
  • Crassipollenites diffusus
  • Crassipollenites rugosus
  • Sandåkra Borehole

Xerophilous

Very scarce

  • Pollen

A Pollen Grain, affinities with Caytoniaceae inside Pteridospermae.

Ovalipollis[113]

  • Ovalipollis sp.
  • Bonnarp Volcano

Xerophilous

Abundant

  • Pollen

A Pollen Grain, affinities with Caytoniaceae inside Pteridospermae.

Pityopollenites[146]

  • Pityopollenites pallidus
  • Sandåkra Borehole

Xerophilous

Relatively common

  • Pollen

A Pollen Grain, affinities with Umkomasiaceae, Peltaspermaceae, Corystospermaceae and Caytoniaceae inside Pteridospermae.

Protopinus[147]

  • Protopinus scanicus
  • Sandåkra Borehole

Xerophilous

Very scarce

  • Pollen

A Pollen Grain, affinities with Caytoniaceae inside Pteridospermae.

Schismatosporites[148]

  • Schismatosporites ovalis
  • Sandåkra Borehole

Xerophilous

Relatively common

  • Pollen

A Pollen Grain, affinities with Umkomasiaceae, Peltaspermaceae, Corystospermaceae and Caytoniaceae inside Pteridospermae.

Vitreisporites[79][117][149][144]

  • Vitreisporites bjuvensis
  • Vitreisporites pallidus
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole

Xerophilous

Scarce, with around a 1% on all samples

  • Pollen

A Pollen Grain, afinnities with Caytoniales inside Gymnospermopsida.

Erdtmanithecales

Genus Species Location General ecology Abundance Material Notes Images

Eucommiidites[79][104][113][150][116]

  • Eucommiidites troedssonii
  • Eucommiidites minor
  • Eucommiidites sp.
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole
  • Bonnarp Volcano

Xerophilous

Abundant, with around a 3-10% on all samples.

  • Pollen

A Pollen Grain, afinnities with Erdtmanithecales inside Spermatophytes. The Gymnosperms that produced Eucommiidites troedsonii pollen possibly dominated the understorey vegetation.

Gnetales

Genus Species Location General ecology Abundance Material Notes Images

Equisetosporites[138]

  • Equisetosporites hallei
  • Sandåkra Borehole

Mediterranean setting indicator

Intervals where is rare to moderately common on the older layers to almost dominant Pollen in some sections up.

  • Pollen

A Pollen Grain, affinities with Ephedraceae inside Gnetopsida. This Pollen grain was thought to be from Equisetaleans, yet was found in Ephedra chinleana

Extant Ephedra, typical example of Ephedraceae. Equisetosporites probably come from a similar or a related Plant

Cycadophyta

Genus Species Location General ecology Abundance Material Notes Images

Bennettitaceaeacuminella[113]

  • Bennettitaceaeacuminella sp.
  • Bonnarp Volcano

Xerophilous

Abundant

  • Pollen

A Pollen Grain, affinities with Cycadeoidaceae and Williamsoniaceae inside Bennettitales. A Genus used to classify Bennetittalean grains of uncertain provenance

Chasmatosporites[79][104][151][144]

  • Chasmatosporites apertus
  • Chasmatosporites elegans
  • Chasmatosporites hians
  • Chasmatosporites crassus
  • Chasmatosporites flavus
  • Chasmatosporites latus
  • Chasmatosporites magnolioides
  • Chasmatosporites major
  • Chasmatosporites minor
  • Chasmatosporites radiatus
  • Chasmatosporites rimatus
  • Chasmatosporites scaber
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole

Xerophilous

Rare, with around a 0.7-1% on all samples. Abundant at Sandåkra

  • Pollen

A pollen grain, incertae sedis inside Cycadopsida, Corystospermaceae and Araucariaceae. 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 maybe come from a related plant

Clavatipollenites[116]

  • Clavatipollenites hughesii
  • KBH2 Borehole at Djupadalsmölla

Xerophilous

Very rare, with less than 0.5% on all samples

  • Pollen

A pollen grain, incertae sedis inside Cycadopsida, Ginkgoaceae, Bennettitales, Axelrodiales and Gnetales.

