Paleoecology and sedimentology of the Upper Cretaceous (Campanian), marine strata at Åsen, Kristianstad Basin, Southern Sweden, Scania

Faisal Javed Iqbal Dissertations in Geology at Lund University, Master’s thesis, no 373 (45 hp/ECTS credits)

Department of Geology Lund University 2013

Paleoecology and sedimentology of Upper Cretace- ous (Campanian), marine strata at Åsen, Kristianstad Basin, southern Sweden.

Master’s thesis Faisal Javed Iqbal

Department of Geology Lund University 2013

Contents

1 Introduction…...... 6 2 Background...... 6 2.1 The Palegeography of the Cretaceous World...... 6 2.1.1 Plate Tectonics and Volcanism...... 6 2.1.2 Climate and Sea level changes...... 8 2.2 Fauna of the Late Cretaceous...... 8 2.2.1 Terrestrial Fauna...... 8 2.2.2 Terrestrial Flora...... 9 2.2.3 Marine Fauna…...... 9 2.3 North America- Western Interior Seaway during the Campanian ...... 9 2.4 Europe during the Campanian ...... 10 2.4.1 Campanian marine fossil assemblages in the Kristianstad Basin - Vertebrate Fauna...... 10 2.4.1.1 Sharks...... 10 2.4.1.2 Mosasaurs...... 12 2.4.1.3 Plesiosaurs...... 12 2.4.1.4 Turtles...... 12 2.4.1.5 Rays...... 12 2.4.1.6 Dinosaurs...... 13 2.4.1.7 Coprolites...... 13 2.4.2 Campanian marine fossil assemblages in the Kristianstad Basin - Invertebrate Fauna ...... 13 2.4.2.1 Belemnites...... 13 2.4.2.2 Oysters...... 14 2.4.2.3 Brachiopods (ex crania)...... 14 2.4.2.4 Inoceramids...... 14 2.4.3 Campanian Flora in the Kristianstad Basin - Fossil wood (Charcoal)…………...... 14 3 Geological Setting...... 16 4 Material and Methods...... 16 4.1 XRF analysis of the sediment samples...... 17 4.2 XRF analysis of the belemnites...... 17 4.3 SEM charcoal analysis...... 17 5 Results...... 18 5.1 Stratigraphy...... 18 5.2 Results of the XRF analysis (sediments)...... 21 5.3 Results of the XRF analysis (belemnites)...... 21 5.4 Results of the SEM (charcoal)...... 23 5.5 Results of the fossils………...... 23 6 Discussion...... 23 6.1 Fossil fauna ...... 23 6.2 XRF (sediments) ...... 28 6.3 XRF (belemnites) ...... 28 6.4 Comparison between the B. mammillatus and B. balsvikensis zones at Åsen ...... 29 6.4.1 Fossil fauna...... 29 6.4.2 Sediment content...... 30 6.4.3 Paleoecology...... 30

Cover Picture: Reconstruction of marine vertebrates in the Kristianstad Basin during the late early Campanian. In the centre is the mosasaur Tylosaurus with a small polycotylid plesiosaur just to the bottom left of it, the latter chasing belemnites and pachydiscid ammonites. In the upper left corner is the aquatic bird Hespe- rornis diving for bony fish, and to the right of it is the plesiosaur Scanisaurus. Two sharks and a marine turtle occupy the right of the picture (illustration by Stefan Sølberg in Sorensen et al. 2013).

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7 Results and discussion about the Charcoal...... 31 7.1 SEM charcoal images (Fig. 13, 14 &15; Table. 4)...... 31 7.2 Binocular charcoal images (Fig. 16, 17 & 18; Table. 5)...... 33 7.3 Result and discussion about the flower/sedges/equisetum (Fig 11) ...... 35 8 Discussion about the eggs/fecal pellets/small coprolites (Fig 12) ...... 35 9 Conclusions...... 35 10 Acknowledgement...... 36 11 References...... … 38 12 Appendix...... 50

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Paleoecology and sedimentology of Upper Cretaceous (Campanian), marine strata at Åsen, Kristianstad Basin, southern Sweden

FAISAL JAVED IQBAL

Iqbal, F. J., 2013: Paleoecology and sedimentology of the Upper Cretaceous (Campanian), marine strata at Åsen, Kristianstad Basin, Southern Sweden, Scania. Dissertations in Geology at Lund University, No. 373, 54 pp. 45 hp (45 ECTS credits).

Abstract: The Campanian marine strata from Åsen, north east part of the Kristianstad Basin (southern Sweden) contain a most diverse vertebrate and invertebrate marine fossil assemblages. A diverse fossil fauna was collected from Campanian deposits from a ca. 4.15m thick section during June-August 2010-2012. The succession is divided into two zones, the latest early Campanian Belemnellocamax mammilatus Zone and early latest Campanian Belem- nellocamax balsvikensis Zone. The B. mammilatus Zone is further divided into distinct units; the Coquina bed, the Green sand bed and the Oyster bank. The B. balsvikensis Zone is divided into Balsvikensis Green and Balsvikensis Yellow. A total of 169 kg material was sorted and the invertebrate fauna collected includes brachiopods, bivalves, belemnites, bryozoans, barnacles, sea urchins and corals with fair amount of vertebrate bones and teeth, and even coprolites. The succession also contains charcoal fragments but most of them are encountered in the lowermost part representing the latest early Campanian marine strata. A quantitative fossil analysis was performed by weight per- centage of the fossils to assess the variation between the different beds of the studied locality. XRF analysis of the sediment samples were performed for elemental analysis of the different beds showing that silica (Si) and calcium (Ca) are the elements constituting the main part of the sediments. Further it is noted that Si and Ca show an inverse relationship through the whole succession. Si shows a decreasing trend all through the succession with highest val- ues in the samples from the terrestrial flood plain deposits and lowest values within the B. balsvikensis Zone. Ca on the other hand, shows the opposite trend with highest values in the B. balsvikensis Zone. XRF analyses of the bel- emnites were further made for temperature proxy based on variations in Sr/Ca ratio between different beds in the sequence. These revealed that the average Sr/Ca values and trend of the belemnites from different beds of the Cam- panian strata show increasing average values of Sr/Ca from the B. mammillatus Zone to the B. balsvikensis Zone. However, as analyses on aragonite were not perfomred furtehr interpretations are out of scope of this study. Char- coal fragments were studied in Scanning Electron Microscopy (SEM) and binocular microscope for identification, and in order to distinguish the source of the charcoal. This study revealed the presence of mainly conifer wood but with some angiosperm wood present. A stratigraphical log was compiled based on the fossil content and sedimen- tological results from the field study. The amount of pelagic fossil fauna and inoceramids identified suggests high sea levels, also the high faunal diversity and the XRF Sr/Ca of the belemnites suggests warm climates and high primary productivity during the latest early Campanian at Åsen. A sea level drop is inferred by the high amount of benthic communities and steinkern identified in the B. balsvikensis Zone, further, the size of the fossil fauna, the presence of the cold water carbonate producing fossil fauna suggest cooling sea temperatures during the early late Campanian at Åsen.

Keywords: Campanian, Belemnellocamax mammilatus Zone, Belemnellocamax balsvikensis Zone, Brachiopods, Bivalves, Belemnites, Bryozoans, Barnacle, Sea urchin, Corals, Coprolites, Shark Teeth, Vertebrate fossils, XRF, Sr/Ca, Steinkern

Supervisors: Vivi Vajda, Elisabeth Einarsson

Faisal Javed Iqbal, Department of Geology, Lund University, Sölvegatan 12, SE-223 62 Lund, Sweden. Email: [email protected]

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Paleoekologi och sedimentologi i marina sediment från övre krita (campan) från Åsen i Kristianstadsbassängen, Skåne

FAISAL JAVED IQBAL

Iqbal, F. J., 2013: Paleoekologi och sedimentologi i marina sediment från övre krita (campan) från Åsen i Kristian- stadsbassängen, Skåne. Examensarbeten i geologi vid Lunds universitet, Nr. 373, 54 sid. 45 hp

Sammanfattning: De marina sen-kretaceiska fossilen från lokalen Åsen som ligger i den nordöstra delen av Kris- tianstadsbassängen i södra Sverige består av en campansk fossilfauna med stor mångfald. Fossil från den ca 4,15 m exponeringen av den marina sanden samlades in under utgrävningar i juni-augusti 2010-2012. Totalt silades och sorterades 169 kg marin sand. Stratigrafin av den marina sanden delas in i två zoner baserat på belemniter: den äldre Belemnellocamax mammilatus zonen som representerar sen tidig Campan samt den yngre Belemnellocamax balsvikensis zonen som representerar tidig sen Campan. Belemnellocamax mammilatus zonen delas i sin tur in i krosslagret, grönsand och ostronbanken. Belemnellocamax balsvikensis zonen delas i sin tur in i balsvikensis grön och balsvikensis gul. De insamlade ryggradslösa fossilen representeras av brachiopoder (armfotingar), musslor, belemniter, bryozoer (mossdjur), rankfotingar (havstulpaner), sjöborrar och koraller. Ryggradsdjurens förhistoriska existens bevisas av fossil från tänder, benfragment och koproliter. Dessutom återfanns kolfragment, men främst i den äldre delen av stratigrafin som kan ha sitt ursprung i det underliggande flodplanet.

Fossilen undersöktes kvantitativt då viktprocent studerades för att bedöma variationen mellan de olika lagrena inom de båda zonerna. Sedimentprover undersöktes med hjälp av XRF-analys för grundämnesanalys, vilket visar att kisel (Si) och kalcium (Ca) är de vanligaste grundämnena i sedimenten. Si och Ca visar ett omvänt förhållande genom hela stratigrafin på Åsen då Si har de högsta värdena i flodplanet och de lägsta värdena i B. balsvikensis zonen me- dan Ca har de högsta värdena i B. balsvikensis zonen och de lägsta värdena i flodplanet. XRF-analyser har även gjorts på belemniter för att undersöka paleotemperaturen baserat på variationen i Sr/Ca förhållandet i de olika lag- rena inom de båda zonerna. Kolfragmenten studerades i både elektronmikroskop (SEM) och ljusmikroskop för identifiering och ursprungsanalys. De flesta kolfragment kommer ifrån barrträd, men spår av lövträd hittades också. Dessutom har en stratigrafisk logg sammanställts utifrån det fossila innehåll och sedimentologiska resultat som framkom under fältstudierna. Sammanfattningsvis verkar det som om att det under sen tidig Campan som represen- teras av B. mammilatus zonen var höga havsnivåer, vilket bevisas av mängden pelagiska fossil samt inoceramider- na. Dessutom ar det troligtvis varmare klimat med hög primär produktivitet, vilket bevisas av den stora mångfalden i faunan tillsammans med XRF-analysen av Sr/Ca förhållandet hos belemniterna. Troligtvis har en havsnivåsänk- ning infunnit sig under tidig sen Campan som representeras av B. balsvikensis zonen, vilket bevisas av den mindre storleken på fossilen samt mängden bentiska fossil varibland en stor mängd stenkärnor har identifierats. Under tidig sen Campan representerat av B. balsvikensis zonen har även en sänkning av paleotemperaturen noterats bevisat av karbonat värdena samt Sr/Ca förhållandet hos belemniterna som framkom vid XRF-analyserna.

