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Faisal Javed IqbalDissertations in Geology at Lund University,Master’s thesis, no 373(45 hp/ECTS credits)
Department of Geology Lund University
2013
Paleoecology and sedimentology of the Upper Cretaceous (Campanian), marine strata at Åsen, Kristianstad Basin, Southern Sweden, Scania
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
2
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).
3
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
4
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).
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]
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.
5
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
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]
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.
6
1 Introduction
Lithological changes and changes in the fossil fau-
na are generally indicative of changes in the environ-
ment. The change in diversity and body size of benthic
and planktonic invertebrate fossil fauna enables the
study of the changes of sea water conditions, organic
influx and sea-level changes (Vajda & Wigforss-
Lange 2006).
During the Late Cretaceous there were repeated epi-
sodes of sea transgression in southern Sweden (Vajda
& Solakius 1999), which resulted in the development
of an archipelago environment and irregular coast line
morphology of the Kristianstad Basin (Christensen
1984). The Upper Cretaceous Campanian deposits at
Åsen, within the Kristianstad Basin in northeast Skåne
consists of marine unconsolidated quartz sand with fair
amount of glauconite (Fig. 1). These sediments yield a
diverse vertebrate fauna including 34 shark species,
five ray species, six mosasaur species, one plesiosaur
species, and minor dinosaur and turtle remains (Table.
1) (Sorensen et al. 2013). The succession also contains
an invertebrate fauna including bivalves, cephalopods,
gastropods, brachiopods, bryozoans, echinoderms and
belemnites (Erlström & Gabrielson 1992; Sorensen &
Surlyk 2010, 2011; Sorensen et al. 2011, 2012).
The main aim of the project is to give an overview of
the total fauna and ecology at Åsen during the Late
Cretaceous. A further aim is to learn the paleontologi-
cal and sedimentological techniques that are used to
study the paleoenviroments; i.e. identification, sorting,
weighing and counting of the fossil fauna from B.
balsvikensis and B. mammilatus zones to investigate
the changes in fossil content in different beds of the
Campanian marine strata. The purpose is also to make
a stratigraphical illustration of the fossil fauna within
the B. balsvikensis and B. mammilatus zones. XRF
analysis of the sediment samples is used to study the
geochemistry of the different beds and XRF analysis
of the belemnites is used to study the variation of the
Sr/Ca between different beds in the sequence. SEM
and binocular microscope is used to study the charcoal
fragments. This study further aim to elucidate changes
in the invertebrate fauna through time, from early to
late Campanian in the Kristianstad Basin providing
additional information on paleotemperatures and sea
level changes.
2 Background
2.1 The Paleogeography of the Cretaceous
World
A super-continent Pangea was formed by the coales-
cence of all pre-existing continent masses during the
late Palaeozoic until Triassic. The southern part of
Pangea is called Gondwana and the northern part is
called Laurasia. The super-continent that started to
break up during the Triassic continued to break up
further during the Creatceous (145-66 Ma) (Gale 2000;
Vajda– Wigforss-Lange 2009). The Cretaceous world
was different from our modern world in several as-
pects, for example by the distribution of the continents
and geography. The period was characterized by ex-
tensive plate movements resulting in extensive rifting
and formation of mantle plumes. The extensive vol-
canism resulted in high concentrations of carbon diox-
ide (CO2), which might be the probable cause of the
global greenhouse conditions (Vajda et al. 2003), with
no polar ice caps (Gale 2000). The sea level-rise dur-
ing the Cretaceous was accompanied by massive struc-
tural changes globally due to plate tectonics. The unu-
sual conditions had profound effects on the initial radi-
ation, diversification and evolution of the flora and
fauna during the Cretaceous (Gale 2000; Friis et al.
2011; Ocampo et al.. 2006).
2.1.1 Plate Tectonics and Volcanism
During the Cretaceous, major changes took place in
7
the configuration of the plates, which resulted in
changes of the paleogeography and affected the carbon
cycle. The major plate-tectonic events included the
continued breakup of the Gondwanan continent
through the Jurassic and the rifting between Africa and
South America during the Early Cretaceous
(Valanginian-Aptian) interval (Kennedy & Cooper
1975). Development of the passage between the north
and south Atlantic during late Albian is evidenced by
the presence of ammonite fauna (Kennedy & Cooper
1975). The rapid opening of the South Atlantic Ocean,
and also the separation and the movement of India
northward took place during the Campanian-
Maastrichtian interval. The collision of India with Asia
also took place at the very end of the Cretaceous
(Jaeger et al. 1989). The rifting between Antarctica
and Australia started at about 132 Ma and the format-
ion of the volcanic centers due to the development of
the Indian Ocean by extensive rifting (Muller et al.
1993; McLoughlin 2001). The extensive spreading of
the Pacific Sea floor caused the development of the
terrains on the Pacific’s border during the mid-
Cretaceous time (Vaughan 1995). During the Cretace-
ous substantial mountain building took place. The sub-
duction and accretion of the northern part of the Tet-
hys throughout the Cretaceous resulted in development
of the Himalayas-East Alpine Orogeny. Also the ob-
duction of Ophiolites over continental crust of Arabian
shield on the southern part of Tethys took place during
the mid-Cretaceous to Campanian (Churkin 1972;
Gale 2000). The uplift of the Andeas and the Rocky
Mountains in America and uplift of the South Chinese
block in Asia were the major orogenic events during
the Late Cretaceous (Gale 2000). The four major Cre-
taceous basaltic flood eruption events took place
during the Cretaceous: 1. The Parana-Etendeka traps
in Brazil during earliest Cretaceous, 2. The Aptian
basalts in Rajmahal eastern India. 3. The Cenomanian
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).
8
basalts of Madagascar and 4. the Deccan traps in India
during the lates Cretaceous and ealiest Paleocene
(Muller et al. 1993).
2.1.2 Climate and sea level changes
The paleogeographical changes during the Cretaceous
were not only due to the plate movements but also
fluctuations in the global sea-level, which was pro-
bably driven by the global tectonic events.(Willumsen
& Vajda 2010) The sea levels during the earliest Cre-
taceous (late Volgian-early Berriasian) were marked
by lowstand (Rawson & Riley 1982) but this was
followed by the Early Cretaceous rapid sea level rises
with little interruption during the Valanginian and
Hauterivian and followed by a regression in the Barre-
mian. During the early Aptian, sea transgressed again
but sea level fell briefly, and then followed by the ove-
rall rise of the sea level during the Cenomanian. The
sea level remained high during the early Turonian, but
dropped during the mid-late Turonian, however at the
end of the Turonian and until the start of the Santo-
nian, a major transgression was seen, which resulted in
the formation of chalk facies (Gale 2000). The inter-
pretation of the global sea-levels for the late Campa-
nian is interpreted differently by different authors; one
show moderate and other show high sea levels i.e.
America and Europe respectively (Haq et al. 1987;
Hardenbol et al. 1998). Sea-level dated from
Maastrichtian indicate a major transgression globally
(Gale 2000; Ramberg et al. 2008). The sea transgres-
sed the vast shelf areas and spread the marine sedi-
ments far beyond the previous area (Hancock &
Kauffman 1979).
Most of the Mesozoic was a Greenhouse world with
no icecaps (Vajda 2001, 2008) and although there was
possibly ice on the poles during the Early Cretaceous,
the poles were ice-free and covered with temperate
forests during the middle and Late Cretaceous (Gale
2000; Vajda & McLoughlin 2005). The hot and humid
climate during the Cretaceous was a result of a combi-
nation of various factors for example the low relief and
even distribution of the continents around the globe,
warm and saline deep-sea oceans and that the conti-
nental interiors were covered with shallow seas (Gale
2000). The carbon dioxide levels were very high
which were attributed to outgassing from magmas
during the fast sea floor spreading, which probably
contributed to the warm climate (Harald & Olaussen
2008). Moderate temperatures prevailed at the poles
with a low temperature gradient from the equator to
the poles. Due to these facts, water masses transported
from the north were not cold and saline enough to sink
and form bottom waters (Gale 2000; Friis et al. 2011).
There was no major current flow from the poles and
no absorption of oxygen in the water over large areas
of the oceans. Combination of all these factors shows
that the average global temperatures during the Creta-
ceous were substantially higher compared to today
(Harald & Olaussen 2008).