Cycadidae[152]

  • Cycadidae Indet.
  • Djupadalsmölla

Xerophilous

Limited to a few fragments

  • Coalified Fragments

A Cycad, afinnities with Cycadidae inside Cycadopsida.

Entylissa[131]

  • Entylissa pyriformis
  • Entylissa reticulata
  • Entylissa tecta
  • Entylissa tenuis
  • Sandåkra Borehole

Xerophilous

Very rare

  • Pollen

A pollen grain, incertae sedis inside Cycadopsida. It also can be pollen from Ginkgoaceae.

Extant Encephalartos, typical example of the Cycas. Monosulcites Pollen is pretty similar to the extant ones of this genus

Monocolpopollenites[140]

  • Monocolpopollenites acerrimus
  • Monocolpopollenites cotidianus
  • Monocolpopollenites fusiformis
  • Monocolpopollenites magnus
  • Monocolpopollenites ovalis
  • Sandåkra Borehole

Xerophilous

Very rare

  • Pollen

A pollen grain, incertae sedis inside Cycadopsida. It also can be pollen from Ginkgoaceae.

Monosulcites[79][104][147][144]

  • Monosulcites punctatus
  • Monosulcites magnolioides
  • Monosulcites echinatus
  • Monosulcites spp.
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole

Xerophilous

Very rare, with less than 0.5% on all samples

  • Pollen

A pollen grain, incertae sedis inside Cycadopsida. It also can be pollen from Ginkgoaceae.

Ptilophyllum[153]

  • Ptilophyllum sp.
  • Korsaröd

Xerophilous

Limited to a single Leaf Impression

  • Leaf Impression

A Bennetite, afinnities with Williamsoniaceae inside Bennettitales. This single impression of a bennettitalean leaf fragment found in a fine ash layer constituted the only foliar remains identified within the volcaniclastic deposit of Korsaröd

Example of Ptilophyllum leaf

Ginkgophyta

Genus Species Location General ecology Abundance Material Notes Images

Ginkgoites[104]

  • Ginkgoites nitidus
  • Korsaröd

Xerophilous

Very rare, with less than 1% on all samples

  • Pollen

A Pollen Grain, affinities with Ginkgoales inside Ginkgophyta. Ginkgoales trend to spike towards the Toarcian in the Northern European region, as seen in the coeval Sorthat Formation of Bornholm

Extant Ginkgo, only surviving example of the Ginkgoaceae. Ginkgoites Pollen is pretty similar to the extant ones of this genus

Coniferophyta

Genus Species Location General ecology Abundance Material Notes Images

Araucariacites[104][144]

  • Araucariacites australis
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Xerophilous

Rare, with around a 1.5% on all samples

  • Pollen

A Pollen Grain, affinites with the family Araucariaceae inside Pinales.

Callialasporites[144]

  • Callialasporites turbatus
  • Callialasporites sp.
  • KBH2 Borehole at Djupadalsmölla

Xerophilous

Rare, with around a 1.5% on all samples

  • Pollen

A Pollen Grain, affinites with the family Araucariaceae inside Pinales.

Cedripites[104]

  • Cedripites sp.
  • Korsaröd

Xerophilous

Rare, with around a 1.5% on all samples

  • Pollen

A Pollen Grain, affinities with Pinaceae inside Coniferophyta.