Nyckelord: Campan, Belemnellocamax mammilatus Zon, Belemnellocamax balsvikensis Zon, Armfotingar, Musslor, Belemniter, Bryozoer, Sjöborre, Korall, Koproliter, Hajtänder, Vertebratfossils, XRF, Sr/Ca, Steinkern

Handledare: Vivi Vajda och Elisabeth Einarsson

Ämnesinriktning: Berggrundsgeologi

Faisal Javed Iqbal, Geologiska institutionen, Lunds Universitet, Sölvegatan 12, 223 62 Lund, Sverige. E-post: [email protected]

5 1 Introduction of the belemnites is used to study the variation of the Sr/Ca between different beds in the sequence. SEM Lithological changes and changes in the fossil fau- and binocular microscope is used to study the charcoal na are generally indicative of changes in the environ- fragments. This study further aim to elucidate changes ment. The change in diversity and body size of benthic in the invertebrate fauna through time, from early to and planktonic invertebrate fossil fauna enables the late Campanian in the Kristianstad Basin providing study of the changes of sea water conditions, organic additional information on paleotemperatures and sea influx and sea-level changes (Vajda & Wigforss- level changes. Lange 2006). 2 Background During the Late Cretaceous there were repeated epi- sodes of sea transgression in southern Sweden (Vajda 2.1 The Paleogeography of the Cretaceous & Solakius 1999), which resulted in the development World of an archipelago environment and irregular coast line morphology of the Kristianstad Basin (Christensen A super-continent Pangea was formed by the coales- 1984). The Upper Cretaceous Campanian deposits at cence of all pre-existing continent masses during the Åsen, within the Kristianstad Basin in northeast Skåne late Palaeozoic until Triassic. The southern part of consists of marine unconsolidated quartz sand with fair Pangea is called Gondwana and the northern part is amount of glauconite (Fig. 1). These sediments yield a called Laurasia. The super-continent that started to diverse vertebrate fauna including 34 shark species, break up during the Triassic continued to break up five ray species, six mosasaur species, one plesiosaur further during the Creatceous (145-66 Ma) (Gale 2000; species, and minor dinosaur and turtle remains (Table. Vajda– Wigforss-Lange 2009). The Cretaceous world 1) (Sorensen et al. 2013). The succession also contains was different from our modern world in several as- an invertebrate fauna including bivalves, cephalopods, pects, for example by the distribution of the continents gastropods, brachiopods, bryozoans, echinoderms and and geography. The period was characterized by ex- belemnites (Erlström & Gabrielson 1992; Sorensen & tensive plate movements resulting in extensive rifting Surlyk 2010, 2011; Sorensen et al. 2011, 2012). and formation of mantle plumes. The extensive vol- canism resulted in high concentrations of carbon diox- The main aim of the project is to give an overview of ide (CO2), which might be the probable cause of the the total fauna and ecology at Åsen during the Late global greenhouse conditions (Vajda et al. 2003), with Cretaceous. A further aim is to learn the paleontologi- no polar ice caps (Gale 2000). The sea level-rise dur- cal and sedimentological techniques that are used to ing the Cretaceous was accompanied by massive struc- study the paleoenviroments; i.e. identification, sorting, tural changes globally due to plate tectonics. The unu- weighing and counting of the fossil fauna from B. sual conditions had profound effects on the initial radi- balsvikensis and B. mammilatus zones to investigate ation, diversification and evolution of the flora and the changes in fossil content in different beds of the fauna during the Cretaceous (Gale 2000; Friis et al. Campanian marine strata. The purpose is also to make 2011; Ocampo et al.. 2006). a stratigraphical illustration of the fossil fauna within the B. balsvikensis and B. mammilatus zones. XRF 2.1.1 Plate Tectonics and Volcanism analysis of the sediment samples is used to study the geochemistry of the different beds and XRF analysis During the Cretaceous, major changes took place in 6 Fig. 1. Simplified geological map of the NE Skåne (southern Sweden), Showing the Kristianstad basin and the location of Åsen. Modified from Norling and Bergstrom (1987). the configuration of the plates, which resulted in 1993; McLoughlin 2001). The extensive spreading of changes of the paleogeography and affected the carbon the Pacific Sea floor caused the development of the cycle. The major plate-tectonic events included the terrains on the Pacific’s border during the mid- continued breakup of the Gondwanan continent Cretaceous time (Vaughan 1995). During the Cretace- through the Jurassic and the rifting between Africa and ous substantial mountain building took place. The sub- South America during the Early Cretaceous duction and accretion of the northern part of the Tet- (Valanginian-Aptian) interval (Kennedy & Cooper hys throughout the Cretaceous resulted in development 1975). Development of the passage between the north of the Himalayas-East Alpine Orogeny. Also the ob- and south Atlantic during late Albian is evidenced by duction of Ophiolites over continental crust of Arabian the presence of ammonite fauna (Kennedy & Cooper shield on the southern part of Tethys took place during 1975). The rapid opening of the South Atlantic Ocean, the mid-Cretaceous to Campanian (Churkin 1972; and also the separation and the movement of India Gale 2000). The uplift of the Andeas and the Rocky northward took place during the Campanian- Mountains in America and uplift of the South Chinese Maastrichtian interval. The collision of India with Asia block in Asia were the major orogenic events during also took place at the very end of the Cretaceous the Late Cretaceous (Gale 2000). The four major Cre- (Jaeger et al. 1989). The rifting between Antarctica taceous basaltic flood eruption events took place and Australia started at about 132 Ma and the format- during the Cretaceous: 1. The Parana-Etendeka traps ion of the volcanic centers due to the development of in Brazil during earliest Cretaceous, 2. The Aptian the Indian Ocean by extensive rifting (Muller et al. basalts in Rajmahal eastern India. 3. The Cenomanian 7 basalts of Madagascar and 4. the Deccan traps in India climate during the Cretaceous was a result of a combi- during the lates Cretaceous and ealiest Paleocene nation of various factors for example the low relief and (Muller et al. 1993). even distribution of the continents around the globe, warm and saline deep-sea oceans and that the conti- 2.1.2 Climate and sea level changes nental interiors were covered with shallow seas (Gale 2000). The carbon dioxide levels were very high The paleogeographical changes during the Cretaceous which were attributed to outgassing from magmas were not only due to the plate movements but also during the fast sea floor spreading, which probably fluctuations in the global sea-level, which was pro- contributed to the warm climate (Harald & Olaussen bably driven by the global tectonic events.(Willumsen 2008). Moderate temperatures prevailed at the poles & Vajda 2010) The sea levels during the earliest Cre- with a low temperature gradient from the equator to taceous (late Volgian-early Berriasian) were marked the poles. Due to these facts, water masses transported by lowstand (Rawson & Riley 1982) but this was from the north were not cold and saline enough to sink followed by the Early Cretaceous rapid sea level rises and form bottom waters (Gale 2000; Friis et al. 2011). with little interruption during the Valanginian and There was no major current flow from the poles and Hauterivian and followed by a regression in the Barre- no absorption of oxygen in the water over large areas mian. During the early Aptian, sea transgressed again of the oceans. Combination of all these factors shows but sea level fell briefly, and then followed by the ove- that the average global temperatures during the Creta- rall rise of the sea level during the Cenomanian. The ceous were substantially higher compared to today sea level remained high during the early Turonian, but (Harald & Olaussen 2008). dropped during the mid-late Turonian, however at the end of the Turonian and until the start of the Santo- 2.2 Fauna of Late Cretaceous nian, a major transgression was seen, which resulted in the formation of chalk facies (Gale 2000). The inter- 2.2.1 Terrestrial fauna pretation of the global sea-levels for the late Campa- nian is interpreted differently by different authors; one The Cretaceous life on land was dominated by the di- show moderate and other show high sea levels i.e. nosaurs which includes the well-known carnivore Ty- America and Europe respectively (Haq et al. 1987; rannosaurus rex. The sauropods, gigantic Jurassic Hardenbol et al. 1998). Sea-level dated from herbivore dinosaurs were joined by the ornithischians Maastrichtian indicate a major transgression globally during the Early Cretaceous. Also the foot prints of the (Gale 2000; Ramberg et al. 2008). The sea transgres- Iguanodon marks the presence in the Lower Cretace- sed the vast shelf areas and spread the marine sedi- ous successions (Lloyd et al. 2008). The diversificat- ments far beyond the previous area (Hancock & ion of new herbivore dinosaurs such as Hadrosaur, Kauffman 1979). Neoceratopsians, Ankylosaurs and the carnivorus di- nosaur group including the giant Carcharodonttosauri- Most of the Mesozoic was a Greenhouse world with nes and smaller Troodontids, Dromaeosaurs and no icecaps (Vajda 2001, 2008) and although there was Omnithomimosaurs took place during the Late Creta- possibly ice on the poles during the Early Cretaceous, ceous (Lloyd et al. 2008). The atmosphere was domi- the poles were ice-free and covered with temperate nated by the pterosaurs (flying reptiles) during the forests during the middle and Late Cretaceous (Gale Cretaceous. Diverse bird’ faunas were present in sizes 2000; Vajda & McLoughlin 2005). The hot and humid varying from finch to Ostrich-like forms. The develop- 8

ment and evolution among the vertebrates, like the and echinoderms were also present during the Cretace- Squamates, Crocodilians and other reptiles took also ous (Harald & Olaussen 2008). The vertebrates of the place during the Cretaceous. The development of the Cretaceous seas were dominated by sea turtles, foetus in the placental mammal’s womb and the ap- Ichthyosaurs (fish lizard), crocodiles, sharks, rays and pearance of the marsupials, which developed exter- the giant marine reptiles, the mosasaurs and the plesio- nally in the mother’s pouch, took place during the Cre- saurs (Harald & Olaussen 2008; Lloyd et al. 2008). In taceous. The mammals during that time were mostly the fresh water enviroments ostracodes and algae, such insectivores, but also some herbivores and omnivores as Botryococcus flourished during the Cretaceous species were present (Gale 2000; Friis et al. 2011; (Guy-Ohlson 1992). Lloyd et al. 2008; Erlström & Gabrielson 1992). 2.3 North America – Western Interior 2.2.2 Terrestrial flora Seaway during the Campanian

The Early Cretaceous flora was partly similar to that of During the Late Cretaceous much of North America the Jurassic, which included ginkgo-related species, was covered by a large inland sea, which was at its bennettitaleans, cycads, ferns and seed-ferns (Vajda maximum limits stretching from the Artic Ocean to the 2001; McLoughlin et al. 1995). However there was a Gulf of Mexico during the late Albian to Maastrichtian big difference, the angiosperms radiated during the (Kauffman 1967, 1977; Zangrel 1960; Coban 1969; Cretaceous with the appearance of the modern oak, McNeil & Caldwell 1981; Bercovici et al. 2009, birch, magnolia (Friis et al. 2011). The explosion of 2012). The development of the Western Interior the angiosperms by replacement of the gymnosperm Seaway started by the extension of the southern em- and ferns provided new radiation for pollinating bayment of the Arctic Ocean throughout the Cretace- insects, moths and butterflies (Friis et al. 2011). Also ous (Friis et al. 2011). The marine strata of the Upper the conifer taxa Sequoia which played important role Cretaceous host large amounts of fossil fauna in- during the Cenozoic, appeared during the Cretaceous cluding invertebrates such as mollusks, brachiopods (Harald & Olaussen 2008; Lloyd et al. 2008). and bivalves (Nicholls & Russell 1990). Vertebrates include, amongst others, mosasaurs, plesiosaurs, turt- 2.2.3 Marine fauna les, pterosaurs, birds, fish, and sharks. The climatic conditions during the Early Cretaceous were dry based In the sea, the coccolithophorid algae flourished, these on the low amount of the terrestrial organic matter in produced vast amount of chalk. Other organisms the sediments (Hallam 1984, 1985) signifying sparce occurring in the Cretaceous marine environments were vegetation. Similarly the presence of high organic con- the foraminifera and the silica sponges and the plank- tent in the sediments suggests a temperate humid cli- tonic foraminifera Globigerina, which provided the mate in North America during the mid-Cretaceous ooze that deposited in the deep sea (Erlström & Gabri- (Gale 2000). The high presence of charcoal (possibly elson 1992). Ammonites and bivalves were also com- from wildfires) in the sediments from the Late Creta- mon in the seas. The Cretaceous marine organisms ceous of North-America suggests dry climatic environ- developed some peculiar morphologies: Baculites and ment (Friis et al. 2011). Macroscaphites developed a linear elongated shell and an open spiral shell respectively. Sea urchins, rudists, The Western-Interior Seaway divided North America bivalves, bryozoans, brachiopods, belemnites, corals

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into several faunal provinces based on the composition The Cretaceous sediments are present in two basins in of the invertebrate faunas such as gastropods (Sohl southern Sweden; in the Malmö Basin and in the Kris- 1967, 1971), cephalopods (Jeletzky 1968, 1971), bi- tianstad Basin (Moberg 1888). Both basins differ in valves and ammonites (Cobban & Reeside 1952; Gill their sedimentological characteristics and lithology, & Coban 1966; Kauffman 1977; Nicholls & Russell and in fossils content. Due to the tectonic activity 1990). during the Santonian, which extended until the Maastrichtian, Skåne was divided by the Romeleåsen The fossil flora along the Western Interior Seaway is fault in southwest and the Linderödsås-Nävlinge- represented by leaves, flowers, pollens, stems and Hallandsås ridge in northeast (Norling & Bergström roots and reveals the presence of ferns, cycads, coni- 1987). Those tectonic activities probably ceased in the fers and angiosperms. The Albian flora, in particlular, northeastern part during the Campanian but continued provides important information about the early radi- in the southwestern part. The sediments in the Malmö ation and diversification of the angiosperms during the and Kristianstad basins were deposited in a near shore mid-Cretaceous (Friis et al. 2011). marine environment (Erlström & Gabrielson 1992).