2.2 Fauna of Late Cretaceous
2.2.1 Terrestrial fauna
The Cretaceous life on land was dominated by the di-
nosaurs which includes the well-known carnivore Ty-
rannosaurus rex. The sauropods, gigantic Jurassic
herbivore dinosaurs were joined by the ornithischians
during the Early Cretaceous. Also the foot prints of the
Iguanodon marks the presence in the Lower Cretace-
ous successions (Lloyd et al. 2008). The diversificat-
ion of new herbivore dinosaurs such as Hadrosaur,
Neoceratopsians, Ankylosaurs and the carnivorus di-
nosaur group including the giant Carcharodonttosauri-
nes and smaller Troodontids, Dromaeosaurs and
Omnithomimosaurs took place during the Late Creta-
ceous (Lloyd et al. 2008). The atmosphere was domi-
nated by the pterosaurs (flying reptiles) during the
Cretaceous. Diverse bird’ faunas were present in sizes
varying from finch to Ostrich-like forms. The develop-
9
and echinoderms were also present during the Cretace-
ous (Harald & Olaussen 2008). The vertebrates of the
Cretaceous seas were dominated by sea turtles,
Ichthyosaurs (fish lizard), crocodiles, sharks, rays and
the giant marine reptiles, the mosasaurs and the plesio-
saurs (Harald & Olaussen 2008; Lloyd et al. 2008). In
the fresh water enviroments ostracodes and algae, such
as Botryococcus flourished during the Cretaceous
(Guy-Ohlson 1992).
2.3 North America – Western Interior
Seaway during the Campanian
During the Late Cretaceous much of North America
was covered by a large inland sea, which was at its
maximum limits stretching from the Artic Ocean to the
Gulf of Mexico during the late Albian to Maastrichtian
(Kauffman 1967, 1977; Zangrel 1960; Coban 1969;
McNeil & Caldwell 1981; Bercovici et al. 2009,
2012). The development of the Western Interior
Seaway started by the extension of the southern em-
bayment of the Arctic Ocean throughout the Cretace-
ous (Friis et al. 2011). The marine strata of the Upper
Cretaceous host large amounts of fossil fauna in-
cluding invertebrates such as mollusks, brachiopods
and bivalves (Nicholls & Russell 1990). Vertebrates
include, amongst others, mosasaurs, plesiosaurs, turt-
les, pterosaurs, birds, fish, and sharks. The climatic
conditions during the Early Cretaceous were dry based
on the low amount of the terrestrial organic matter in
the sediments (Hallam 1984, 1985) signifying sparce
vegetation. Similarly the presence of high organic con-
tent in the sediments suggests a temperate humid cli-
mate in North America during the mid-Cretaceous
(Gale 2000). The high presence of charcoal (possibly
from wildfires) in the sediments from the Late Creta-
ceous of North-America suggests dry climatic environ-
ment (Friis et al. 2011).
The Western-Interior Seaway divided North America
ment and evolution among the vertebrates, like the
Squamates, Crocodilians and other reptiles took also
place during the Cretaceous. The development of the
foetus in the placental mammal’s womb and the ap-
pearance of the marsupials, which developed exter-
nally in the mother’s pouch, took place during the Cre-
taceous. The mammals during that time were mostly
insectivores, but also some herbivores and omnivores
species were present (Gale 2000; Friis et al. 2011;
Lloyd et al. 2008; Erlström & Gabrielson 1992).
2.2.2 Terrestrial flora
The Early Cretaceous flora was partly similar to that of
the Jurassic, which included ginkgo-related species,
bennettitaleans, cycads, ferns and seed-ferns (Vajda
2001; McLoughlin et al. 1995). However there was a
big difference, the angiosperms radiated during the
Cretaceous with the appearance of the modern oak,
birch, magnolia (Friis et al. 2011). The explosion of
the angiosperms by replacement of the gymnosperm
and ferns provided new radiation for pollinating
insects, moths and butterflies (Friis et al. 2011). Also
the conifer taxa Sequoia which played important role
during the Cenozoic, appeared during the Cretaceous
(Harald & Olaussen 2008; Lloyd et al. 2008).
2.2.3 Marine fauna
In the sea, the coccolithophorid algae flourished, these
produced vast amount of chalk. Other organisms
occurring in the Cretaceous marine environments were
the foraminifera and the silica sponges and the plank-
tonic foraminifera Globigerina, which provided the
ooze that deposited in the deep sea (Erlström & Gabri-
elson 1992). Ammonites and bivalves were also com-
mon in the seas. The Cretaceous marine organisms
developed some peculiar morphologies: Baculites and
Macroscaphites developed a linear elongated shell and
an open spiral shell respectively. Sea urchins, rudists,
bivalves, bryozoans, brachiopods, belemnites, corals
10
into several faunal provinces based on the composition
of the invertebrate faunas such as gastropods (Sohl
1967, 1971), cephalopods (Jeletzky 1968, 1971), bi-
valves and ammonites (Cobban & Reeside 1952; Gill
& Coban 1966; Kauffman 1977; Nicholls & Russell
1990).
The fossil flora along the Western Interior Seaway is
represented by leaves, flowers, pollens, stems and
roots and reveals the presence of ferns, cycads, coni-
fers and angiosperms. The Albian flora, in particlular,
provides important information about the early radi-
ation and diversification of the angiosperms during the
mid-Cretaceous (Friis et al. 2011).
2.4 Europe during the Campanian
During the Late Cretaceous, most of Europe was cove-
red by shallow epicontinental seas. Shallow marine
continental sediments of Campanian age are present in
Northwestern Europe (Harald & Olaussen, 2008; Gale
2000). The shallow sea covered an area from the Bri-
tish Isles to the eastern Caspian Sea and from the
Fennoscandian Craton in the north to the Alpine fold
in the south (Harald & Olaussen 2008; Gale 2000).
The development of the Alps in the south, and
spreading of the North Atlantic oceanic crust and in
the west, were probable cause of the intensive volca-
nism and earthquake in the central parts of the paleo-
Europe. During the Campanian the sea-level reached
to its maximum, over 100m higher compared to pre-
sent sea-level (Ziegler 1990). The intense sea-level
rise linked the southern warmer waters with northern
colder waters, which resulted in lowering the water
temperatures in western and central Europe (Harald &
Olaussen 2008). The connection between the seas to
the south and north resulted in the exchange between
the faunal provinces (Matsumoto 1973).
The Cretaceous sediments are present in two basins in
southern Sweden; in the Malmö Basin and in the Kris-
tianstad Basin (Moberg 1888). Both basins differ in
their sedimentological characteristics and lithology,
and in fossils content. Due to the tectonic activity
during the Santonian, which extended until the
Maastrichtian, Skåne was divided by the Romeleåsen
fault in southwest and the Linderödsås-Nävlinge-
Hallandsås ridge in northeast (Norling & Bergström
1987). Those tectonic activities probably ceased in the
northeastern part during the Campanian but continued
in the southwestern part. The sediments in the Malmö
and Kristianstad basins were deposited in a near shore
marine environment (Erlström & Gabrielson 1992).
2.4.1 Campanian marine fossil assembla-
ges in the Kristianstad Basin - Vertebrate
fauna
The major vertebrate fossil assemblages in the Campa-
nian strata of the Kristianstad Basin are sharks, turtles,
plesiosaurs, mosasaurs, rays and dinosaurs. They are
described below:
2.4.1.1 Sharks
38 predatory shark species have been identified from
the Kristianstad Basin in the lower Campanian strata
(Siverson 1992a, b, 1993, 1995; Rees 1999; Sorensen
et al. 2012). All of the identified species predominant-
ly fed on bony fishes, cephalopods and invertebrates
(Siverson 1992a). Shark species such as Cretoxyrhina
mantelli and Squalicorax kaupi (Agassiz 1843) have
played important role in the marine ecosystem as they
occupy the top of the food web (Shamada 1997; Shi-
mada & Cicimurri 2005). Teeth from large nektonic
shark species embedded in the partially digested mo-
sasaur bones were found in sediments from Western
Interior Seaway of North America and shows that they
fed, or scavenged, on those giant reptiles (Everhart et
11
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).
12
al. 1995; Shimada 1997).
34 out of the 38 shark species encountered in North
America were identified from the Åsen locality
(Siversson 1992). All identified species were active
predators (Sorensen 2013). Some of the sharks from
Åsen fed on everything they encounter, whereas some
fed on the smaller fishes and invertebrates (Siverson
1992a). Carcharias aasensis (Siverson 1992a) and
Cretalamna appendiculata (Agassiz 1843) occupied
the top of the food chain (Siverson 1992a). The bite
marks of shark's teeth are common on reptile bones
from the Kristianstad Basin (Einarsson et al. 2010).