Extant Cedrus Cone, example of the Pinidae. Cedripites is similar to the pollen found on this genus

Cerebropollenites[104][113][145][144]

  • Cerebropollenites mesozoicus
  • Cerebropollenites macroverrucosus
  • Cerebropollenites thiergartii
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole
  • Bonnarp Volcano

Xerophilous

Around 1-1.5% on all the samples

  • Pollen

A Pollen Grain, affinities with both Sciadopityaceae and Miroviaceae inside Pinopsida. This Pollen resemblance with extant Sciadopitys suggest that Miroviaceae can be an extinct lineage of sciadopityaceaous-like plants

Extant Sciadopitys. Cerebropollenites likely come from a related plant

Classopollis[79][104][113][154][144]

  • Classopollis classoides
  • Classopollis meyerianus
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd
  • Sandåkra Borehole
  • Bonnarp Volcano

Extreme Dry Indicator

Moderately abundant, with around a 1.5-9% on all samples

  • Pollen

A Pollen Grain, affinities with Cheirolepidiaceae inside Coniferophyta.

Exesipollenites[144]

  • Exesipollenites tumulus
  • KBH2 Borehole at Djupadalsmölla

Xerophilous

Moderately abundant, with around a 1.5-9% on all samples

  • Pollen

A Pollen Grain, affinities with Cupressaceae inside Coniferophyta. Lowland (coastal) indicator

Granulonapites[139]

  • Granulonapites baculatus
  • Granulonapites punctatus
  • Sandåkra Borehole

Xerophilous

Rare to moderately common in upper samples

  • Pollen

A Pollen Grain, affinites with the family Araucariaceae inside Pinales.

Inaperturopollenites[155]

  • Inaperturopollenites flavus
  • Inaperturopollenites globulus
  • Inaperturopollenites irregularis
  • Inaperturopollenites orbiculatus
  • Inaperturopollenites sublevis
  • Inaperturopollenites triangularis
  • Sandåkra Borehole

Xerophilous

Diverse and moderately abundant

  • Pollen

A Pollen Grain, affinities with Taxodiaceae and Cupressaceae inside Coniferophyta. Its abundance at Sandåkra Borehole can indicate the presence of a Taxodium Swamp-like habitat

Parvisaccites[156]

  • Parvisaccites enigmatus
  • Korsaröd

Xerophilous

Rare to moderately abundant

  • Pollen

A Pollen Grain, affinities with Podocarpaceae and Cupressaceae inside Coniferophyta.

Extant Podocarpus Cone, example of the Podocarpaceae. Parvisaccites is similar to the pollen found on this genus

Perinopollenites[79][104][144]

  • Perinopollenites elatoides
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Xerophilous

The most abundant pollen recovered locally, with near a 30% on some samples

  • Pollen

A Pollen Grain, affinities with Taxodiaceae and Cupressaceae inside Coniferophyta. Afilited with extant genera such as Thuja plicata

Extant Thuja Cone, example of the Cupressaceae. Perinopollenites is similar to the pollen found on this genus

Piceoideae[157]

  • Piceoideae Indeterminate
  • Djupadalsmölla

Xerophilous

Rare, limited to 2 specimens

  • Coalified Fragments

A Conifer, afinnities with Piceoideae inside Pinaceae. One of the branched leaves found wears ano organic structure that resembles extant Spruces. Likely belong to the common Picea-like foliage of Pityocladus

Pinoideae[158]

  • Pinoideae Indeterminate A
  • Pinoideae Indeterminate B
  • Djupadalsmölla

Xerophilous

Abundant, up to 20 especimens

  • Coalified Fragments

A Conifer, afinnities with Pinoideae inside Pinaceae. In the initial revision of Djupadalsmölla Nathorst determined that some of the plant remains come from conifers. Latter revision showed that three clearly distinct species are represented and two resemble the genus Pinus. Likely belong to Schizolepidopsis, as is the most common Pinaceae in the Eurasian region in the Jurassic

Pinuspollenites[104][144]

  • Pinuspollenites minimus
  • Pinuspollenites pinoides
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Xerophilous

Around a 1–4.5% on the samples.

  • Pollen

A Pollen Grain, affinities with Pinaceae inside Coniferophyta.