2.4.1 Campanian marine fossil assembla- ges in the Kristianstad Basin - Vertebrate 2.4 Europe during the Campanian fauna During the Late Cretaceous, most of Europe was cove- red by shallow epicontinental seas. Shallow marine The major vertebrate fossil assemblages in the Campa- continental sediments of Campanian age are present in nian strata of the Kristianstad Basin are sharks, turtles, Northwestern Europe (Harald & Olaussen, 2008; Gale plesiosaurs, mosasaurs, rays and dinosaurs. They are 2000). The shallow sea covered an area from the Bri- described below: tish Isles to the eastern Caspian Sea and from the 2.4.1.1 Sharks Fennoscandian Craton in the north to the Alpine fold in the south (Harald & Olaussen 2008; Gale 2000). 38 predatory shark species have been identified from The development of the Alps in the south, and the Kristianstad Basin in the lower Campanian strata spreading of the North Atlantic oceanic crust and in (Siverson 1992a, b, 1993, 1995; Rees 1999; Sorensen the west, were probable cause of the intensive volca- et al. 2012). All of the identified species predominant- nism and earthquake in the central parts of the paleo- ly fed on bony fishes, cephalopods and invertebrates Europe. During the Campanian the sea-level reached (Siverson 1992a). Shark species such as Cretoxyrhina to its maximum, over 100m higher compared to pre- mantelli and Squalicorax kaupi (Agassiz 1843) have sent sea-level (Ziegler 1990). The intense sea-level played important role in the marine ecosystem as they rise linked the southern warmer waters with northern occupy the top of the food web (Shamada 1997; Shi- colder waters, which resulted in lowering the water mada & Cicimurri 2005). Teeth from large nektonic temperatures in western and central Europe (Harald & shark species embedded in the partially digested mo- Olaussen 2008). The connection between the seas to sasaur bones were found in sediments from Western the south and north resulted in the exchange between Interior Seaway of North America and shows that they the faunal provinces (Matsumoto 1973). fed, or scavenged, on those giant reptiles (Everhart et

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Diet Position Sharks Anomotodon hermani, Siverson, 1992a,b C N Archaeolamna kopingensis (Davis, 1890) C N Galeorhinus sp. C N Carcharias aasenensis Siverson, 1992a,b C N Carcharias latus (Davis, 1890) C N Carcharias tenuis (Davis, 1890) C N Cederstroemia nilsi Siverson, 1995 C NB Chiloscyllium sp. Cretodus borodini, Cappetta and Case, 1975 C N Cretalamna appendiculata (Agassiz, 1843) C N Cretorectolobus sp. C NB Hemiscyllium hermani, Müller, 1989 C N Heterodontus sp. 1 C NB Heterodontus sp. 2 C NB Hybodus sp. C N Palaeogaleus sp. C N Paraorthacodus andersoni (Case, 1978) C N Paranomotodon sp. C N Paraorthacodus conicus (Davis, 1890) C N Pararhincodon spp. C NB Paratriakis? sp. C N Polyacrodus siversoni, Rees, 1999 C N Polyacrodus sp. C N Pseudocorax laevis (Leriche, 1906) C N Scapanorhynchus perssoni Siverson, 1992a,b C N Scyliorhinidae sp. 1 C N Scyliorhinus’germanicus (Herman, 1982) C NB Serratolamna sp. C N Squalicorax kaupi (Agassiz, 1843) C N Squalidae spp. C N Squatina spp. C NB Squatirhina sp. C NB Synechodus sp. 1 C N Synechodus sp. 2 C N Rays Rhinobatos casieri, Herman, 1977 C NB Rhinobatos sp. 1 C NB Rhinobatos sp. 2 C NB Rhinobatos sp. 3 C NB Mosasaurs Dollosaurus sp. C N Hainosaurus sp. C N Eonatator sternbergi (Wiman, 1920) C N Platecarpus? sp. N Tylosaurus ivoensis (Persson, 1963) C N Plesiosaurs Scanisaurus sp. P N Turtles Turtle remains O M Dinosaurs Leptoceratopsidae sp. H T Table. 1. Fossil vertebrate species found at Åsen locality, their inferred diet and position in the water column, C=Carnivore, P=Piscivore, O=Omnivore, H=Herbivore, N=Nektonic, NB=Nektobenthic, T=Terrestrial (Modified from Sorensen et al. 2013). 11

al. 1995; Shimada 1997). Six plesiosaurs species have been identified so far from the lower Campanian of the Kristianstad Basin 34 out of the 38 shark species encountered in North (Person 1959, 1963, 1990). Plesiosaurs were reptiles America were identified from the Åsen locality that were adapted to the marine realm, however they at (Siversson 1992). All identified species were active times sought shelter in non-marine environments to predators (Sorensen 2013). Some of the sharks from protect their juveniles (Vajda & Raine 2010). Po- Åsen fed on everything they encounter, whereas some lycotylida was a carnivorus plesiosaur genus with short fed on the smaller fishes and invertebrates (Siverson neck and large, elongated head. The Elasmosurus were 1992a). Carcharias aasensis (Siverson 1992a) and instead long necked, short headed and piscivorus Cretalamna appendiculata (Agassiz 1843) occupied (Massare 1987). These were free swimmers and their the top of the food chain (Siverson 1992a). The bite body could reach up to 14m in length, and weigh up to marks of shark's teeth are common on reptile bones several tons (Massare 1988). They mostly fed on small from the Kristianstad Basin (Einarsson et al. 2010). fishes and cephalopods (Massare 1987; McHenry et al. 2005). However, some of them were also benthic gra- 2.4.1.2 Mosasaurs zers, fed on the invertebrates and used gastrolites to help digesting the food (McHenry et al. 2005; Taylor Mosasaurs were the giant reptiles that occupied the top 1987). So far a polycotylid plesiosaur (Einarsson et al. of the food web during the Late Cretaceous in the seas 2010) and an elasmosaurid plesiosaur Scanisaurus (Russel 1967). Six mosasaur species have previously have been identified from Åsen (Persson 1959). been identified from Kristiansand Basin in the upper lower Campanian strata (Lindgren & Siverson 2002, 2.4.1.4 Turtles 2004, 2005; Lindgren 2004, 2005a). Mosasaurs were carnivores and primarily fed on fish and cephalopods Turtle remains are found at almost every locality in the (Lindgren et al. 2010). All six mosasaur species were Kristianstad Basin; however, only two turtle taxa have found in the lower Campanian strata at Åsen been identified (Persson 1959, 1963; Scheyer et al. (Lindgren 2004, 2005a, b). The largest mosasaur 2012). The turtle’s diet is poorly known, however bi- living in the shallow sea of the Kristianstad Basin was valve remains are found in protostegid turtle’s body Tylosaurus ivoensis (Person 1963). The length of its cavity from Australia in Lower Cretaceous deposits lower jaw reached 1.5 m (Lindgren 2004). The broken (Kear 2006). The large amount of turtle’s remains sug- tooth crowns at the apex of this species suggest that it gests that they were abundant in shallow seas might have fed on fleshy animals (Lindgren 2004). (Sorenson et al. 2013). The remains of the turtles were Clidates propython (Cope 1869) and Eonatator stern- also found in the Camapanian strata from Åsen bergi (Wiman 1920) are both small sized mosasaurs (Soresson et al. 2013). reaching a length of 6 m, and these are common in the Kristianstad Basin. Also the presence of teeth and ver- tebrae from the juvenile Clidates propython (Cope 1869) shows that the locality provided protection for 2.4.1.5 Rays the smaller mosasaurs from larger predators (Lindgren & Siverson 2002, 2004). Totally six ray species have been identified in the Campanian strata from the Kristianstad Basin, all be- 2.4.1.3 Plesiosaurs longing to the Rhinobatoidae (Siverson 1993). They

12

were nektobenthic carnivores, which mostly fed on sasaurs and plesiosaurs (Månsby 2009; Eriksson et al. invertebrates and small fishes (Last & Stevens 2009). 2011). Five of the Rhinobatoidae species were encountered in the Campanian sediments at Åsen, whereas rays are 2.4.2 Campanian marine fossil assembla- rarely found from the other localities in the Kristian- ges in the Kristianstad Basin - Inverte- stad Basin, indicating that they preferred more murky, brate fauna estuarine waters as compared to their modern counter parts (Last & Stevens 2009). 2.4.2.1 Belemnites

2.4.1.6 Dinosaurs Belemnites are an extinct group of marine cephalopods whose ‘‘guard’’ the calcitic body part is mostly found Leptoceratopsid dinosaurs have been identified based in Jurassic and Cretaceous deposits. They had ten arms on the teeth remains from Åsen (Lindgren et al. 2007). like the modern squid which helped them to swim The Leptoceratopsid is a small sized and horned, her- (Stevens 1965; Kröger et al. 2011). The body of the bivore dinosaur, which mostly fed on cycads, conifers belemnites were divided into three parts; the posterior and ferns (You & Dodson 2004). ostrum (guard), the middle phragmocone and the front pro-ostracum (Saelen 1989; Doyle 1990). Belemnites 2.4.1.7 Coprolites had an important position in marine realms as predator on smaller organisms and as prey for large marine rep- Coprolites are fossilized feces and provide a source of tiles (Doyle & Macdonald 1993; Cicimurri & Everhart information about the diet, digestive physiology and 2001; Rexfort & Mutterlose 2006). Several species and trophic levels of the paleoecosystems (Hunt et al. sub species of the belemnites were identified be- 1994; Eriksson et al. 2011). Coprolites can be distin- longing to the genera such as Actinocamax ( Miller guished and identified based on composition as well as 1823), Gonioteuthis (Bayle 1879), Belemnitella the morphology. They can vary in shape, size and (d’Orbigny 1840), Belemnellocamax (Naidin 1864) composition. Systematically spiral shaped coprolites and Belemnella (Nowak 1913) from the Campanian are linked to fish (Häntzschel et al. 1968). The irregu- strata of the Kristianstad Basin (Christensen 1975). lar unspiraled shaped coprolites have generally been Belemnites secrete different parts of calcareous shells linked to dinosaurs and may be related to mosasaurs in different water depths, therefore each secreted part and plesiosaurs then collected from marine sediments shows temperature of an area where it has grown, and (Månsby 2009; Eriksson et al. 2011; Thulborn 1991). the whole belemnite reflects the average temperature The size of the coprolites may vary. For example the of the area where it grew (Spaeth et al. 1971). The coprolite of a theropod dinosaur may reach up to 40 δO18 values of belemnite rostra show that the mean cm in length and 15 cm in width (Chin et al. 1998). temperature during the Campanian in the Kristianstad Two groups of lamniform sharks have been identified basin was 20˚C to 24˚C (Lowenstam & Epstein 1954). from Åsen based on coprolites. The coprolites were The diversity changes of the belemnites are related to linked to macrophagous sharks, based on the morpho- the changes in the Earth’s environment such as sea- logy (a heterotropic mode of spiraling). The other type level changes, climatic changes and mass extinction of coprolite, containing mollusk were interpreted to be events (Christensen 2002).The belemnites got extinct produced by durophagous sharks. Further, large unspi- at the K-Pg boundary along with the ammonites ralled coprolites were interpreted as related to mo-

13

(Russel 1979; Alroy et al. 2008). From Åsen, belemni- inner shelf environments (Erlström & Gabrielson tes of the genus Belemnellocamax are used as key-taxa 1992). Additionally, findings of Crania craniolaris for the uppermost lower Campanian and the lowermost (Linnaeus 1958) and the ichnogenus Podichnus have upper Campanian representing the Belemnellocamax been identified from Åsen (Bromley & Surlyk 1973; mammilatus and Belemnellocamax balsvikensis Sorensen & Surlyk 2008). respectively (Christensen 1975). B. balsvikensis differ from B. mammilatus by its deep alveoli (Brotzen 2.4.2.4 Inoceramids 1960). Also the presence of conellae along with the Inoceramids belong to a bivalve family that appeared white layer in B. balsvikensis distinguishes it from B. in the Permian and became extinct at the end of the mammilatus (Christensen 1975). Cretaceous (Hilbrecht & Harries 1992). They were 2.4.2.2 Oysters dominant among the bottom communities especially in the benthic oxygen restricted environments (Kauffman Oysters are bivalves that are found in coastal waters, & Harries 1992). Numerous inoceramid fragments and also in estuarine areas. They are attached to the have been collected from the sediments of the Kristi- substrate such as stones, pilings, and mangroves. The anstad Basin (Lundgren 1974: Erlström & Gabrielson oyster Acutostrea incurva was identified from Åsen in 1992). Campanian strata (Nilson 1827). The oysters from Åsen have thin shells with common fragile imbricat- 2.4.3 Campanian flora in the Kristianstad ions and xenomorphic structures (Sorensen & Surlyk Basin - Fossil wood (Charcoal) 2008). On the right valve the Oysters have mostly xe- nomorphic longitudinal cylindrical ornamentation and Fossil wood is found throughout the fossil record as it on the left side they have attachment imprint, which is is resistant to both biotic and abiotic factors. Charred attributed to attachments to the deciduous trees in the wood fragments provide essential insights into the mangrove (Christensen 1975). Many oyster valves are plants anatomy (Herenedeen 1991a, b; McLoughlin et found together with complete shells in the Campanian al 1995). However, the charcoalified wood is fragmen- sediments (Sorensen & Surlyk 2008). tary and brittle so the information acquired from the charcoal about the host plant is incomplete (Friis et al. 2.4.2.3 Brachiopods (ex crania) 2011). Charcoal has been found within the Kristian- stad Basin and dated to late Santonian to early Campa- Brachiopoda are invertebrates that have sedentary fee- nian based on megaspores (Koppelhus and Batten ding behavior and have great resemblance with the 1989) and also based on the overlain strata identified mollusks (Rudwick 1970), however; their body axes, to be of early Campanian age based on the presence of gill, foot and gonads are different from the mollusks Belemnellocamax mammillatus (Christensen 1975; (Pennington & Striker 2001). Several brachiopods Friis et al. 2011). The initial radiation and diversificat- species have been identified from the Kristianstad Ba- ion of the angiosperms (flowering plants) took place sin (Lundgren 1885; Hägg 1947) amongst those the during the Early Cretaceous beginning from c. 135 Ma genera Crania (Carlsson 1958) and Terebratula (Friis et al. 2011). A world known Campanian (Hadding 1919). Also the inarticulate brachiopod charcoalified flora from Åsen have been described by Isocrania ignabergensis has been identified. This is a (Friis & Skarby 1981). The assemblages contain wood thin-shelled species, characteristic of higher energy from angiosperms but also reproductive organs, lea- 14