2.4.1.2 Mosasaurs
Mosasaurs were the giant reptiles that occupied the top
of the food web during the Late Cretaceous in the seas
(Russel 1967). Six mosasaur species have previously
been identified from Kristiansand Basin in the upper
lower Campanian strata (Lindgren & Siverson 2002,
2004, 2005; Lindgren 2004, 2005a). Mosasaurs were
carnivores and primarily fed on fish and cephalopods
(Lindgren et al. 2010). All six mosasaur species were
found in the lower Campanian strata at Åsen
(Lindgren 2004, 2005a, b). The largest mosasaur
living in the shallow sea of the Kristianstad Basin was
Tylosaurus ivoensis (Person 1963). The length of its
lower jaw reached 1.5 m (Lindgren 2004). The broken
tooth crowns at the apex of this species suggest that it
might have fed on fleshy animals (Lindgren 2004).
Clidates propython (Cope 1869) and Eonatator stern-
bergi (Wiman 1920) are both small sized mosasaurs
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
the smaller mosasaurs from larger predators (Lindgren
& Siverson 2002, 2004).
2.4.1.3 Plesiosaurs
Six plesiosaurs species have been identified so far
from the lower Campanian of the Kristianstad Basin
(Person 1959, 1963, 1990). Plesiosaurs were reptiles
that were adapted to the marine realm, however they at
times sought shelter in non-marine environments to
protect their juveniles (Vajda & Raine 2010). Po-
lycotylida was a carnivorus plesiosaur genus with short
neck and large, elongated head. The Elasmosurus were
instead long necked, short headed and piscivorus
(Massare 1987). These were free swimmers and their
body could reach up to 14m in length, and weigh up to
several tons (Massare 1988). They mostly fed on small
fishes and cephalopods (Massare 1987; McHenry et al.
2005). However, some of them were also benthic gra-
zers, fed on the invertebrates and used gastrolites to
help digesting the food (McHenry et al. 2005; Taylor
1987). So far a polycotylid plesiosaur (Einarsson et al.
2010) and an elasmosaurid plesiosaur Scanisaurus
have been identified from Åsen (Persson 1959).
2.4.1.4 Turtles
Turtle remains are found at almost every locality in the
Kristianstad Basin; however, only two turtle taxa have
been identified (Persson 1959, 1963; Scheyer et al.
2012). The turtle’s diet is poorly known, however bi-
valve remains are found in protostegid turtle’s body
cavity from Australia in Lower Cretaceous deposits
(Kear 2006). The large amount of turtle’s remains sug-
gests that they were abundant in shallow seas
(Sorenson et al. 2013). The remains of the turtles were
also found in the Camapanian strata from Åsen
(Soresson et al. 2013).
2.4.1.5 Rays
Totally six ray species have been identified in the
Campanian strata from the Kristianstad Basin, all be-
longing to the Rhinobatoidae (Siverson 1993). They
13
were nektobenthic carnivores, which mostly fed on
invertebrates and small fishes (Last & Stevens 2009).
Five of the Rhinobatoidae species were encountered in
the Campanian sediments at Åsen, whereas rays are
rarely found from the other localities in the Kristian-
stad Basin, indicating that they preferred more murky,
estuarine waters as compared to their modern counter
parts (Last & Stevens 2009).
2.4.1.6 Dinosaurs
Leptoceratopsid dinosaurs have been identified based
on the teeth remains from Åsen (Lindgren et al. 2007).
The Leptoceratopsid is a small sized and horned, her-
bivore dinosaur, which mostly fed on cycads, conifers
and ferns (You & Dodson 2004).
2.4.1.7 Coprolites
Coprolites are fossilized feces and provide a source of
information about the diet, digestive physiology and
trophic levels of the paleoecosystems (Hunt et al.
1994; Eriksson et al. 2011). Coprolites can be distin-
guished and identified based on composition as well as
the morphology. They can vary in shape, size and
composition. Systematically spiral shaped coprolites
are linked to fish (Häntzschel et al. 1968). The irregu-
lar unspiraled shaped coprolites have generally been
linked to dinosaurs and may be related to mosasaurs
and plesiosaurs then collected from marine sediments
(Månsby 2009; Eriksson et al. 2011; Thulborn 1991).
The size of the coprolites may vary. For example the
coprolite of a theropod dinosaur may reach up to 40
cm in length and 15 cm in width (Chin et al. 1998).
Two groups of lamniform sharks have been identified
from Åsen based on coprolites. The coprolites were
linked to macrophagous sharks, based on the morpho-
logy (a heterotropic mode of spiraling). The other type
of coprolite, containing mollusk were interpreted to be
produced by durophagous sharks. Further, large unspi-
ralled coprolites were interpreted as related to mo-
sasaurs and plesiosaurs (Månsby 2009; Eriksson et al.
2011).
2.4.2 Campanian marine fossil assembla-
ges in the Kristianstad Basin - Inverte-
brate fauna
2.4.2.1 Belemnites
Belemnites are an extinct group of marine cephalopods
whose ‘‘guard’’ the calcitic body part is mostly found
in Jurassic and Cretaceous deposits. They had ten arms
like the modern squid which helped them to swim
(Stevens 1965; Kröger et al. 2011). The body of the
belemnites were divided into three parts; the posterior
ostrum (guard), the middle phragmocone and the front
pro-ostracum (Saelen 1989; Doyle 1990). Belemnites
had an important position in marine realms as predator
on smaller organisms and as prey for large marine rep-
tiles (Doyle & Macdonald 1993; Cicimurri & Everhart
2001; Rexfort & Mutterlose 2006). Several species and
sub species of the belemnites were identified be-
longing to the genera such as Actinocamax ( Miller
1823), Gonioteuthis (Bayle 1879), Belemnitella
(d’Orbigny 1840), Belemnellocamax (Naidin 1864)
and Belemnella (Nowak 1913) from the Campanian
strata of the Kristianstad Basin (Christensen 1975).
Belemnites secrete different parts of calcareous shells
in different water depths, therefore each secreted part
shows temperature of an area where it has grown, and
the whole belemnite reflects the average temperature
of the area where it grew (Spaeth et al. 1971). The
δO18 values of belemnite rostra show that the mean
temperature during the Campanian in the Kristianstad
basin was 20˚C to 24˚C (Lowenstam & Epstein 1954).
The diversity changes of the belemnites are related to
the changes in the Earth’s environment such as sea-
level changes, climatic changes and mass extinction
events (Christensen 2002).The belemnites got extinct
at the K-Pg boundary along with the ammonites
14
(Russel 1979; Alroy et al. 2008). From Åsen, belemni-
tes of the genus Belemnellocamax are used as key-taxa
for the uppermost lower Campanian and the lowermost
upper Campanian representing the Belemnellocamax
mammilatus and Belemnellocamax balsvikensis
respectively (Christensen 1975). B. balsvikensis differ
from B. mammilatus by its deep alveoli (Brotzen
1960). Also the presence of conellae along with the
white layer in B. balsvikensis distinguishes it from B.
mammilatus (Christensen 1975).
2.4.2.2 Oysters
Oysters are bivalves that are found in coastal waters,
and also in estuarine areas. They are attached to the
substrate such as stones, pilings, and mangroves. The
oyster Acutostrea incurva was identified from Åsen in
Campanian strata (Nilson 1827). The oysters from
Åsen have thin shells with common fragile imbricat-
ions and xenomorphic structures (Sorensen & Surlyk
2008). On the right valve the Oysters have mostly xe-
nomorphic longitudinal cylindrical ornamentation and
on the left side they have attachment imprint, which is
attributed to attachments to the deciduous trees in the
mangrove (Christensen 1975). Many oyster valves are
found together with complete shells in the Campanian
sediments (Sorensen & Surlyk 2008).
2.4.2.3 Brachiopods (ex crania)
Brachiopoda are invertebrates that have sedentary fee-
ding behavior and have great resemblance with the
mollusks (Rudwick 1970), however; their body axes,
gill, foot and gonads are different from the mollusks
(Pennington & Striker 2001). Several brachiopods
species have been identified from the Kristianstad Ba-
sin (Lundgren 1885; Hägg 1947) amongst those the
genera Crania (Carlsson 1958) and Terebratula
(Hadding 1919). Also the inarticulate brachiopod
Isocrania ignabergensis has been identified. This is a
thin-shelled species, characteristic of higher energy
inner shelf environments (Erlström & Gabrielson
1992). Additionally, findings of Crania craniolaris
(Linnaeus 1958) and the ichnogenus Podichnus have
been identified from Åsen (Bromley & Surlyk 1973;
Sorensen & Surlyk 2008).