Example of extant Pinus, Pinuspollenites laves are considered related with it

Protocedroxylon[159]

  • Protocedroxylon lindleyanum
  • Protocedroxylon spp.
  • Korsaröd
  • Karup
  • Färingtofta
  • Snalleröd

Xerophilous

Moderately Common

  • Carbonised pieces
  • Isolated Logs
  • Isolated Branches
  • Stump remains

A Conifer, affinities with Pinaceae (Probably Abietoideae) inside Coniferophyta. Referred originally to the genus Cedroxylon, using it as referente to stablish an Eocene age for the Volcanic deposits, as similar wood from Eocene age was recovered from Jutland.[159] This genus has been found to not be suitable to Mesozoic woods, and material similar to the one recovered on the volcanic deposits has been assigned to Tiloxylon (=Protocedroxylon), also known in the Toarcian of Greenland.[160]

Protopiceoxylon[161]

  • Protopiceoxylon spp.
  • Djupadalsmölla

Xerophilous

Moderately Common

  • Isolated Branches
  • Stump remains

A Conifer, affinities with Piceoideae inside Coniferophyta. Various Pinus-like woods are identified in Djupadalsmölla, they indicate that the lava flows flowed from a nearby forest setting.

Protophyllocladoxylon[162]

  • Protophyllocladoxylon spp.
  • Korsaröd
  • Djupadal
  • Karup
  • Färingtofta
  • Snalleröd
  • Sjöberga
  • Säte
  • Knutshög
  • Bonnarp

Xerophilous

Dominant Wood Type, up to 50 specimens

  • Carbonised pieces
  • Isolated fragments
  • Chaotic aggregations in a weathered volcaniclastic agglomerate
  • Large logs

A Conifer, affinities with Cupressaceae inside Coniferophyta. The main diagnostic wood recovered locally, identified based on the possession of uniseriate rays with smooth walls, pointed oblique oopores, absence of axial parenchyma, and tracheid radial walls. It resembles the extant Thuja plicata, but hosts a mean rings more similar to Juniperus thurifera. A few fossil wood specimens clearly represent portions of large trunks, with at least one fragment derived from a trunk of 1.68 m in diameter (with stimated c.5.3 m). Although, most specimens represent lateral branches or even roots of small size (10 cm long and 5 cm wide).[163] Its growth rings are distinct in all wood samples.[164]

Permineralized Protophyllocladoxylon wood with a branch trace recovered from the volcaniclastic sediments at Korsaröd

Podocarpidites[144]

  • Podocarpidites spp.
  • KBH2 Borehole at Djupadalsmölla

Xerophilous

Around a 1–3.9% on all the samples

  • Pollen

A Pollen Grain, affinities with Podocarpaceae inside Coniferophyta.

Phyllocladiidites[144]

  • Phyllocladiidites spp.
  • KBH2 Borehole at Djupadalsmölla

Xerophilous

Around a 0.2% on all the samples

  • Pollen

A Pollen Grain, Incertade sedis affinities inside Coniferophyta

Quadraeculina[104][144]

  • Quadraeculina anaellaeformis
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Xerophilous

Around a 1–3.9% on all the samples

  • Pollen

A Pollen Grain, affinities with Podocarpaceae and Pinaceae inside Coniferophyta.

Rugaletes[104]

  • Rugaletes sp.
  • Korsaröd

Xerophilous

Around a 7.5% on all the samples

  • Pollen

A pollen grain, incertae sedis inside Coniferophyta.

Spheripollenites[104][116]

  • Spheripollenites psilatus
  • Spheripollenites subgranulatus
  • Spheripollenites spp.
  • KBH2 Borehole at Djupadalsmölla
  • Korsaröd

Extreme Dry Indicator

Scarce, around a 1.5–3.9% on all the samples

  • Pollen

A Pollen Grain, affinities with Cheirolepidiaceae inside Coniferophyta.