Fig. 2. photograph showing diffrerent invertebrate fossil fauna sorted from the Campanian marine strata at Åsen. Corals (microbacia) = G1, Inoceramids = G2, Barnacles = G3 and Steinkern (inoceramids) = G4. 15

ves; along with conifers twigs (Friis et al. 2011). of the archipelago environment and irregular coast line About 100 taxa and 20 flowers species have been iden- morphology (Christensen 1984). The quartz rich sedi- tified from the charred fossil plant assemblages from ment and kaolin of fluviatile origin are present in the Åsen (Friis et al. 2011 and references therein). The basal part of the sedimentary portion (Erlström & Ga- presence of mostly same taxa in the lower and upper brielson, 1992). During the latest early Campanian the part of the sequence shows that no hiatus occurs (Friis calcareous sandstone, conglomerates boulder beds, et al. 2011). The charcoalified plant fragments along calcarenites and calcisiltit were deposited (Christensen with the lignitised compression fossils are present 1984). This corresponds to the local biozone throughout the succession; angiosperms flowers, seeds Belemnellocamax mammillatus Zone that can be corre- and fruit fragments are a few millimeters in size but lated to the Belmnitella mucronata senior/Genioteuthis conifers and twigs are found in comparativelty large quadra gracilis Zone in Germany (Christensen 1975). size (Friis et al. 2011). The larger specimens identified Åsen is located in the north-east part of the Kristian- from the locality are all conifers while the angi- stads Basin and the sediments consist of marine sands osperms identified are small fragments (Nykvist 1957; that rest on the lacustrine clay and argillaceous sedi- Herendeen 1991a). ments. The Cretaceous marine stratum at the locality is rich in macro-invertebrate fossils (Lundgren 1934; 3 Geological Setting Lindgren & Siverson 2002). Glacially tectonised, un- consolidated sand overly the upper Santonian/ Campa- The Kristianstad Basin is located in Skåne, southern nian lacustrine clayey argillaceous sediments (Friis & Sweden and has c. 200 m thick Cretaceous marine Skarby 1981; Siverson 1992) and 3.5 m of these sedi- strata exposed (Christensen 1984). During the Cretace- ments are exposed at the Åsen locality. The sediments ous period the basin was formed due to the southwest in the exposed succession were sampled at different dipping of the crystalline bedrocks, with faults for- levels. The rock unit is divided into two parts, the lo- ming the Nävlingeåsen and Linderödsåsen horsts wer Campanian B. mammillatus Zone and the upper (Bergström & Sundquist 1978). The northern part of Campanian B. balsvikensis Zone (Christensen 1975; the Kristianstad Basin is irregular, having several Up- Lindgren et al. 2007). per Cretaceous outliers exposed outside the basin and on the south-eastern part of the basin lies the Hanö The sediments of the B. mammillatus Zone consist of Bay. However, the sedimentary bedrock extends into sandstones with calcarenites and calcirudites and host the Baltic Sea (Bergström & Sundquist 1978). During vertebrate bones and teeth, belemnites, oysters, corals the latest Triassic to Middle Jurassic the temperature and coprolites. The coquina bed at approximately 1.5 was warm and humid, the weathering of the kaolinized m below the erosional layer in the B. mammillatus bedrock resulted in the development of the uneven Zone, consist of green, coarse quartz sand with belem- topography of the Precambrian bedrock (Lidmar & nites, oysters, coprolites and vertebrate bones and Bergström 1982). shark teeth (Rees 1999; Lindgren et al. 2007).

Four major transgressive pulses occurred during the 4 Materials and Methods Cretaceous (Bergström & Sundquist 1978), which covered the crystalline Precambrian rocks (that were The samples were collected from different beds during exposed at the time) and resulted in the development excavations in summer, 2010 - 2012 from the Campa-

16

nian successions at Åsen, southern Sweden (Fig. 3). 4 micron thick polypropylene film. The samples were The fieldwork was performed by several volunteers analyzed during 240 sec with a portable Niton XL3t and students under the supervision of Elisabeth Einars- XRF analyzer at Department of Geology, Lund Uni- son. The samples were first sieved in the field and the versity. The data reduction was performed with the larger fossil fragments were separated. The remaining program NDTREL-08. The analytical data from the samples were collected in bags, got marked with the different samples are shown in the Appendix II. name of the bed they were collected from, and with the excavation date. The samples were processed by wet 4.2 XRF analysis of the Belemnites sieving through a mesh size of 2,00 mm and were then Belemnites were also analysed with the XRF method. put into steel dishes and dried in the oven over night. Several belemnites were collected from different lay- After the samples dried, the sediments were separated ers from the Belemnellocamax mammillatus Zone and from the fossils and were put in different plastic boxes from the Belemnellocamax balsvikensis Zone. Diffe- marked with sample names as designated on the plas- rent belemnite specimens from the same beds were tic sample bags. Different fossil fragments i.e. the analyzed in order to obtain the Sr/Ca variations for shells without any structure and the belemnites were paleotemperature estimation. The phragmocone and put in large plastic boxes whereas the shells with rostrum solidum of the belemnites were analyzed to structures, charcoal, corals, shark teeth, bone frag- check the variation within single belemnites. Before ments, were put in the same colored box sorted into the analysis the belemnites were cut in halves, and the different segments (Fig. 2). cut surface was gently polished using a carborundum Among the sorted fossils the following groups were disc. Some belemnites have an inorganic phragmocone identified and used for the diversity and paleoenviro- layer in the center surrounded by brown rostrum so- ment study in this project; shark teeth, plesiosaur lida. In most cases, the belemnite phragmocones and tooth, gastrolits, charcoal, belemnites, shells, the rostrum solida are formed by brown calcite. The inoceramids, sea urchins (spines and plates), bryozo- samples were analyzed in 240 sec with a portable Ni- ans, barnacles and coprolites. The sorted fossils were ton XL3t XRF analyzer at Department of Geology, weighted and put in a comprehensive excel data set. Lund University. The data reduction was done with the Total amount of the sample along with fossils in per- program NDTREL-08. The data is presented as parts centages are presented in the Appendix I. The percen- per million (ppm). In this study the values of Si, Ca tages (based on weight) of the fossil and sediment and Sr were estimated (see Appendix III). The average components from the different beds within the value is taken from all the belemnites of the same bed. sequence, are presented as pie charts (Fig. 4; Fig. 5;

Fig . 6) 4.3 SEM Charcoal Analysis

4.1 XRF analysis of the sediment samples The primary tool used to study the charcoal was the binocular microscope as is a fast and inexpensive In total 19 samples were collected from the different method. The binocular microscope is used to sort the beds and were crushed to powder for the XRF analy- charcoal from bones and sediments. Also the weathe- sis. A small amount was taken from each powdered red and well-preserved (=fine) charcoal fragments sample after making sure that the sample was homoge- were separated by using the binocular microscope. neous. The powder was placed in a sample cup with a Then to further anatomical study of the charcoal frag- 17

ments, Scanning Electron Microscopy (SEM) techni- 2. The middle ‘’Greensand layer” lies on top of the que was used. Only the unweathered and well preser- Coquina bed which has a thickness of c. 0.5 m. The ved charcoal fragments were studied under SEM. bed contains glauconitic-rich quartz sand, including fossils such as bryozoans, sea urchins, belemnites, A Hitachi S-3400N SEM at Department of Geology, inoceramid fragments, charcoalified fragments, copro- Lund University. The samples were gold plated and lites, shark teeth, bony fishes vertebrae, and bone frag- mounted on a 3mm diameter SEM stub. Imaging was ments. done using the secondary electron detector. 3. The following uppermost ‘’Oyster bank’’ , named 5 Results based on the high oyster abundances has a thickness of c. 0.6 m. The bed is predominantly represented by 5.1 Stratigraphy calcareous chalk with an abundant oyster fauna. The bed also contains other fossils fragments such bryozo- The stratigraphy of the studied locality is mostly based ans, sea urchins (plates & spines), micrabacia, incore- on belemnite zonations (Christensen 1975), and other mids and coprolites. fossil groups such as oysters. The different beds are in this study named based on the lithology and the colour 4. The lower ‘’Balsvikensis Green’’ named after the of the sediments. Fig. 4 shows the different beds wit- green coloured glauconitic-rich sand with a thickness hin the stratigraphical succession along with the fossil of c. 1.2 m. ‘’Balsvikensis Green’’ consists of gra- content. The total thickness of the exposed marine nules, pebbles, carbonate cemented nodules and stein- Campanian strata at the study site reaches about 4,15 kerns. The fragmented fossils of the ‘’Balsvikensis m (Einarsson et al. in press). A total of 169 kg material Green’’ including belemnites, inocermids, micrabacia, was sorted and a diverse fauna with coprolites barnacles, bryozoans, coprolites, shark teeth, bony and charcoalified phytoclasts as an additional compo- fishes vertebrae and tiny amount of charcoal frag- nent was identified. The studied succession is divided ments. into two zones, the lower B. mammilatus Zone of latest 5. The uppermost bed ‘’Balsvikensis Yellow’’ named early Campanian and upper, younger B. balsvikensis based on the yellow colored sand, is quartz-rich and Zone of earliest late Campanian age (Christensen reaches a thickness of c. 1.6 m. In the base of this bed 1975). a slightly whiter interval can be detected but is herein The lower B. mammilatus Zone is further divided into included in ‘’Balsvikensis Yellow’’. This bed contains three beds; granules, pebbles, carbonate cemented nodules and steinkerns. ‘’Balsvikensis Yellow’’ further contains 1. The lowermost ‘’Coquina bed’’ based on its content fossil fragments of oysters, belemnites, bryozoans, of fossil shells with a thickness of c. 0.25 m. The co- micrabacia, barnacles, sea urchins, inoceramids, quina bed is a mottled storm deposit consisting of gre- coprolites, gastrolites, shark teeth, plesiosaur tooth, enish coarse sand, rich in fragmented oysters, belemni- bony fishes vertebrae and small quantity of charcoal tes, inoceramids, micrabacia, coprolites, shark teeth, fragments. Quaternary cover is present above the up- bony fishes vertebrae and bone fragments. The bed per B. balsvikensis Zone. also contains charcoalified wood, both micro and macro sized.

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Fig. 3. Phtograph showing the Quaternary cover and two beds of Campanian marine strata at Åsen represen- ting the B. balsvikensis Zone (earliest late Campanian). Quaternary cover = QL, Ballsvikensis Yellow = BY & Balsvikensis Green = BG.

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Fig. 4. Stratigraphical log showing the lithology and fossil fauna of the Campanian marine strata at Åsen representing the B. mammillatus Zone (latest early Campanian) and the B. balsvikensis Zone (earliest late Campanian).

20

5.2 Results of the XRF analysis balsvikensis Zone. (Sediments) Stronitium (Sr) has slightly higher values in the flood plain deposits as compared to the overlying B. mam- Analytical XRF data from the 19 sediment samples millatus Zone. Sr shows an increasing trend in the two from the Campanian succession at Åsen (Fig. 5, Table zones, with lowest values in the samples from the B. 6) is represented by Si, Ca, Fe, K, Al, P and Sr. mammillatus Zone and highest values within the . Silica (Si) and calcium (Ca) are the major elements balsvikensis Zone found in the analysis; Si and Ca show an inverse relat- The following components occur as trace elements; Zr, ionship through the whole succession. Si shows a Ba, Zn, Th, U, Nb, Mo, Y and Rb in the 19 analyzed decreasing trend all through the succession with samples and these elements occur in almost same pro- highest values in the samples from the terrestrial flood portions throughout the succession (Table 6). plain deposits and lowest values within the B. balsvi- kensis Zone. Ca on the other hand shows the opposite 5.3 Results of the XRF analysis trend with highest values in the B. balsvikensis Zone. (Belemnites) Average Iron (Fe) content in the flood plain deposit is The results from the geochemical measurements of the slightly over 1% but it fluctuates a lot in the flood belemnite rostra shows that the phragmocone have plain sediments. Fe shows a decreasing trend all +2 through the succession with highest values in the consistently lower values of Sr compared to the +2 samples from the Coquina bed and lowest values wit- rostrum solidum. The Sr values detected from the hin the B. balsvikensis Zone. secondary calcite phragmocone reach 400 ppm to 650 ppm; whereas Sr+2 values from the primary calcite Potassium (K) shows a decreasing trend all through phragmocone reach values 800 ppm to 900 ppm. The the succession with highest values in the samples from Sr+2 values for the rostra solida range from 1000 ppm the terrestrial flood plain deposits and lowest values to 1200 ppm (Table. 8). within the B. balsvikensis Zone, however K shows a small peak in the Green sand bed. As the results from the rostrum solidum is considered more reliable, this study focuses on those results. The Aluminium (Al) shows a decreasing trend all through Sr+2 values from rostrum solidum of belemnites from the flood plain deposit with highest values in the the B. mammillatus Zone range between 1000 ppm and samples from the lower part of the flood plain deposit. 1100 ppm and from the B. balsvikensis Zone between Al shows no trend through the whole succession; ho- 1100 ppm and 1200 ppm. wever Al shows a small peak at the boundary between the B. mammilatus Zone and B. balsvikensis Zone. The average Sr/Ca values and trend of the belemnites from different beds of the Campanian strata (Table 2; Phosphorus (P) shows slightly high values in the flood Fig. 5) show increasing average values of Sr/Ca from plain deposits as compared to the overlying B. mam- the B. mammillatus Zone to the B. balsvikensis Zone millatus Zone. P shows an increasing trend in the two which indicate possibly lowering in paleotemperature zones, with lowest values in the samples from the B. or diagenetic alteration. mammillatus Zone and highest values within the B.