2.4.2.4 Inoceramids
Inoceramids belong to a bivalve family that appeared
in the Permian and became extinct at the end of the
Cretaceous (Hilbrecht & Harries 1992). They were
dominant among the bottom communities especially in
the benthic oxygen restricted environments (Kauffman
& Harries 1992). Numerous inoceramid fragments
have been collected from the sediments of the Kristi-
anstad Basin (Lundgren 1974: Erlström & Gabrielson
1992).
2.4.3 Campanian flora in the Kristianstad
Basin - Fossil wood (Charcoal)
Fossil wood is found throughout the fossil record as it
is resistant to both biotic and abiotic factors. Charred
wood fragments provide essential insights into the
plants anatomy (Herenedeen 1991a, b; McLoughlin et
al 1995). However, the charcoalified wood is fragmen-
tary and brittle so the information acquired from the
charcoal about the host plant is incomplete (Friis et al.
2011). Charcoal has been found within the Kristian-
stad Basin and dated to late Santonian to early Campa-
nian based on megaspores (Koppelhus and Batten
1989) and also based on the overlain strata identified
to be of early Campanian age based on the presence of
Belemnellocamax mammillatus (Christensen 1975;
Friis et al. 2011). The initial radiation and diversificat-
ion of the angiosperms (flowering plants) took place
during the Early Cretaceous beginning from c. 135 Ma
(Friis et al. 2011). A world known Campanian
charcoalified flora from Åsen have been described by
(Friis & Skarby 1981). The assemblages contain wood
from angiosperms but also reproductive organs, lea-
15
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.
16
ves; along with conifers twigs (Friis et al. 2011).
About 100 taxa and 20 flowers species have been iden-
tified from the charred fossil plant assemblages from
Åsen (Friis et al. 2011 and references therein). The
presence of mostly same taxa in the lower and upper
part of the sequence shows that no hiatus occurs (Friis
et al. 2011). The charcoalified plant fragments along
with the lignitised compression fossils are present
throughout the succession; angiosperms flowers, seeds
and fruit fragments are a few millimeters in size but
conifers and twigs are found in comparativelty large
size (Friis et al. 2011). The larger specimens identified
from the locality are all conifers while the angi-
osperms identified are small fragments (Nykvist 1957;
Herendeen 1991a).
3 Geological Setting
The Kristianstad Basin is located in Skåne, southern
Sweden and has c. 200 m thick Cretaceous marine
strata exposed (Christensen 1984). During the Cretace-
ous period the basin was formed due to the southwest
dipping of the crystalline bedrocks, with faults for-
ming the Nävlingeåsen and Linderödsåsen horsts
(Bergström & Sundquist 1978). The northern part of
the Kristianstad Basin is irregular, having several Up-
per Cretaceous outliers exposed outside the basin and
on the south-eastern part of the basin lies the Hanö
Bay. However, the sedimentary bedrock extends into
the Baltic Sea (Bergström & Sundquist 1978). During
the latest Triassic to Middle Jurassic the temperature
was warm and humid, the weathering of the kaolinized
bedrock resulted in the development of the uneven
topography of the Precambrian bedrock (Lidmar &
Bergström 1982).
Four major transgressive pulses occurred during the
Cretaceous (Bergström & Sundquist 1978), which
covered the crystalline Precambrian rocks (that were
exposed at the time) and resulted in the development
of the archipelago environment and irregular coast line
morphology (Christensen 1984). The quartz rich sedi-
ment and kaolin of fluviatile origin are present in the
basal part of the sedimentary portion (Erlström & Ga-
brielson, 1992). During the latest early Campanian the
calcareous sandstone, conglomerates boulder beds,
calcarenites and calcisiltit were deposited (Christensen
1984). This corresponds to the local biozone
Belemnellocamax mammillatus Zone that can be corre-
lated to the Belmnitella mucronata senior/Genioteuthis
quadra gracilis Zone in Germany (Christensen 1975).
Åsen is located in the north-east part of the Kristian-
stads Basin and the sediments consist of marine sands
that rest on the lacustrine clay and argillaceous sedi-
ments. The Cretaceous marine stratum at the locality is
rich in macro-invertebrate fossils (Lundgren 1934;
Lindgren & Siverson 2002). Glacially tectonised, un-
consolidated sand overly the upper Santonian/ Campa-
nian lacustrine clayey argillaceous sediments (Friis &
Skarby 1981; Siverson 1992) and 3.5 m of these sedi-
ments are exposed at the Åsen locality. The sediments
in the exposed succession were sampled at different
levels. The rock unit is divided into two parts, the lo-
wer Campanian B. mammillatus Zone and the upper
Campanian B. balsvikensis Zone (Christensen 1975;
Lindgren et al. 2007).
The sediments of the B. mammillatus Zone consist of
sandstones with calcarenites and calcirudites and host
vertebrate bones and teeth, belemnites, oysters, corals
and coprolites. The coquina bed at approximately 1.5
m below the erosional layer in the B. mammillatus
Zone, consist of green, coarse quartz sand with belem-
nites, oysters, coprolites and vertebrate bones and
shark teeth (Rees 1999; Lindgren et al. 2007).
4 Materials and Methods
The samples were collected from different beds during
excavations in summer, 2010 - 2012 from the Campa-
17
nian successions at Åsen, southern Sweden (Fig. 3).
The fieldwork was performed by several volunteers
and students under the supervision of Elisabeth Einars-
son. The samples were first sieved in the field and the
larger fossil fragments were separated. The remaining
samples were collected in bags, got marked with the
name of the bed they were collected from, and with the
excavation date. The samples were processed by wet
sieving through a mesh size of 2,00 mm and were then
put into steel dishes and dried in the oven over night.
After the samples dried, the sediments were separated
from the fossils and were put in different plastic boxes
marked with sample names as designated on the plas-
tic sample bags. Different fossil fragments i.e. the
shells without any structure and the belemnites were
put in large plastic boxes whereas the shells with
structures, charcoal, corals, shark teeth, bone frag-
ments, were put in the same colored box sorted into
different segments (Fig. 2).
Among the sorted fossils the following groups were
identified and used for the diversity and paleoenviro-
ment study in this project; shark teeth, plesiosaur
tooth, gastrolits, charcoal, belemnites, shells,
inoceramids, sea urchins (spines and plates), bryozo-
ans, barnacles and coprolites. The sorted fossils were
weighted and put in a comprehensive excel data set.
Total amount of the sample along with fossils in per-
centages are presented in the Appendix I. The percen-
tages (based on weight) of the fossil and sediment
components from the different beds within the
sequence, are presented as pie charts (Fig. 4; Fig. 5;
Fig . 6)
4.1 XRF analysis of the sediment samples
In total 19 samples were collected from the different
beds and were crushed to powder for the XRF analy-
sis. A small amount was taken from each powdered
sample after making sure that the sample was homoge-
neous. The powder was placed in a sample cup with a
4 micron thick polypropylene film. The samples were
analyzed during 240 sec with a portable Niton XL3t
XRF analyzer at Department of Geology, Lund Uni-
versity. The data reduction was performed with the
program NDTREL-08. The analytical data from the
different samples are shown in the Appendix II.
4.2 XRF analysis of the Belemnites
Belemnites were also analysed with the XRF method.
Several belemnites were collected from different lay-
ers from the Belemnellocamax mammillatus Zone and
from the Belemnellocamax balsvikensis Zone. Diffe-
rent belemnite specimens from the same beds were
analyzed in order to obtain the Sr/Ca variations for
paleotemperature estimation. The phragmocone and
rostrum solidum of the belemnites were analyzed to
check the variation within single belemnites. Before
the analysis the belemnites were cut in halves, and the
cut surface was gently polished using a carborundum
disc. Some belemnites have an inorganic phragmocone
layer in the center surrounded by brown rostrum so-
lida. In most cases, the belemnite phragmocones and
the rostrum solida are formed by brown calcite. The
samples were analyzed in 240 sec with a portable Ni-
ton XL3t XRF analyzer at Department of Geology,
Lund University. The data reduction was done with the
program NDTREL-08. The data is presented as parts
per million (ppm). In this study the values of Si, Ca
and Sr were estimated (see Appendix III). The average
value is taken from all the belemnites of the same bed.