Sulcatisporites[155]

  • Sulcatisporites pinoides
  • Sulcatisporites quadratus
  • Sandåkra Borehole

Xerophilous

+1% on the samples

  • Pollen

A Pollen Grain, affinities with Pinaceae inside Coniferophyta.

Taedaepollenites[165]

  • Taedaepollenites scaurus
  • Taeniaesporites kraeuseli
  • Sandåkra Borehole

Xerophilous

+1% on the samples

  • Pollen

A Pollen Grain, affinities with Pinaceae inside Coniferophyta.

Arachnida

Genus Species Location Material Notes Images

Oribatida[126]

Oribatida Indeterminate

  • Korsaröd
  • Excavated areas filled with coprolites

A mite, incertae sedis inside Oribatida. On the petiole of Osmundastrum puchellum excavations up to 715 μm in diameter are evident, filled with pellets that resemble the coprolites of Oribatid mites found also on Paleozoic and Mesozoic Woods.[126]

example of Oribatida mite

See also

References

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  108. Tralau (1973) p. 151
  109. Nilsson (1958) p. 5
  110. Nilsson (1958) p. 6
  111. Nilsson (1958) p. 10
  112. Tralau (1973) p. 154
  113. Bölau & Kockel-Brosius (1965) p. 55
  114. Wahlquist (2023) p.22
  115. Nilsson (1958) p. 33
  116. Wahlquist (2023), p23
  117. Vajda, Linderson & McLoughlin (2016) p. 132
  118. Nilsson (1958) p. 29
  119. Tralau (1973) p. 130
  120. McLoughlin & Bomfleur (2016) p. 88
  121. McLoughlin & Bomfleur (2016) p. 89
  122. Nilsson (1958) p. 31
  123. Nilsson (1958) p. 32
  124. Tralau (1973) p. 132
  125. Nilsson (1958) p. 34
  126. McLoughlin & Bomfleur (2016) p. 91
  127. Nilsson (1958) p. 38
  128. Tralau (1973) p. 134
  129. Nilsson (1958) p. 40
  130. Nilsson (1958) p. 41
  131. Nilsson (1958) p. 43
  132. Tralau (1973) p. 131
  133. Nilsson (1958) p. 50
  134. Bomfleur, Grimm & McLoughlin (2014) p. 5
  135. Bomfleur, Grimm & McLoughlin (2015) p. 6
  136. Bomfleur, Grimm & McLoughlin (2015) p. 8
  137. Bomfleur, McLoughlin & Vajda (2014) p. 1378
  138. Nilsson (1958) p. 45
  139. Nilsson (1958) p. 47
  140. Nilsson (1958) p. 51
  141. Tralau (1973) p. 133
  142. Nilsson (1958) p. 30
  143. Tralau (1973) p. 135
  144. Wahlquist (2023) p.21
  145. Nilsson (1958) p. 35
  146. Nilsson (1958) p. 42
  147. Nilsson (1958) p. 44
  148. Nilsson (1958) p. 46
  149. Nilsson (1958) p. 52
  150. Nilsson (1958) p. 46
  151. Nilsson (1958) p. 36
  152. Tullberg & Nathorst (1880) p. 232
  153. McLoughlin & Bomfleur (2016) p. 87
  154. Nilsson (1958) p. 37
  155. Nilsson (1958) p. 48
  156. Tralau (1973) p. 136
  157. Eichstädt (1883) p. 115
  158. Eichstädt (1883) p. 114
  159. Tralau (1973) p. 121
  160. Philippe & Bamford (2008) p. 193
  161. Eichstädt (1883) p. 415
  162. Vajda, Linderson & McLoughlin (2016) p. 138
  163. Vajda, Linderson & McLoughlin (2016) p. 136
  164. Vajda, Linderson & McLoughlin (2016) p. 137
  165. Nilsson (1958) p. 49

Bibliography

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