21

B. B.

Zone Zone

Zone andZone the

B. balsvikensis

Zone, the

B. mammillatus

ones

o z

B. mammillatus

Zone at Åsen.

Graph showing XRF showing trends Graph elementsof (Si, Ca, Fe, P,K, Al, Sr) of in weight% sediments from beds different of

Fig. 5.

and Flood plain deposit.The trendlast Graph on the showing trend the of Sr/Ca of belemnites the differentfrom Beds tw from balsvikensis 22

Layers name Depth (m) Phragmocone Rostrum solida

Si Ca Sr Sr/Ca Si Ca Sr Sr/Ca

Balsvikensis gul 3.75 1665 408050 658 0.0016 2057 424638 1129 0.0026

Balsvikensis vit 3.35 1319 378400 342 0.0009 2065 407662 1106 0.0027

Balsvikensis Green 1.5 1976 423430 1175 0.0027 1964 426846 1098 0.0025

Oyster bank 1.05 1769 401242 684 0.0017 2049 410025 1007 0.0024

Green sand 0.5 2666 426468 499 0.0011 2282 428040 1056 0.0024

Coquina Bed 0.25 3337 417935 961 0.0022 2598 433504 1060 0.0024 Table. 2. XRF values of elements (Si, Ca & Sr) in ppm and the Sr/Ca of belemnites from the phragmocone and rostrum solidum from the different beds of Campanian marine strata at Åsen.

5.4 Results of the SEM 5.5 Results of the fossils

The charcoal fragments were analyzed under the bi- The following fossils are identified and included in nocular microscope and were sorted into two datasets this project: shark teeth, plesiosaur tooth, gastrolits, grouped in weathered charcoal and fine charcoal. The charcoal, belemnites, shells, inoceramids, sea urchins charcoal is weighted and graphs were drawn to see the (spines & plates), bryozoans, barnacles and coprolites. changes in the different layers. The images taken un- The pie graphs show the overall decrease in the faunal der binocular microscope and SEM are used to identi- diversity in the B. balsvikensis Zone as compare to the fy the plants. The images were taken from different B. mammilatus Zone. The carbonate content in the B. angles to see anatomical structures of the wood. Most balsvikensis Zone is higher than in the B. mammilatus of the analyzed fragments are gymnosperms but few of Zone. The bar graphs for the charcoal, inoceramids, them are angiosperms. Most of the identified gym- coprolites, vertebrates’ bones and teeth show high pe- nosperms are conifers e.g, pinaceae. From the ana- aks in the B. mammilatus Zone; however the benthic lyzed fragments different cell vessels were identified, community like the micrabacia, barnacles, sea urchins that helped to identify what plant group they belong to and bryozoan show high peaks in the B. balsvikensis (Fig. 13, 14, 15, 16, 17 & 18). Zone. For SEM analysis less than 10 charcoal fragments were chosen, SEM images were taken from different angles to study the anatomy of the plant cell. The ima- 6 Discussion ges taken of the charcoal fragments were Transverse Section (TS), Radial Longitudinal Section (RLS), 6.1 Fossil Fauna Tangential Longitudinal Section (TLS) and Stem Exte- rior (Ext). The studied succession at Åsen comprise ca. 4,15 m sediments and these are divided into two zones, the B.

23

G2

G3 G4

G5

Fig. 6. The result of the overall fossil fauna and sediment content in the different beds of Campanian marine strata at Åsen. G1 showing Coquina bed, G2 showing Green sand bed and G3 showing Oysterbank bed repre- senting the B. mammillatus Zone. G4 showing Balsvikensis Green bed and G5 showing Balsvikensis Yellow bed representing the B. balsvikensis Zone. Blue color showing the Granules and pebbles, red color for carbonate cemented nodules, green color for Granules, pebbles & carbonate cemented nodules (same as red and blue to- gether), purple for total fossil content and light blue for Steinkern.

24

Balsvikensis Yellow Balsvikensis Yellow

Balsvikensis Green Balsvikensis Green

Oysterbank Oysterbank

Greensand Greensand

Coquina Bed Coquina Bed

0 0.2 0.4 0.6 0 20 40 60 80

G1 Fine Charcoal Weathered Charcoal G2 Belemnites Shell fragments

Balsvikensis Yellow Balsvikensis Yellow Balsvikensis Green Balsvikensis Green Oysterbank Oysterbank Greensand Greensand Coquina Bed Coquina Bed 0 0.5 1 1.5 2 0 0.05 0.1 0.15 Bony fishes vertebrae Shark teeth Coprolites G3 G4 Bone fragments

Fig. 7. Bar Graph showing fossil fauna weight% from different beds of the Campanian strata at Åsen. Coquina bed, Green sand and Oysterbank representing the B. mammillatus Zone. Balsvikensis Green and Balsvikensis Yellow representing the B. balsvikensis Zone. G1 showing Charcoal weight% (red for fine charcoal & blue for weathered charcoal). G2 showing belemnites and shell fragments weight% (red for belemnites & blue for shell fragments). G3 showing coprolite weight%. G4 showing vertebrate bones and teeth weight% (green for bony fishes vertebrate, red for shark teeth and blue for vertebrate bones). mammilatus Zone and the B. balsvikensis Zone, addit- ion of the coquina bed (G1, Fig. 6 ). ionally the succession is divided in beds based on the ii. The bed also contains high amount of charcoal lithological and fossil content. The succession is divid- and wood fragments which might be reworked ed into following entities: from the underlying flood plain deposits or 1. The coquina bed is the lowermost bed within the B. derived from the terrestrial flora transported mammilatus Zone which is dominated by a variety of from land into the marine basin (G1, Fig. 7). fossils such as shell fragments, belemnites, charcoal, iii. The amount of bone fragments, charcoal, shell coprolites, inoceramids, vertebrate bones and teeth. It fragments and belemnites in small bed suggests is a dark green glauconitic rich quartz sand bed. a storm deposit (G1, G2, Fig. 7). i. The amount of the fossil fauna and the compo- iv. The presence of dark green glauconitic rich sition of the sediments suggest high energy, quartz sand and siliceous preservation of the inner shelf sea environment during the deposit- fossils such as charcoal and shell fragments 25

Balsvikensis Yellow

Balsvikensis Green

Oysterbank

Greensand

Coquina Bed

0 0.05 0.1 0.15 0.2 0.25 0.3 0.35

G5 Bryozoa Sea urchin plates Sea urchin spines Barnacle Micrabacia Inoceramid

Fig. 8. Bar Graph showing weight% of benthic fauna from different beds of the Campanian strata at Åsen. Co- quina bed, Green sand and Oysterbank representing the B. mammillatus Zone. Balsvikensis Green and Balsvi- kensis Yellow representing the B. balsvikensis Zone. G5 showing the benthic fossil fauna representing by blue for inoceramids, red for micrabacia, green for Barnacle, purple for sea urchins spines, light blue for sea urchin plates and orange for Bryozoa.

suggests low sedimentation rate in an open ma- i. The fossil fauna and sediments content suggests rine environment at the water and sediment inner shelf environment because the charac- interface (Odin & Matter 1981; Vajda & Solak- teristics of the sediments develop the benthic ius 1999). The sea water is under-saturated in community (Gray 1981) (G2, Fig. 6). silica. The amount of coprolites, vertebrate bo- ii. The more abundant micrabacia, barnacles and nes and teeth also suggests high food availa- sea urchins compared to the underlying coquina bility and high sea levels for the vertebrates to bed may suggest more oxygenated bottom con- grow (G3, G4, Fig. 7). ditions and also fairly clear waters or may be a small episodic drop in sea level (regression) 2. Green Sand is the second layer of the B. mammi- (G5, Fig. 8). latus Zone and lies above the Coquina bed. It has a higher diversity of the fauna compared to the lower iii. The decrease in coprolites suggests decrease in coquina layer (G2, Fig. 6). the suspended food availability (G3, Fig. 7).

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3. The oyster bank is the upper most layer of the B. ture and drop in sea level, which helped the mammilatus Zone and is dominated by fossil shell benthic communities to flourish (G5, Fig. 8). fragments specially oysters. The overall diversity of iii. The higher amount of the belemnites compared the fossils fauna increases slightly compared to the to the layers in the B. mammilatus zone is due lower Green sand bed. to a drop in sea level that created a shallow i. The amount of glauconitic sand and the silice- marine environment (G2, Fig. 8). ous fauna suggest starvation of sediments due iv. The lower amount of bone fragments and hig- to transgression (G3, Fig. 6) her amount of coprolites (from smaller sharks ii. The bed contains high amount of oyster’s and bony fishes) and shark teeth compared to fauna, which have attachment imprint, which is oysterbank could be due to that the big verte- attributed to attachments to wood, likely rafted brate animals might moved to the deep waters, in from the terrestrial environment (Christensen this show the fall in water level (G3, G4, Fig. 1975). 7). iii. The increase in vertebrate bones suggests high 5. Balsvikensis Yellow is the uppermost bed within the food availability because increase in primary B. balsvikeinsis Zone and is dominated by the sedi- productivity led to more resources for higher ment content of carbonate cemented nodules and the animals in the tropic level (G2, G4, Fig. 6). fossil content is dominated by the invertebrate fau- naThe high amount of carbonate cemented nodules iv. The decrease in amount of coprolites might be suggests shallow water, semi lagoonal environment. related to the presence of the amount of shark The high amount of carbonates may be due to either teeth that also are decreasing. The presence of fall in sea level drop in temperature or both (G5, Fig. high amount of bone fragments could be related 6 ). to the presence of big vertebrate animals that are indirectly related to higher sea levels (G4, Fig. 7). i. The overall fossil diversity is half to that of the 4. The Balsvikensis Green is the lower bed within the Balsvikensis Green but mostly affected the pe- B. balsvikensis Zone of the early late Campanian. The lagic invertebrates and vertebrates which is also overall fossil faunal diversity is lower compared to the an indication of fall in sea level that then Oyster bank and the silisiclastic component is higher. constrained them to move into open sea. It (G4, Fig. 6). could also be due to the decrease in the primary productivity, lack of food, which constrained i. The decrease in overall siliceous fauna, sug- them to move towards better food sources (G5, gests low primary productivity. This decrease Fig. 4; G5, Fig. 7). in primary production affected the abundance and size of the fossil fauna. (G4, Fig. 6). ii. The presence of the benthic invertebrate com- munity along with the carbonates also supports ii. The increase in amount of the benthic commu- the idea for drop in temperature because bent- nity also supports the idea of drop in tempera- hic invertebrates collected from the bed are