4.3 SEM Charcoal Analysis
The primary tool used to study the charcoal was the
binocular microscope as is a fast and inexpensive
method. The binocular microscope is used to sort the
charcoal from bones and sediments. Also the weathe-
red and well-preserved (=fine) charcoal fragments
were separated by using the binocular microscope.
Then to further anatomical study of the charcoal frag-
18
2. The middle ‘’Greensand layer” lies on top of the
Coquina bed which has a thickness of c. 0.5 m. The
bed contains glauconitic-rich quartz sand, including
fossils such as bryozoans, sea urchins, belemnites,
inoceramid fragments, charcoalified fragments, copro-
lites, shark teeth, bony fishes vertebrae, and bone frag-
ments.
3. The following uppermost ‘’Oyster bank’’ , named
based on the high oyster abundances has a thickness of
c. 0.6 m. The bed is predominantly represented by
calcareous chalk with an abundant oyster fauna. The
bed also contains other fossils fragments such bryozo-
ans, sea urchins (plates & spines), micrabacia, incore-
mids and coprolites.
4. The lower ‘’Balsvikensis Green’’ named after the
green coloured glauconitic-rich sand with a thickness
of c. 1.2 m. ‘’Balsvikensis Green’’ consists of gra-
nules, pebbles, carbonate cemented nodules and stein-
kerns. The fragmented fossils of the ‘’Balsvikensis
Green’’ including belemnites, inocermids, micrabacia,
barnacles, bryozoans, coprolites, shark teeth, bony
fishes vertebrae and tiny amount of charcoal frag-
ments.
5. The uppermost bed ‘’Balsvikensis Yellow’’ named
based on the yellow colored sand, is quartz-rich and
reaches a thickness of c. 1.6 m. In the base of this bed
a slightly whiter interval can be detected but is herein
included in ‘’Balsvikensis Yellow’’. This bed contains
granules, pebbles, carbonate cemented nodules and
steinkerns. ‘’Balsvikensis Yellow’’ further contains
fossil fragments of oysters, belemnites, bryozoans,
micrabacia, barnacles, sea urchins, inoceramids,
coprolites, gastrolites, shark teeth, plesiosaur tooth,
bony fishes vertebrae and small quantity of charcoal
fragments. Quaternary cover is present above the up-
per B. balsvikensis Zone.
ments, Scanning Electron Microscopy (SEM) techni-
que was used. Only the unweathered and well preser-
ved charcoal fragments were studied under SEM.
A Hitachi S-3400N SEM at Department of Geology,
Lund University. The samples were gold plated and
mounted on a 3mm diameter SEM stub. Imaging was
done using the secondary electron detector.
5 Results
5.1 Stratigraphy
The stratigraphy of the studied locality is mostly based
on belemnite zonations (Christensen 1975), and other
fossil groups such as oysters. The different beds are in
this study named based on the lithology and the colour
of the sediments. Fig. 4 shows the different beds wit-
hin the stratigraphical succession along with the fossil
content. The total thickness of the exposed marine
Campanian strata at the study site reaches about 4,15
m (Einarsson et al. in press). A total of 169 kg material
was sorted and a diverse fauna with coprolites
and charcoalified phytoclasts as an additional compo-
nent was identified. The studied succession is divided
into two zones, the lower B. mammilatus Zone of latest
early Campanian and upper, younger B. balsvikensis
Zone of earliest late Campanian age (Christensen
1975).
The lower B. mammilatus Zone is further divided into
three beds;
1. The lowermost ‘’Coquina bed’’ based on its content
of fossil shells with a thickness of c. 0.25 m. The co-
quina bed is a mottled storm deposit consisting of gre-
enish coarse sand, rich in fragmented oysters, belemni-
tes, inoceramids, micrabacia, coprolites, shark teeth,
bony fishes vertebrae and bone fragments. The bed
also contains charcoalified wood, both micro and
macro sized.
19
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.
20
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).
21
5.2 Results of the XRF analysis
(Sediments)
Analytical XRF data from the 19 sediment samples
from the Campanian succession at Åsen (Fig. 5, Table
6) is represented by Si, Ca, Fe, K, Al, P and Sr.
Silica (Si) and calcium (Ca) are the major elements
found in the analysis; Si and Ca show an inverse relat-
ionship through the whole succession. Si shows a
decreasing trend all through the succession with
highest values in the samples from the terrestrial flood
plain deposits and lowest values within the B. balsvi-
kensis Zone. Ca on the other hand shows the opposite
trend with highest values in the B. balsvikensis Zone.
Average Iron (Fe) content in the flood plain deposit is
slightly over 1% but it fluctuates a lot in the flood
plain sediments. Fe shows a decreasing trend all
through the succession with highest values in the
samples from the Coquina bed and lowest values wit-
hin the B. balsvikensis Zone.
Potassium (K) shows a decreasing trend all through
the succession with highest values in the samples from
the terrestrial flood plain deposits and lowest values
within the B. balsvikensis Zone, however K shows a
small peak in the Green sand bed.
Aluminium (Al) shows a decreasing trend all through
the flood plain deposit with highest values in the
samples from the lower part of the flood plain deposit.
Al shows no trend through the whole succession; ho-
wever Al shows a small peak at the boundary between
the B. mammilatus Zone and B. balsvikensis Zone.
Phosphorus (P) shows slightly high values in the flood
plain deposits as compared to the overlying B. mam-
millatus Zone. P shows an increasing trend in the two
zones, with lowest values in the samples from the B.
mammillatus Zone and highest values within the B.
balsvikensis Zone.
Stronitium (Sr) has slightly higher values in the flood
plain deposits as compared to the overlying B. mam-
millatus Zone. Sr shows an increasing trend in the two
zones, with lowest values in the samples from the B.
mammillatus Zone and highest values within the .
balsvikensis Zone
The following components occur as trace elements; Zr,
Ba, Zn, Th, U, Nb, Mo, Y and Rb in the 19 analyzed
samples and these elements occur in almost same pro-
portions throughout the succession (Table 6).
5.3 Results of the XRF analysis
(Belemnites)
The results from the geochemical measurements of the
belemnite rostra shows that the phragmocone have
consistently lower values of Sr+2
compared to the
rostrum solidum. The Sr+2 values detected from the
secondary calcite phragmocone reach 400 ppm to 650
ppm; whereas Sr+2 values from the primary calcite
phragmocone reach values 800 ppm to 900 ppm. The
Sr+2 values for the rostra solida range from 1000 ppm
to 1200 ppm (Table. 8).
As the results from the rostrum solidum is considered
more reliable, this study focuses on those results. The
Sr+2 values from rostrum solidum of belemnites from
the B. mammillatus Zone range between 1000 ppm and
1100 ppm and from the B. balsvikensis Zone between
1100 ppm and 1200 ppm.
The average Sr/Ca values and trend of the belemnites
from different beds of the Campanian strata (Table 2;
Fig. 5) show increasing average values of Sr/Ca from
the B. mammillatus Zone to the B. balsvikensis Zone
which indicate possibly lowering in paleotemperature
or diagenetic alteration.
22
Fig
. 5
. G
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23
5.4 Results of the SEM
The charcoal fragments were analyzed under the bi-
nocular microscope and were sorted into two datasets
grouped in weathered charcoal and fine charcoal. The
charcoal is weighted and graphs were drawn to see the
changes in the different layers. The images taken un-
der binocular microscope and SEM are used to identi-
fy the plants. The images were taken from different
angles to see anatomical structures of the wood. Most
of the analyzed fragments are gymnosperms but few of
them are angiosperms. Most of the identified gym-
nosperms are conifers e.g, pinaceae. From the ana-
lyzed fragments different cell vessels were identified,
that helped to identify what plant group they belong to
(Fig. 13, 14, 15, 16, 17 & 18).
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-
ges taken of the charcoal fragments were Transverse
Section (TS), Radial Longitudinal Section (RLS),
Tangential Longitudinal Section (TLS) and Stem Exte-
rior (Ext).
5.5 Results of the fossils
The following fossils are identified and included in
this project: shark teeth, plesiosaur tooth, gastrolits,
charcoal, belemnites, shells, inoceramids, sea urchins
(spines & plates), bryozoans, barnacles and coprolites.