27

cold water carbonate producers (Nelson 1988; content. The value of Ca in the B. balsvikensis Zone Rao 1997). are high as compared to the B. mammilatus Zone, which can be due to the presence of carbonate cemen- iii. The presence of steinkerns in the upper strata ted nodules in the B. balsvikensis Zone. within the B. balsvikensis Zone suggests drop in sea level because the steinkern are internal The K high values of the flood plain deposit fits quite molds of an organism, that are formed by the well because K is incorporates in the clay minerals digenetic alteration of the unstable minerals in a lattice due to it large size. Its moderate values in the B. fauna and which were replaced by the calcite to mammilatus Zone are related to the abundance of fos- form carbonate shells in a carbonate rich envi- sil fauna presence because K is strongly adsorbed and ronment (Manning & Dockery III 1992) (G1, incorporated in organic life forms (Wedepohl 1978). Fig. 9). So the declining trend of K in the B. balsvikensis Zone might be related to the low fauna diversity. 6.2 XRF (sediments) The high values of Al in the flood plain deposits fits The high value of the Si in the non-marine flood plain quite well because Al is essential metal of the alumini- might be due to the aluminiumsilicate minerals due to umsilsicate clay mineral. The amount of Al in the B. the fact that the lithology of the flood plain is mostly mammilatus Zone and B. balsvikensis Zone is linear fine clay. Similarly the value of Si is high in B. mam- because of the lithology that is composed mostly of milatus Zone as compare to B. balsvikensis Zone, due sand. to the higher amount of glauconitic sand. The high Si content in the glauconitic sand with high fossil content High Sr concentrations in the flood plain deposits is also suggests low deposition and therefore there is lot related to the fact that Sr incorporates into the clay of fossils during the late early Campanian. The high Si minerals by the weathering of the source rocks. The content also coincides with the abundant biotic life in high values of the Sr in the B. balsvikensis Zone as the B. mammilatus Zone as compare to the B. balsvike- compare to the B. mammilatus Zone are due to the nis Zone. The dissolved skeletal Si from diatoms and high carbonate content in the B. balsvikensis Zone. other silica utilizing organisms after their death en- The Sr increasing trend in the upward part of the suc- riches the deep water by settling through the water cession might be due to the paleotemperature, since Sr column (DeMaster 1981). Due to starved sediments concentration increases in the carbonates due to the the relative abundance of fossils increase within the drop in temperature (Kinsman 1969). sediments. The low value of the Si in the B. Balsvi- 6.3 XRF (belemnites) kensis Zone is may be due vice versa conditions both for sediments and fossil fauna as compared to B. mam- The difference of Sr values in the secondary calcite milatus Zone. phragmocone and that of the primary calcite phragmocone might be due to diagenetic alteration. The calcium peaks are inversed to Si in the whole The phragmocone is composed of aragonite and arago- sequence; the low Ca values in the flood plain can be nite is replaced by the calcite after a long deposition related to the lithology because it is terrestrial flood (Kinsman 1969; Spaeth et al 1971). The distribution of plain. The low value of Ca in the B. mammilatus Zone Sr in aragonite and calcite is ~8. There is a difference is related to the presence of the high siliciclastic fossil

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between the Sr/Ca in the phragmocone and rostrum from B. mammilatus zone (1000-1100 ppm) to B. solidum that might be due to biochemical fractionation balsvikensis zone (1100-1200 ppm). (Kinsman 1969). 6.4 Comparison between the two zones (B. The difference of the Sr concentrations between the mammillatus and B. balsvikensis) at Åsen. phragmocone and rostrum solidum may be due to the fact that the belemnites developed different body parts The fossil fauna comprises both vertebrate and inverte- at different water depths. The phragmocone consists of brate fossil fragments, and the sediment content sug- aragonite, and aragonite do not preserve well because gests inner shelf marine environment. The B. mammi- it is altered by genetic processes and converts it to latus and the B. balsvikensis zones are significantly calcite. The rostrum solidum of the belemnite is the different in sediment and fossil content. The B. mam- well preserved part (Spaeth et al. 1971, 1971b; milatus Zone has an overall high diversity of fossils Kinsman 1969). In this study the rostrum solidum is compare to the B. balsvikensis Zone (Table. 3; Fig. 9). used instead of the phragmocone since the XRF values of the rostrum solidum are consistent whereas there is 6.4.1 Fossil fauna a large fluctuations in the XRF values of The overall diversity of the fossil fauna decreases in phragmocone. the upper B. balsvikensis Zone; however the amount of The Sr values obtained from the rostrum solidum of benthic fauna increase in the B. balsvikensis Zone. the belemnites from different beds of the Campanian This is an indication of drop in sea level, which favo- marine strata at Åsen coincide with the measured va- red the benthic communities to flourish (G5, Fig. 10). lue calculated by Kinsman 1969. The Sr values in cal- The presence of abundant fossil fauna including verte- cite are in the belemnites from Åsen measured to brate remains and fossil shell fragments of bivalves between 1000 ppm to 1200 ppm. According to suggests high sea levels and high food availability Kinsman (1969)” the Sr concentrations of calcite pre- during the latest early Campanian of the B. mammi- cipitated from sea water is about 1200 ppm” During latus Zone. Also higher temperatures because of the normal sea water conditions organisms deposit the high primary productivity resulted in the overall faunal calcite at about 25°C and the Sr concentration in Cal- diversity (G2, G3, G4, Fig. 10). The relatively high cite would be 1000 to 1200ppm and at the same time abundance of charcoal in the latest early Campanian 8200ppm in aragonmite. (Kinsman 1969). The Sr va- strata may be a result of high erosion rate on land with lues from the rostrum solidum in the calcite of some high influx of sediment and charcoal, another scenario belemnites in the B. balsvikensis Zone is above 1200 is that the charcoal was reworked from the lower flood ppm but as the aragonitic value has not been measured plain deposits (G1, Fig. 10). Further, the presence of in this study, the interpretations of sea temperature is steinkerns in the upper strata of the B. balsvikensis out of scope for this study. Zone suggests drop in sea level because the steinkern are internal molds of an organism that are formed by The values of Sr/Ca based on the rostrum solidum of the digenetic alteration of the unstable minerals in a belemnites from the B. mammilatus zone and B. fauna and which were replaced by the calcite to form balsvikensis zone increase slightly i.e 0.0024 to carbonate shells in a carbonate rich environment 0.0027. There is a slight increase in Sr concentration (Manning & Dockery III 1992) (G1, Fig. 9). The decrease in amount of the bone fragments of the B. 29

balsvikensis Zone suggests either drop in sea level or the late early Campanian. The litholog of the B. mam- indirectly decreas in primary productivity by the drop milatus Zone contains glauconite that together with the in temperature. The increase of amount of coprolites presence of siliceous fossil fauna supports the idea of might be related to the amount of shark teeth in the B. high sea levels, and starved sediments. Whereas the balsvikensis zone (G3, G4 Fig 10). The presence of the lithology of the B. balsvikensis Zone containing quartz benthic invertebrate community along with the carbo- rich sand with high amount of carbonates together nates also supports the interpretation for drop in tem- with lower diversity and smaller size of the fossil perature because benthic invertebrate collected from fauna suggests either drop in sea level, drop in tempe- the bed are cold water carbonate producers (Nelson ratures or both (Fig. 7. G2). 1988; Rao 1997). 6.4.3 Paleoecology 6.4.2 Sediment content The lithology and high faunal diversity together with The change in lithology and fossil fauna either its’s the bigger size of the fossil suggests warmer paleotem- diversity or size when comparing the B. balsvikensi- peratures and high sea-levels during the latest early sand B. mammilatus zones of Åsen suggests either Campanain time represented in the B. mammilatus drop in sea level or drop in temperature, or both (Fig. Zone. The larger body sizes together with the high 9. G1).The presence of sand on top of the non-marine amount of the bone fragments of the large vertebrate flood plain and no carbonates in the B. mammilatus animals suggests high food availability for the preda- Zone suggests high sea level or transgression during tors and the prey. This high food availability supported

B. mammilatus B. balsvikensis

Zone Zone Overall Fauna High Low Granules and Pebbles (2-64 mm) High Low Charcoal High Low Bone Fragments High Low Shell Fragments High Low Bony Fishe vertebrae High Low Shark Teeth Low High Belemnites Low High Inoceramids Low High Carbonate Nodules Low High Steinkern Low High Granules and pebbles cemented in carbonate nodules Low High Micrabacia Low High Barnacle Low High Sea Urchin (spines & Plates) Low High Bryozoa Low High Table. 3. Comparison of the fossil fauna and sediment between the zones in Campanian marine strata of Åsen.

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the idea for the high primary production in the surface The charcoal fragment image a01, a02, a03, a04, a05, waters, this high productivity help the pelagic commu- a06 and a07 were referred to as conifer due to the pre- nity to groom and diversify. Reduction in the body sence of homogeneous tracheid and equal ray cells, size together with the decrease in the bones fragments containing two to four circular cross-filed pits. The of the vertebrate animals suggests drop in sea-levels charcoal fragment is highly weathered and compres- during the earliest late Campanain time representing sed, and may be transported from a distant source. by the B. balsvikensis Zone. Also the presence of the The charcoal fragment image b01 and b02 is from a benthic invertebrate community along with the higher conifer based on the fact that the cells are homogene- amount of carbonates also supports the idea for drop in ous. The fragment is charcoalified wood and it is soft, temperature since benthic invertebrate collected from brittle and lightweight. the earliest late Campanian strata are cold water carbo- nate producers (Nelson 1988; Rao 1997). The charcoal fragment image d01 and d02 is referred to a conifer because the cells are homogeneous, it may 7 Results and Discussion about have been deeply buried and transported because the Charcoal charcoal fragment is highly compressed, hard and he- avy weight. 7.1 SEM Charcoal Images (Fig. 13, 14 & 15; Table. 4)

G1 G2

Fig. 9. Pie graphs showing the overall fossil and sediments content from the B. mammillatus (latest early Campanian) and B. balsvikensis (earliest late Campanian) zones at Åsen.

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Fine Charcoal Belemnites

Weathered Charcoal Shell fragments

0 0.05 0.1 0.15 0 20 40 60 80 G1 G2

Bony fishes vertebrae

Shark teeth Coprolites

Bone fragments

0 0.5 1 1.5 G3 0 0.05 0.1 G4

Bryozoa

Sea urchin plates

Sea urchin spines

Barnacle

Micrabacia Legends

Inoceramid B. balsvikensis zone

B. mammilaus zone G5 0 0.05 0.1 0.15 0.2

Fig. 10. Bar graphs showing weight% of different fossils from the B. mammillatus (latest early Campanian) and B. balsvikensis (earliest late Campanian) zones at Åsen.

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Img, scale View plant type comment No a01 3mm TLS Conifer compressed charcoalified wood

a02 500µm TS Conifer compressed charcoalified wood

a03 500µm TLS Conifer compressed charcoalified wood

a04 500µm RLS Conifer compressed charcoalified wood showing ray cells with pits

a05 100µm RLS Conifer compressed charcoalified wood showing ray cells with pits

a06 500µm TLS & RLS Conifer compressed charcoalified wood showing ray cells with pits

a07 100µm TLS Conifer compressed charcoalified wood showing ray cells with pits

b01 2mm TLS Conifer Charcoal fragment

b02 3mm TLS Conifer burnt charcoal fragment

c01 3mm Ext Equisetum? stem with leaves

c02 3mm Ext Equisetum? stem with leaves

c03 4mm Ext Equisetum? stem with leaves

d01 4mm TLS Conifer Strongly compressed charcoalified wood

d02 200µm TLS Conifer compressed charcoalified wood

e01 4mm Ext Conifer compressed charcoalified wood

e02 400µm RLS Conifer compressed charcoalified wood showing broad ray cells & from outer part of the stem f01 3mm RLS Angiosperm compressed charcoal wood

f02 200µm TS & RLS Angiosperm compressed charcoal wood showing the large vessels and parenchyma cells

f03 500µm RLS Angiosperm compressed charcoal wood showing the large vessels and parenchyma cells

g01 500µm RLS Conifer compressed charcoal wood showing the broad ray cells

h01 100µm RLS Conifer compressed charcoal wood showing cross-filed pits & vertical tracheid

Table. 4. SEM results of charcoal fragments. Transverse Section = TS, Radial Longitudinal Section = RLS & Tangential Longitudinal Section = Stem Exterior = EXT

The charcoal image e01 and e02 is considered to be a The charcoal fragment in image g01 is referred to a conifer because the cells are homogeneous. The frag- conifer due to the presence of broad ray cells; it seems ment was buried and transported because it is highly to be transported because it is highly weathered and weathered and compressed; the ray cells are much shows strong burial compression. larger and broader so it may be from a different spe- The charcoal fragment image h01 is referred to a coni- cies than those mentioned earlier. fer due to the presence of ray cells of uniform charac-

The charcoal in image f01, f02 and f03 are referred to ter with circular cross-filed pits. an angiosperm plant due to large vessel cells and small 7.2 Binocular Microscope Charcoal Images parenchyma cells. The charcoal fragment is long trans- ported, because it is highly weathered and compressed. (Fig. 18, 19 & 20; Table. 5)

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Img. No View plant type comment

16-1 TLS Conifer Charcoalified wood, soft and brittle in appearance

16-2 TLS Conifer Charcoalified wood, soft and brittle in appearance

16-3 TLS Conifer Compressed lignite fragment

16-4 TLS Conifer Compressed lignite fragment

16-5 TLS Conifer Compressed charcoal wood

16-7 TLS Conifer Compressed lignite fragment

16-8 TLS Conifer Compressed lignite fragment

16-9 TLS Conifer Compressed charcoalified wood

05-1 RLS Conifer Compressed lignite fragment

05-2 RLS Conifer Charcoalified wood, soft and brittle in appearance

05-3 RLS Conifer Charcoalified wood, soft and brittle in appearance

05-4 RLS Conifer Compressed lignite fragment

08-1 RLS Conifer Compressed lignite fragment

10-1 RLS Conifer Compressed lignite fragment

10-2 RLS Conifer Compressed lignite fragment

10-3 RLS Conifer Charcoalified wood, soft and brittle in appearance

10-4 RLS Conifer Charcoalified wood, soft and brittle in appearance

16-13 RLS Conifer Compressed lignite fragment

22-1 RLS Conifer Compressed lignite fragment

22-2 RLS Conifer Charcoalified wood, soft and brittle in appearance

Table. 5. Identification of charcoal fragments based om binocular microscopic analuzes: Transverse Section = TS, Radial Longitudnal Section = RLS & Tangential Longitudnal Section = Stem Exterior = EXT