The pie graphs show the overall decrease in the faunal
diversity in the B. balsvikensis Zone as compare to the
B. mammilatus Zone. The carbonate content in the B.
balsvikensis Zone is higher than in the B. mammilatus
Zone. The bar graphs for the charcoal, inoceramids,
coprolites, vertebrates’ bones and teeth show high pe-
aks in the B. mammilatus Zone; however the benthic
community like the micrabacia, barnacles, sea urchins
and bryozoan show high peaks in the B. balsvikensis
Zone.
6 Discussion
6.1 Fossil Fauna
The studied succession at Åsen comprise ca. 4,15 m
sediments and these are divided into two zones, the B.
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.
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
24
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.
G2
G3
G5
G4
25
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-
ionally the succession is divided in beds based on the
lithological and fossil content. The succession is divid-
ed into following entities:
1. The coquina bed is the lowermost bed within the B.
mammilatus Zone which is dominated by a variety of
fossils such as shell fragments, belemnites, charcoal,
coprolites, inoceramids, vertebrate bones and teeth. It
is a dark green glauconitic rich quartz sand bed.
i. The amount of the fossil fauna and the compo-
sition of the sediments suggest high energy,
inner shelf sea environment during the deposit-
ion of the coquina bed (G1, Fig. 6 ).
ii. The bed also contains high amount of charcoal
and wood fragments which might be reworked
from the underlying flood plain deposits or
derived from the terrestrial flora transported
from land into the marine basin (G1, Fig. 7).
iii. The amount of bone fragments, charcoal, shell
fragments and belemnites in small bed suggests
a storm deposit (G1, G2, Fig. 7).
iv. The presence of dark green glauconitic rich
quartz sand and siliceous preservation of the
fossils such as charcoal and shell fragments
0 0.2 0.4 0.6
Coquina Bed
Greensand
Oysterbank
Balsvikensis Green
Balsvikensis Yellow
Fine Charcoal Weathered Charcoal
0 20 40 60 80
Coquina Bed
Greensand
Oysterbank
Balsvikensis Green
Balsvikensis Yellow
Belemnites Shell fragments
0 0.05 0.1 0.15
Coquina Bed
Greensand
Oysterbank
Balsvikensis Green
Balsvikensis Yellow
Coprolites
0 0.5 1 1.5 2
Coquina Bed
Greensand
Oysterbank
Balsvikensis Green
Balsvikensis Yellow
Bony fishes vertebrae Shark teeth
Bone fragments
G1 G2
G3 G4
26
i. The fossil fauna and sediments content suggests
inner shelf environment because the charac-
teristics of the sediments develop the benthic
community (Gray 1981) (G2, Fig. 6).
ii. The more abundant micrabacia, barnacles and
sea urchins compared to the underlying coquina
bed may suggest more oxygenated bottom con-
ditions and also fairly clear waters or may be a
small episodic drop in sea level (regression)
(G5, Fig. 8).
iii. The decrease in coprolites suggests decrease in
the suspended food availability (G3, Fig. 7).
suggests low sedimentation rate in an open ma-
rine environment at the water and sediment
interface (Odin & Matter 1981; Vajda & Solak-
ius 1999). The sea water is under-saturated in
silica. The amount of coprolites, vertebrate bo-
nes and teeth also suggests high food availa-
bility and high sea levels for the vertebrates to
grow (G3, G4, Fig. 7).
2. Green Sand is the second layer of the B. mammi-
latus Zone and lies above the Coquina bed. It has a
higher diversity of the fauna compared to the lower
coquina layer (G2, Fig. 6).
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.
0 0.05 0.1 0.15 0.2 0.25 0.3 0.35
Coquina Bed
Greensand
Oysterbank
Balsvikensis Green
Balsvikensis Yellow
Bryozoa Sea urchin plates Sea urchin spines Barnacle Micrabacia InoceramidG5
27
3. The oyster bank is the upper most layer of the B.
mammilatus Zone and is dominated by fossil shell
fragments specially oysters. The overall diversity of
the fossils fauna increases slightly compared to the
lower Green sand bed.
i. The amount of glauconitic sand and the silice-
ous fauna suggest starvation of sediments due
to transgression (G3, Fig. 6)
ii. The bed contains high amount of oyster’s
fauna, which have attachment imprint, which is
attributed to attachments to wood, likely rafted
in from the terrestrial environment (Christensen
1975).
iii. The increase in vertebrate bones suggests high
food availability because increase in primary
productivity led to more resources for higher
animals in the tropic level (G2, G4, Fig. 6).
iv. The decrease in amount of coprolites might be
related to the presence of the amount of shark
teeth that also are decreasing. The presence of
high amount of bone fragments could be related
to the presence of big vertebrate animals that
are indirectly related to higher sea levels (G4,
Fig. 7).
4. The Balsvikensis Green is the lower bed within the
B. balsvikensis Zone of the early late Campanian. The
overall fossil faunal diversity is lower compared to the
Oyster bank and the silisiclastic component is higher.
(G4, Fig. 6).
i. The decrease in overall siliceous fauna, sug-
gests low primary productivity. This decrease
in primary production affected the abundance
and size of the fossil fauna. (G4, Fig. 6).
ii. The increase in amount of the benthic commu-
nity also supports the idea of drop in tempera-
ture and drop in sea level, which helped the
benthic communities to flourish (G5, Fig. 8).
iii. The higher amount of the belemnites compared
to the layers in the B. mammilatus zone is due
to a drop in sea level that created a shallow
marine environment (G2, Fig. 8).
iv. The lower amount of bone fragments and hig-
her amount of coprolites (from smaller sharks
and bony fishes) and shark teeth compared to
oysterbank could be due to that the big verte-
brate animals might moved to the deep waters,
this show the fall in water level (G3, G4, Fig.
7).
5. Balsvikensis Yellow is the uppermost bed within the
B. balsvikeinsis Zone and is dominated by the sedi-
ment content of carbonate cemented nodules and the
fossil content is dominated by the invertebrate fau-
naThe high amount of carbonate cemented nodules
suggests shallow water, semi lagoonal environment.
The high amount of carbonates may be due to either
fall in sea level drop in temperature or both (G5, Fig.
6 ).
i. The overall fossil diversity is half to that of the
Balsvikensis Green but mostly affected the pe-
lagic invertebrates and vertebrates which is also
an indication of fall in sea level that then
constrained them to move into open sea. It
could also be due to the decrease in the primary
productivity, lack of food, which constrained
them to move towards better food sources (G5,
Fig. 4; G5, Fig. 7).
ii. The presence of the benthic invertebrate com-
munity along with the carbonates also supports
the idea for drop in temperature because bent-
hic invertebrates collected from the bed are
28
cold water carbonate producers (Nelson 1988;
Rao 1997).
iii. The presence of steinkerns in the upper strata
within the B. balsvikensis Zone suggests drop
in sea level because the steinkern are internal
molds of an organism, that are formed by the
digenetic alteration of the unstable minerals in a
fauna and which were replaced by the calcite to
form carbonate shells in a carbonate rich envi-
ronment (Manning & Dockery III 1992) (G1,
Fig. 9).
6.2 XRF (sediments)
The high value of the Si in the non-marine flood plain
might be due to the aluminiumsilicate minerals due to
the fact that the lithology of the flood plain is mostly
fine clay. Similarly the value of Si is high in B. mam-
milatus Zone as compare to B. balsvikensis Zone, due
to the higher amount of glauconitic sand. The high Si
content in the glauconitic sand with high fossil content
also suggests low deposition and therefore there is lot
of fossils during the late early Campanian. The high Si
content also coincides with the abundant biotic life in
the B. mammilatus Zone as compare to the B. balsvike-
nis Zone. The dissolved skeletal Si from diatoms and
other silica utilizing organisms after their death en-
riches the deep water by settling through the water
column (DeMaster 1981). Due to starved sediments
the relative abundance of fossils increase within the
sediments. The low value of the Si in the B. Balsvi-
kensis Zone is may be due vice versa conditions both
for sediments and fossil fauna as compared to B. mam-
milatus Zone.
The calcium peaks are inversed to Si in the whole
sequence; the low Ca values in the flood plain can be
related to the lithology because it is terrestrial flood
plain. The low value of Ca in the B. mammilatus Zone
is related to the presence of the high siliciclastic fossil
content. The value of Ca in the B. balsvikensis Zone
are high as compared to the B. mammilatus Zone,
which can be due to the presence of carbonate cemen-
ted nodules in the B. balsvikensis Zone.