The charcoal in images 16-1 and 16-2 are considered dinal and ray cells are homogeneous. They have been to be conifer wood due to the presence of homogene- buried and transported because they are extremely ous ray cells, the specimens may be charcoalified frag- compressed, hard and heavyweight. ments from a wildfire, because they are black, glassy, soft, brittle and lightweight. The charcoal in images 16 The specimens illustrated in 05-1, 05-4, 08-1, 16-13, -3, 16-4, 16-7 and 16-8 are conifers based on the pre- 10-1, 10-2 and 22-1 are referred to conifer wood, ba- sence of longitudinal and homogeneous ray cells. They sed on the fact that the longitudinal and ray cells are have brown compressed layers and may be lignified. homogeneous. They are lignitized, because they are They are hard, compressed and heavy which might be compressed, hard, heavyweight and are brown. The because they have been buried, lithified and transpor- images 05-2, 05-3, 10-3, 10-4 and 22-2 are also refer- ted from a distant source. The charcoal in images 16-5 red to conifer wood; however they are burnt and 16-9 are referred to conifers, because the longitu- (charcoalified) wood from wildfires because they are soft, brittle, lightweight and glassy. 34

are mostly found in glauconitic sand deposits (Bell & Goodell 1967; Tooms et al 1970; Giresse & Odin 7.3 Results and discussion about the flo- 1973). They are often produced by filter feeding org- wer/sedges/equisetum (Table. 4; Fig. 11) anisms in shallow water (Moore 1939). These are elo- ngated cylinders with rounded edges having latitudinal The charcoal fragment image c01, c02 and c03 is ten- lines around the cylindrical body. Total lengths of the tatively referred to as Equisetum stem node, because all clustered cylinders are 5 mm and 1mm in width. the stem of Equisetum in early development has leaves The size of the individual cylinder is 0.8 to 1 mm in that form a whorl around the stem at each node. Anot- length and 0.1 to 0.3 mm in width. Further studies are her possibility, however more unlikely is that this fos- neede in order to link them to an organism. sil represents a Sedge, Sedge have blade-like leaves that whorl around the stem attached to nodes in the 9 Conclusion early development but when they grow up the leaves detach the blade from the stem leaving exposed nodes. A marine succession of Campanian age from Åsen, Kristianstads Basin, southern Sweden 8 Discussion about the eggs/fecel pellets/ was studied bed by bed to describe and assess small coprolites (Fig. 12) the diversity and abundance of the fossils as- semblages by study the fossil and sediment A small cemented cluster of cylindrical shaped bodies content. The studied Campanian marine strata were found in the sediments from the B. mammilatus at Åsen is divided into two zones, the latest Zone. The image Fp-1 and Fp-2 referred to as eggs at early Campanian B. mammilatus Zone and the the first sight, but their close morphological study un- earliest late Campanian B. balsvikensis Zone. der the binocular microscope suggests them to be The B. mammilatus Zone is further divided into faecal pellets. Faecal pellets are argillaceous matter several beds: the Coquina bed, the Green sand that contain variable amount of organic matter, they

c01 c02

Fig. 11. SEM images of the charcoal fragment referred to as Equisetum from the B. mammillatus Zone at Åsen. c01 and c02 showing broken leaves attached to the node of broken stem of Equisetum (3mm).

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bed and the Oyster bank. The B. balsvikensis amount of the vertebrate bone fragments and Zone was divided into the beds; Balsvikensis increase in amount of the benthic communities Green and Balsvikensis Yellow. including micrabacia, barnacles, sea urchins, and bryozoas, along with the high amount of The rich fossil fauna including brachiopods, carbonate content of the deposits of the B. bivalves, belemnites, bryozoans, barnacles, sea balsvikensis Zone suggests drop in sea level urchins and corals with fair amount of copro- during the earliest late Campanian. lites, vertebrate bones and teeth, indicates that the area was a well-functioning ecosystem dur- The smaller size of the fossil fauna and pres- ing latest early and earliest late Campanian. ence of the cold water carbonate producing fossil fauna from the B. balsvikensis Zone sug- The sediment content and the fossil fauna sug- gest drop in water temperatures during the ear- gest an inner shelf environment. liest late Campanian at Åsen.

The high amount of vertebrate bone fragments 10 Acknowledgement together with the decreasing benthic communi- ty suggests high sea levels during the latest First I want to thank my Supervisor Vivi Vajda. I ap- early Campanian representing by the B. mam- preciate all her contributions of time, ideas, and guid- milatus Zone. ance to make my Master’s thesis project productive and stimulating. I am also thankful for the excellent The high amount of charcoal in the sediments example she has provided as a successful woman geo- belonging to the B. mammilatus Zone suggests scientist and professor. I would also like to express my high erosion rate on land with high influx of sincere gratitude to my Co-supervisor Elisabeth sediments, or it may be reworked from the low- Einarsson for the continuous support during my Mas- er flood plain deposits. ter’s thesis project, for her patience, motivation and The presence of the Steinkern, the decrease in enthusiasm. Her guidance helped me in all the time of

Fp-1 Fp-2

Fig. 12. Plate showing Binocular images of Faecal pellets collected from the B. mammillatus Zone at Åsen. Fp- 1 & Fp-2 showing cluster of cylindrical bodies of faecal pellets (5mm). 36

research and writing of this thesis. The joy and enthu- siasm she has for her research was contagious and mo- tivational for me, even during tough times of my The- sis project. Besides them, I would like to thank An- toine Bercovici (for his help in SEM analysis of char- coal) and Stephen McLoughlin for their encourage- ment and insightful comments for the SEM charcoal images. My sincere thanks also go to Leif Johansson for his help in the XRF analysis. I would also want to thank Klara, Daniel, Hani and Andrea for helping me in sorting the fossils. Special thanks to Ahmad, Majid, Ateeq and Saeed for helping me during my stay in Sweden. Last but not the least; I would like to thank my parents who provided me moral and financial sup- port throughout my life. Your love and prayers made all of this possible.

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a01

a03 a04

a05 a06

Fig. 13. SEM images of charcoal fragments collected from the B. mammillatus Zone at Åsen. a01 showing coni- fer having equal ray cells (3mm), a02 showing homogeneous tracheid of conifer (500 µm), a03 showing ray cell with pits of conifer (500µm), a04 showing tracheids and ray cells of conifer (500µm), a05 showing ray cells with pits of conifer (100µm) and a06 showing tracheiods and ray cells of conifer (500µm).

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a07 b01

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c03

d02 d01

Fig. 14. SEM images of charcoal fragments collected from the B. mammillatus Zone at Åsen. a07 showing ray cells with pits of conifer (100µm), b01showing homogeneous ray cells of conifer (2mm), b02 showing conifer having homogenous ray cells (3mm), c03 showing broken leaves attached to the node of broken stem of Equise- tum (4mm), d01 showing compressed wood of conifer (200µm) and d02 showing compressed wood of conifer (4mm).

45

f03

e01

e02

g01

f01

h01 f02

Fig. 15. SEM images of charcoal fragments collected from the B. mammillatus Zone at Åsen. e01 showimg compressed wood of exterior part stem of conifer (4mm), e02 showing broad ray cells and from outer part of a stem of conifer (400µm), f01compresssed wood of angiosperm (3mm), f02 showing large vessels and parenchyma cells of angiosperm (200µm), f03 showing large vessels and parenchyma cells of angiosperm (500µm), g01 showing broad ray cells of conifer (500µm) and h01 showing cross field pits and vertical tracheid of conifer (100µm). 46

16-1 16-2

16-4 16-9

16-4

16-3 16-8

16-7 16-5

Fig. 16. Binocular images of charcoal fragments collected from the B. mammillatus Zone at Åsen. 16-1& 16-2 showing homogenous ray cells of conifer, 16-3, 16-4, 16-7 & 16-8 showing longitudinal and homogeneous ray cells of conifer, 16-5 & 16-9 showing longitudinal and ray cells of compressed wood of conifer.

47

05-1 05-3

05-2 16-13

08-1 05-4

Fig. 17. Binocular images of charcoal fragments collected from the B. mammillatus Zone at Åsen. 05-1, 05-4, 08-1 and 16-3 showing longitudinal and homogenous ray cells of conifers. 05-2 and 05-3 showing homogenous ray cells of charcolified wood of conifer.

48

10-1 10-2

10-4

10-3

22-1 22-2

Fig. 18. Binocular microscope images of charcoal fragments collected from the B. mammillatus Zone at Åsen. 10-1, 10-2 & 22-1 showing longitudinal and homogenous ray cells of conifers. 10-3, 10-4 and 22-2 showing homogenous ray cells of charcolified conifer wood

49

12 Appendix

Coquina Greensand Oysterbank Balsvikensis Green Balsvikensis Yellow Bed Total sample (total amount 282 501 783 1239 4387.5 sieved sand in kg) Sieved sand (less than 2mm) 276.08115 498.11885 774.24259 1229.03893 4329.91271

Granules & Pebbles (2-64 2.46149 0.56579 3.02728 4.495 0.15633 mm) Carbonate cemented no- 0.00013 0.04246 0 0 29.77139 dules Graules, Pebbles & Carbo- 0 0 0 0.688 13.91762 nate cemented nodules TOTAL FOSSILS (kg) 3.45723 2.2729 5.73013 4.77389 13.25337

Steinkern 0 0 0 0.00418 0.48858

Weathered charcoal 0.017563 0.00355 0.021113 0.00154 0.00231

Well-preserved charcoal 0.00028 0.000168 0.000448 0.0003 0

Shell fragments 1.92144 1.42324 3.34468 1.98 5.14436

Belemnites 1.48722 0.82197 2.30919 2.692 7.91578

Inoceramids 0.00104 0.00114 0.00218 0.00096 0.01134

Microbacia 0.00033 0.00201 0.00234 0.01547 0.01234

Barnacles 0 0.00014 0.00014 0.00075 0.00906

Sea urchin spines 0 0.00061 0.00061 0.00434 0.01704

Sea urchin plates 0 0 0 0.00017 0.00084

Bryozoans 0 0.00004 0.00004 0.00087 0.00125

Bone fragments 0.01299 0.00712 0.02011 0.02223 0.01471

Shark teeth 0.00627 0.00764 0.01391 0.01485 0.01541

Bony fishe vertebrae 0.00052 0.00038 0.0009 0.00124 0.00342

Coprolites 0.00279 0.00095 0.00374 0.00684 0.00955

Others 0.00679 0.00394 0.01073 0.03217 0.09361

Plesiosaur tooth 0 0 0 0.00016 0.00146

Gastrolits 0 0 0 0 0.00089

Appendix I. Quantitative data: weight (kg) of fossil fauna (each group) and of sediment excavated at Åsen.