The K high values of the flood plain deposit fits quite
well because K is incorporates in the clay minerals
lattice due to it large size. Its moderate values in the B.
mammilatus Zone are related to the abundance of fos-
sil fauna presence because K is strongly adsorbed and
incorporated in organic life forms (Wedepohl 1978).
So the declining trend of K in the B. balsvikensis Zone
might be related to the low fauna diversity.
The high values of Al in the flood plain deposits fits
quite well because Al is essential metal of the alumini-
umsilsicate clay mineral. The amount of Al in the B.
mammilatus Zone and B. balsvikensis Zone is linear
because of the lithology that is composed mostly of
sand.
High Sr concentrations in the flood plain deposits is
related to the fact that Sr incorporates into the clay
minerals by the weathering of the source rocks. The
high values of the Sr in the B. balsvikensis Zone as
compare to the B. mammilatus Zone are due to the
high carbonate content in the B. balsvikensis Zone.
The Sr increasing trend in the upward part of the suc-
cession might be due to the paleotemperature, since Sr
concentration increases in the carbonates due to the
drop in temperature (Kinsman 1969).
6.3 XRF (belemnites)
The difference of Sr values in the secondary calcite
phragmocone and that of the primary calcite
phragmocone might be due to diagenetic alteration.
The phragmocone is composed of aragonite and arago-
nite is replaced by the calcite after a long deposition
(Kinsman 1969; Spaeth et al 1971). The distribution of
Sr in aragonite and calcite is ~8. There is a difference
29
between the Sr/Ca in the phragmocone and rostrum
solidum that might be due to biochemical fractionation
(Kinsman 1969).
The difference of the Sr concentrations between the
phragmocone and rostrum solidum may be due to the
fact that the belemnites developed different body parts
at different water depths. The phragmocone consists of
aragonite, and aragonite do not preserve well because
it is altered by genetic processes and converts it to
calcite. The rostrum solidum of the belemnite is the
well preserved part (Spaeth et al. 1971, 1971b;
Kinsman 1969). In this study the rostrum solidum is
used instead of the phragmocone since the XRF values
of the rostrum solidum are consistent whereas there is
a large fluctuations in the XRF values of
phragmocone.
The Sr values obtained from the rostrum solidum of
the belemnites from different beds of the Campanian
marine strata at Åsen coincide with the measured va-
lue calculated by Kinsman 1969. The Sr values in cal-
cite are in the belemnites from Åsen measured to
between 1000 ppm to 1200 ppm. According to
Kinsman (1969)” the Sr concentrations of calcite pre-
cipitated from sea water is about 1200 ppm” During
normal sea water conditions organisms deposit the
calcite at about 25°C and the Sr concentration in Cal-
cite would be 1000 to 1200ppm and at the same time
8200ppm in aragonmite. (Kinsman 1969). The Sr va-
lues from the rostrum solidum in the calcite of some
belemnites in the B. balsvikensis Zone is above 1200
ppm but as the aragonitic value has not been measured
in this study, the interpretations of sea temperature is
out of scope for this study.
The values of Sr/Ca based on the rostrum solidum of
belemnites from the B. mammilatus zone and B.
balsvikensis zone increase slightly i.e 0.0024 to
0.0027. There is a slight increase in Sr concentration
from B. mammilatus zone (1000-1100 ppm) to B.
balsvikensis zone (1100-1200 ppm).
6.4 Comparison between the two zones (B.
mammillatus and B. balsvikensis) at Åsen.
The fossil fauna comprises both vertebrate and inverte-
brate fossil fragments, and the sediment content sug-
gests inner shelf marine environment. The B. mammi-
latus and the B. balsvikensis zones are significantly
different in sediment and fossil content. The B. mam-
milatus Zone has an overall high diversity of fossils
compare to the B. balsvikensis Zone (Table. 3; Fig. 9).
6.4.1 Fossil fauna
The overall diversity of the fossil fauna decreases in
the upper B. balsvikensis Zone; however the amount of
benthic fauna increase in the B. balsvikensis Zone.
This is an indication of drop in sea level, which favo-
red the benthic communities to flourish (G5, Fig. 10).
The presence of abundant fossil fauna including verte-
brate remains and fossil shell fragments of bivalves
suggests high sea levels and high food availability
during the latest early Campanian of the B. mammi-
latus Zone. Also higher temperatures because of the
high primary productivity resulted in the overall faunal
diversity (G2, G3, G4, Fig. 10). The relatively high
abundance of charcoal in the latest early Campanian
strata may be a result of high erosion rate on land with
high influx of sediment and charcoal, another scenario
is that the charcoal was reworked from the lower flood
plain deposits (G1, Fig. 10). Further, the presence of
steinkerns in the upper strata of the B. balsvikensis
Zone suggests drop in sea level because the steinkern
are internal molds of an organism that are formed by
the digenetic alteration of the unstable minerals in a
fauna and which were replaced by the calcite to form
carbonate shells in a carbonate rich environment
(Manning & Dockery III 1992) (G1, Fig. 9). The
decrease in amount of the bone fragments of the B.
30
balsvikensis Zone suggests either drop in sea level or
indirectly decreas in primary productivity by the drop
in temperature. The increase of amount of coprolites
might be related to the amount of shark teeth in the B.
balsvikensis zone (G3, G4 Fig 10). The presence of the
benthic invertebrate community along with the carbo-
nates also supports the interpretation for drop in tem-
perature because benthic invertebrate collected from
the bed are cold water carbonate producers (Nelson
1988; Rao 1997).
6.4.2 Sediment content
The change in lithology and fossil fauna either its’s
diversity or size when comparing the B. balsvikensi-
sand B. mammilatus zones of Åsen suggests either
drop in sea level or drop in temperature, or both (Fig.
9. G1).The presence of sand on top of the non-marine
flood plain and no carbonates in the B. mammilatus
Zone suggests high sea level or transgression during
the late early Campanian. The litholog of the B. mam-
milatus Zone contains glauconite that together with the
presence of siliceous fossil fauna supports the idea of
high sea levels, and starved sediments. Whereas the
lithology of the B. balsvikensis Zone containing quartz
rich sand with high amount of carbonates together
with lower diversity and smaller size of the fossil
fauna suggests either drop in sea level, drop in tempe-
ratures or both (Fig. 7. G2).
6.4.3 Paleoecology
The lithology and high faunal diversity together with
the bigger size of the fossil suggests warmer paleotem-
peratures and high sea-levels during the latest early
Campanain time represented in the B. mammilatus
Zone. The larger body sizes together with the high
amount of the bone fragments of the large vertebrate
animals suggests high food availability for the preda-
tors and the prey. This high food availability supported
B. mammilatus
Zone
B. balsvikensis
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.
31
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.
the idea for the high primary production in the surface
waters, this high productivity help the pelagic commu-
nity to groom and diversify. Reduction in the body
size together with the decrease in the bones fragments
of the vertebrate animals suggests drop in sea-levels
during the earliest late Campanain time representing
by the B. balsvikensis Zone. Also the presence of the
benthic invertebrate community along with the higher
amount of carbonates also supports the idea for drop in
temperature since benthic invertebrate collected from
the earliest late Campanian strata are cold water carbo-
nate producers (Nelson 1988; Rao 1997).
7 Results and Discussion about
Charcoal
7.1 SEM Charcoal Images (Fig. 13, 14 &
15; Table. 4)
The charcoal fragment image a01, a02, a03, a04, a05,
a06 and a07 were referred to as conifer due to the pre-
sence of homogeneous tracheid and equal ray cells,
containing two to four circular cross-filed pits. The
charcoal fragment is highly weathered and compres-
sed, and may be transported from a distant source.
The charcoal fragment image b01 and b02 is from a
conifer based on the fact that the cells are homogene-
ous. The fragment is charcoalified wood and it is soft,
brittle and lightweight.
The charcoal fragment image d01 and d02 is referred
to a conifer because the cells are homogeneous, it may
have been deeply buried and transported because the
charcoal fragment is highly compressed, hard and he-
avy weight.
G1 G2
32
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.
0 0.05 0.1 0.15
Weathered Charcoal
Fine Charcoal
0 20 40 60 80
Shell fragments
Belemnites
0 0.05 0.1
Coprolites
0 0.5 1 1.5
Bone fragments
Shark teeth
Bony fishes vertebrae
0 0.05 0.1 0.15 0.2
Inoceramid
Micrabacia
Barnacle
Sea urchin spines
Sea urchin plates
Bryozoa
Legends
B. balsvikensis zone
B. mammilaus zone
G1 G2
G3 G4
G5
33
The charcoal image e01 and e02 is considered to be a
conifer because the cells are homogeneous. The frag-
ment was buried and transported because it is highly
weathered and compressed; the ray cells are much
larger and broader so it may be from a different spe-
cies than those mentioned earlier.