50

25

-

9.54386 0.00235

0.427608 0.484012 0.283295 2.168611 0.165195 0.041124 0.000603 0.002434 0.044804 0.001808 0.002659 0.005668

26.742253 59.942544

57.20970184 0.713378426 18.03312347 0.537882536 0.396386364 2.612308811 0.378527823 0.045914946

20

-

54.0296 0.00491

0.397589 6.667417 0.602022 1.486084 2.943297 0.207971 0.020143 0.000793 0.001558 0.046571 0.001379 0.005065 0.001273

33.183625

0.02248966

70.98972896 0.663297729 12.59808442 0.669027049 2.079328733 3.545495566 0.476544749

Flood Plain Plain Deposit Flood

1

-

1.41844

0.298289 5.285213 5.810982 2.445575 0.058868 0.019953 0.000567 0.001609 0.038295 0.002152 0.014567 0.004537 0.001228

32.025016 52.165709

68.51111673 0.497635539 9.986409964 1.576312372 8.130726014 2.945939645 0.134890135 0.022277525

0

0.30955

5.248638 0.644053 0.485883 2.366931 0.149402 0.023445 0.000735 0.001017 0.031206 0.001126 0.002451 0.004311 0.001207

25.366011 64.365181

Boundary

54.26550733 0.516422265 9.917301501 0.715736099 0.679847494 2.851205083 0.342339743 0.026176343

0.125

0.120755 1.511317 4.888431 0.021835 0.008489 0.004333

1.8150145 2.5433125 0.0604505 0.0122145 63.999541 0.0001585 0.0006575 0.0007225 0.0010175

24.9370705

53.34787492 0.201455567 2.855633472 2.017025614 6.839892655 3.063674238 0.138516276 0.013637489

Coquina bed Coquina

0 0 0

0.5

1.57421

0.209013 1.068272 8.951823 3.145908 0.026576 57.53075 0.001024 0.016182 0.010685 0.005385 0.004871 0.132065

26.393706

Green sand Green

56.46405525 0.348696388 2.974469795 1.187170674 12.52539074 3.789560777 0.029672104

1.05

0.979511 9.098715 1.760031 0.014033 0.001003 0.000413

21.73858233 46.50534919 0.109099667 0.182010974 1.121441333 2.118963399 1.088530574 12.73092203 2.120133343 0.048805333 0.111832541 0.015667845 65.03220467 0.000140667 0.025785667 0.000882333 0.008856333 0.004018667

Oyster bank Oyster

0

1.35

Layer

3.91366

0.238924 1.187705 7.428947 1.961564 0.122057 0.019584 0.000966 0.021399 0.001814 0.012137 0.003817 0.001677

Erosional Erosional

24.365603 60.673569

52.1253345 7.39486057 0.27968141

0.398596909 1.319896567 10.39458264 2.362899994 0.021865536

0

1.5

0.106654 1.252865 0.836067 1.319531 0.269341 0.004314 0.000372 0.000698 0.019276 0.001147 0.014401 0.002431

15.548964 17.959095 61.958125

33.26389869 0.177930868 2.367288418 0.929121257 25.12836572 1.589507043 0.617167967 0.004816581

Balsvikensis Green Balsvikensis

1.95

1.15743

0.109686 0.731335 0.666592 0.206585 0.018963 0.000298 0.000767 0.028182 0.001213 0.013627 0.002159 0.000777

14.913972 15.740911 66.369538

31.9054603 0.74078369 0.02117219

0.182989154 1.381857483 22.02468267 1.394240178 0.473368869

0

3.35

0.73462 0.00132 0.01821

9.436977 0.080052 0.577616 0.455881 0.410504 0.010986 0.000628 0.023416 0.001409 0.001057

24.254061 63.941894

20.1885249

0.133550752 1.091405432 0.506620555 33.93628215 0.884923252 0.940628866 0.012265869

. XRF values of . values elements of XRF differentfrom beds study at the in site Åsen.

0

Balsvikensis Yellow Balsvikensis

3.75

0.09247 0.95276 0.00031

0.398351 0.415613 0.001878

13.055697 0.9135965 0.0202925 0.0008595 0.0112375 0.0014285 0.0201955

1.80024002 27.1133875 56.9309565

27.93005259 0.154267701 0.442687466 37.93705179 1.100518344 0.952335628 0.022656576

/

2 2 5

P

K Y

Si

Ti Sr

Zr

Fe

Ca Ba Zn Dep

AppendixII Rb

O3 O3

Th

Ele/

AL

Bal Mo

Al2 Fe2

SiO

TiO

P2O

CaO BaO K2O

51

Appendix III. XRF elements (Si, Ca & Sr) values in ppm and Sr/Ca of belemnite’s phragmocone and rostrum so- lidum from different layers of the Campanian marine strata.

Phragmocone Rostrum solida SAMPLE Si Ca Sr Si Ca Sr Sr/Ca Sr/Ca

balsvikensis-Yellow-A 2793 418055 1209 0.0028 2724 422686 1263 0.0029

balsvikensis-Yellow-B 1397 390414 571 0.0014 2512 418173 1120 0.0026

balsvikensis-Yellow-C 838 399547 662 0.0016 1576 434510 1106 0.0025

balsvikensis-Yellow-D 1211 411522 460 0.0011 3076 428805 1168 0.0027

balsvikensis-Yellow-E 1554 416179 507 0.0012 1593 416697 1089 0.0026

balsvikensis-Yellow-B 1662 423927 1232 0.0029 1667 434388 1122 0.0025

balsvikensis-Yellow-C 2318 416316 1106 0.0026 2401 428943 1144 0.0026

balsvikensis-Yellow-D 1712 419594 1035 0.0024 676 421663 1097 0.0026

balsvikensis-Yellow-E 2046 418053 1247 0.0029 1782 405744 1251 0.0030

balsvikensis-Yellow-A 1518 385102 579 0.0015 2140 446247 1065 0.0023

balsvikensis-Yellow-B 1226 407463 559 0.0013 2422 420990 1112 0.0026

balsvikensis-Yellow-C 3290 395993 278 0.0007 3793 416185 1067 0.0025

balsvikensis-Yellow-D 2064 396661 751 0.0018 1975 421728 1055 0.0025

balsvikensis-Yellow-E 894 430393 524 0.0012 2229 417735 1235 0.0029

balsvikensis-Yellow-A 877 423000 319 0.0007 1555 426435 1106 0.0025

balsvikensis-Yellow-B 1391 404802 924 0.0022 1298 422669 1135 0.0026

balsvikensis-Yellow-C 2596 417175 1179 0.0028 2254 422139 1114 0.0026

balsvikensis-Yellow-E 2369 384416 472 0.0012 2734 403843 1094 0.0027

balsvikensis-Yellow-A 2717 369734 851 0.0023 1481 412707 1174 0.0028

balsvikensis-Yellow-B 790 408853 772 0.0018 1662 428191 1070 0.0024

balsvikensis-Yellow-C 902 418316 280 0.0006 2897 432427 1081 0.0024

balsvikensis-Yellow-D 783 402085 821 0.0020 2373 426349 1114 0.0026

balsvikensis-Yellow-E 1375 415425 395 0.0009 1607 424274 1123 0.0026

balsvikensis-Yellow-A 1304 424750 259 0.0006 1971 433752 1195 0.0027

balsvikensis-Yellow-C 985 432252 174 0.0004 3651 422852 1162 0.0027

balsvikensis-Yellow-E 1015 396790 347 0.0008 2012 429253 1002 0.0023

balsvikensis-Yellow-A 1686 411650 225 0.0005 2282 432139 979 0.0022

balsvikensis-Yellow-B 2048 419984 1177 0.0028 2052 430619 1033 0.0023

balsvikensis-Yellow-C 2220 416272 601 0.0014 1708 428367 1214 0.0028

balsvikensis-Yellow-D 1343 410659 463 0.0011 1328 415458 1164 0.0028 52

Phragmocone Rostrum solida SAMPLE Si Ca Sr Si Ca Sr Sr/Ca Sr/Ca balsvikensis-Yellow-E 2277 392478 1110 0.0028 1547 434120 1226 0.0028 balsvikensis-Yellow-A 1429 396218 599 0.0015 1589 423100 1236 0.0029 balsvikensis-Yellow-C 2734 380271 461 0.0012 1511 429909 1248 0.0029 balsvikensis-Yellow-E 1252 419364 255 0.0006 1888 424608 1041 0.0024

Balsvikensis-Vit-B 2403 307932 558 0.0018 2028 393055 1193 0.0030

Balsvikensis-Vit-C 1572 387840 181 0.00046 3218 404446 1130 0.0027

Balsvikensis-Vit-D 666 408629 318 0.0007 1127 414226 1123 0.0027

Balsvikensis-Vit-E 634 409201 312 0.0007 1889 418924 980 0.0023

Balsvikensis-Green-D 2591 401382 1166 0.0029 1364 407902 1140 0.0027

Balsvikensis-Green-E 1070 414034 957 0.0023 1659 421979 1101 0.0026

Balsvikensis-Green-A 2347 416935 1138 0.0027 1106 428017 1075 0.0025

Balsvikensis-Green-B 1690 422972 1226 0.0028 2262 427503 1136 0.0026

Balsvikensis-Green-A 1727 434201 1285 0.0029 1543 430898 1069 0.0024

Balsvikensis-Green-D 2088 428564 1361 0.0031 2221 432262 1186 0.0027

Balsvikensis-Green-B 2354 427265 1388 0.0032

Balsvikensis-Green-A 2674 423898 1349 0.0031 1619 432455 1081 0.0025

Balsvikensis-Green-B 1900 425854 1103 0.0025 1962 431430 1163 0.0026

Balsvikensis-Green-A 1072 427810 985 0.0023 2867 425739 1041 0.0024

Balsvikensis-Green-C 2222 434820 971 0.0022 3043 430282 990 0.0023

Oyster bank-A 3817 287209 182 0.0006 2893 352460 1022 0.0028

Oyster bank-B 1350 362466 316 0.0008 2416 395412 965 0.0024

Oyster bank-C 3348 410496 1029 0.0025

Oyster bank-D 1452 407072 284 0.0006 1586 423257 1042 0.0024

Oyster bank-E 1858 414957 166 0.0004 1120 436449 988 0.0022

Oyster bank-A 3821 408893 366 0.0008 2038 429787 1146 0.0026

Oyster bank-B 3098 429365 1036 0.0024 2726 431352 1069 0.0024

Oyster bank-C 1706 416098 848 0.0020 1905 436948 1062 0.0024

Oyster bank-D 2082 434183 1020 0.0023 2264 434644 978 0.0022

Oyster bank-A 1319 435001 815 0.0018 1716 432451 1012 0.0023

Oyster bank-B 1362 428875 722 0.0016 1842 418951 1063 0.0025

Oyster bank-C 870 424498 472 0.0011 2710 429555 1032 0.0024 53

Phragmocone Rostrum solida SAMPLE Si Ca Sr Si Ca Sr Sr/Ca Sr/Ca

Oyster bank-D 1117 437934 870 0.0019 1267 424547 1031 0.0024

Oyster bank-E 1529 423288 769 0.0018 3634 434496 1136 0.0026

Oyster bank-A 997 422395 530 0.0012 2368 439412 1028 0.0023

Oyster bank-B 1262 432296 353 0.0008 1104 435228 1020 0.0023

Oyster bank-C 2206 432173 1038 0.0024 1840 435144 1122 0.0025

Oyster bank-D 2041 431653 1240 0.0028 2543 438045 975 0.0022

Oyster bank-E 1489 439504 829 0.0018 1722 437492 1035 0.0023

Oyster bank-A 2283 432957 1179 0.0027 2538 436064 1139 0.0026

Oyster bank-B 1901 431546 962 0.0022 1823 435440 986 0.0022

Oyster bank-C 1278 437509 272 0.0006 2528 429309 1135 0.0026

Oyster bank-D 1614 437228 1065 0.0024 1284 439315 1148 0.0026

Oyster bank-E 2004 422733 1084 0.0025 2016 434381 1035 0.0023

Green sand-A 1443 422963 1019 0.0024 1949 429709 1032 0.0024

Green sand-B 2961 414915 268 0.0006 2007 437407 1048 0.0023

Green sand-C 1457 434712 391 0.0008 2662 432699 1145 0.0026

Green sand-D 1656 433026 574 0.0013 1855 428302 1050 0.0024

Green sand-E 5815 426727 246 0.0005 2941 412086 1006 0.0024

Coquina bed-A 1526 362214 159 0.0004 1549 436880 902 0.0020

Coquina bed-B 3066 421213 1226 0.0029 2748 431424 1186 0.0027

Coquina bed-C 2572 427396 1222 0.0028 3614 429799 1187 0.0027

Coquina bed-D 1619 433802 213 0.0004 1587 431171 982 0.0022

Coquina bed-E 10684 421952 1139 0.0026 4647 434182 1103 0.0025

Coquina bed-A 4468 378224 1048 0.0027 4679 433299 1105 0.0025

Coquina bed-B 2700 425234 281 0.0006 2715 426869 917 0.0021

Coquina bed-C 2732 432049 1131 0.0026 1810 432364 1078 0.0024

Coquina bed-D 1895 437006 1067 0.0024 1911 437144 1029 0.0023

Coquina bed-E 2410 434798 1087 0.0024 2670 434286 1114 0.0025

Coquina bed-A 2286 437975 1093 0.0024 1025 435397 1039 0.0023

Coquina bed-B 4697 407668 1247 0.0030 3143 432707 1014 0.0023

Coquina bed-C 2758 432657 1380 0.0031 1676 440038 1133 0.0025

Coquina bed-D 3311 398913 1164 0.0029 54 Tidigare skrifter i serien 336. Nilsson, Lawrence, 2013: The alteration ”Examensarbeten i Geologi vid Lunds mineralogy of Svartliden, Sweden. (30 universitet”: hp) 337. Bou-Rabee, Donna, 2013: Investigations 323. Gabrielsson, Johan, 2012: Havsisen i of a stalactite from Al Hota cave in Oman arktiska bassängen – nutid och framtid i and its implications for palaeoclimatic ett globalt uppvärmningsperspektiv. (15 reconstructions. (45 hp) hp) 338. Florén, Sara, 2013: Geologisk guide till 324. Lundgren, Linda, 2012: Variation in rock Söderåsen – 17 geologiskt intressanta quality between metamorphic domains in platser att besöka. (15 hp) the lower levels of the Eastern Segment, 339. Kullberg, Sara, 2013: Asbestkontamination Sveconorwegian Province. (45 hp) av dricksvatten och associerade risker. 325. Härling, Jesper, 2012: The fossil wonders (15 hp) of the Silurian Eramosa Lagerstätte of 340. 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