The charcoal in image f01, f02 and f03 are referred to
an angiosperm plant due to large vessel cells and small
parenchyma cells. The charcoal fragment is long trans-
ported, because it is highly weathered and compressed.
The charcoal fragment in image g01 is referred to a
conifer due to the presence of broad ray cells; it seems
to be transported because it is highly weathered and
shows strong burial compression.
The charcoal fragment image h01 is referred to a coni-
fer due to the presence of ray cells of uniform charac-
ter with circular cross-filed pits.
7.2 Binocular Microscope Charcoal Images
(Fig. 18, 19 & 20; Table. 5)
Img,
No scale View plant type comment
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
34
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
to be conifer wood due to the presence of homogene-
ous ray cells, the specimens may be charcoalified frag-
ments from a wildfire, because they are black, glassy,
soft, brittle and lightweight. The charcoal in images 16
-3, 16-4, 16-7 and 16-8 are conifers based on the pre-
sence of longitudinal and homogeneous ray cells. They
have brown compressed layers and may be lignified.
They are hard, compressed and heavy which might be
because they have been buried, lithified and transpor-
ted from a distant source. The charcoal in images 16-5
and 16-9 are referred to conifers, because the longitu-
dinal and ray cells are homogeneous. They have been
buried and transported because they are extremely
compressed, hard and heavyweight.
The specimens illustrated in 05-1, 05-4, 08-1, 16-13,
10-1, 10-2 and 22-1 are referred to conifer wood, ba-
sed on the fact that the longitudinal and ray cells are
homogeneous. They are lignitized, because they are
compressed, hard, heavyweight and are brown. The
images 05-2, 05-3, 10-3, 10-4 and 22-2 are also refer-
red to conifer wood; however they are burnt
(charcoalified) wood from wildfires because they are
soft, brittle, lightweight and glassy.
35
are mostly found in glauconitic sand deposits (Bell &
Goodell 1967; Tooms et al 1970; Giresse & Odin
1973). They are often produced by filter feeding org-
anisms in shallow water (Moore 1939). These are elo-
ngated cylinders with rounded edges having latitudinal
lines around the cylindrical body. Total lengths of the
all clustered cylinders are 5 mm and 1mm in width.
The size of the individual cylinder is 0.8 to 1 mm in
length and 0.1 to 0.3 mm in width. Further studies are
neede in order to link them to an organism.
9 Conclusion
A marine succession of Campanian age from
Åsen, Kristianstads Basin, southern Sweden
was studied bed by bed to describe and assess
the diversity and abundance of the fossils as-
semblages by study the fossil and sediment
content. The studied Campanian marine strata
at Åsen is divided into two zones, the latest
early Campanian B. mammilatus Zone and the
earliest late Campanian B. balsvikensis Zone.
The B. mammilatus Zone is further divided into
several beds: the Coquina bed, the Green sand
7.3 Results and discussion about the flo-
wer/sedges/equisetum (Table. 4; Fig. 11)
The charcoal fragment image c01, c02 and c03 is ten-
tatively referred to as Equisetum stem node, because
the stem of Equisetum in early development has leaves
that form a whorl around the stem at each node. Anot-
her possibility, however more unlikely is that this fos-
sil represents a Sedge, Sedge have blade-like leaves
that whorl around the stem attached to nodes in the
early development but when they grow up the leaves
detach the blade from the stem leaving exposed nodes.
8 Discussion about the eggs/fecel pellets/
small coprolites (Fig. 12)
A small cemented cluster of cylindrical shaped bodies
were found in the sediments from the B. mammilatus
Zone. The image Fp-1 and Fp-2 referred to as eggs at
the first sight, but their close morphological study un-
der the binocular microscope suggests them to be
faecal pellets. Faecal pellets are argillaceous matter
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).
36
bed and the Oyster bank. The B. balsvikensis
Zone was divided into the beds; Balsvikensis
Green and Balsvikensis Yellow.
The rich fossil fauna including brachiopods,
bivalves, belemnites, bryozoans, barnacles, sea
urchins and corals with fair amount of copro-
lites, vertebrate bones and teeth, indicates that
the area was a well-functioning ecosystem dur-
ing latest early and earliest late Campanian.
The sediment content and the fossil fauna sug-
gest an inner shelf environment.
The high amount of vertebrate bone fragments
together with the decreasing benthic communi-
ty suggests high sea levels during the latest
early Campanian representing by the B. mam-
milatus Zone.
The high amount of charcoal in the sediments
belonging to the B. mammilatus Zone suggests
high erosion rate on land with high influx of
sediments, or it may be reworked from the low-
er flood plain deposits.
The presence of the Steinkern, the decrease in
amount of the vertebrate bone fragments and
increase in amount of the benthic communities
including micrabacia, barnacles, sea urchins,
and bryozoas, along with the high amount of
carbonate content of the deposits of the B.
balsvikensis Zone suggests drop in sea level
during the earliest late Campanian.
The smaller size of the fossil fauna and pres-
ence of the cold water carbonate producing
fossil fauna from the B. balsvikensis Zone sug-
gest drop in water temperatures during the ear-
liest late Campanian at Åsen.
10 Acknowledgement
First I want to thank my Supervisor Vivi Vajda. I ap-
preciate all her contributions of time, ideas, and guid-
ance to make my Master’s thesis project productive
and stimulating. I am also thankful for the excellent
example she has provided as a successful woman geo-
scientist and professor. I would also like to express my
sincere gratitude to my Co-supervisor Elisabeth
Einarsson for the continuous support during my Mas-
ter’s thesis project, for her patience, motivation and
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).
37
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.
38
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44
a01
a04 a03
a06 a05
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).
45
a07 b01
b02
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).
46
e01
e02
f01
f02
g01
f03
h01
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).
47
16-1 16-2
16-3
16-4
16-7 16-5
16-9
16-8
16-4
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.
48
05-1 05-3
05-2
05-4 08-1
16-13
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.
49
10-1 10-2
10-3
10-4
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
50
Coquina
Bed Greensand Oysterbank Balsvikensis Green Balsvikensis Yellow
Total sample (total amount
sieved sand in kg) 282 501 783 1239 4387.5
Sieved sand (less than 2mm) 276.08115 498.11885 774.24259 1229.03893 4329.91271
Granules & Pebbles (2-64
mm) 2.46149 0.56579 3.02728 4.495 0.15633
Carbonate cemented no-
dules 0.00013 0.04246 0 0 29.77139
Graules, Pebbles & Carbo-
nate cemented nodules 0 0 0 0.688 13.91762
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.
12 Appendix
51
Ap
pen
dix
II.
XR
F v
alues
of
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ents
fro
m d
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eds
at t
he
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ite
in Å
sen.
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3.3
5
1.9
5
1.5
1
.35
1.0
5
0.5
0
.125
0
-1
-20
-25
Si
13.0
55
697
9.4
369
77
14.9
13
972
15.5
48
964
24.3
65
603
21.7
38
582
33
26.3
93
706
24.9
37
070
5
25.3
66
011
32.0
25
016
33.1
83
625
26.7
42
253
SiO 2
27.9
30
052
59
20.1
88
524
9
31.9
05
460
3
33.2
63
898
69
52.1
25
334
5
46.5
05
349
19
56.4
64
055
25
53.3
47
874
92
54.2
65
507
33
68.5
11
116
73
70.9
89
728
96
57.2
09
701
84
Ti
0.0
924
7
0.0
800
52
0.1
096
86
0.1
066
54
0.2
389
24
0.1
090
996
67
0.2
090
13
0.1
207
55
0.3
095
5
0.2
982
89
0.3
975
89
0.4
276
08
TiO 2
0.1
542
677
01
0.1
335
507
52
0.1
829
891
54
0.1
779
308
68
0.3
985
969
09
0.1
820
109
74
0.3
486
963
88
0.2
014
555
67
0.5
164
222
65
0.4
976
355
39
0.6
632
977
29
0.7
133
784
26
AL
0.9
527
6
0.5
776
16
0.7
313
35
1.2
528
65
3.9
136
6
1.1
214
413
33
1.5
742
1
1.5
113
17
5.2
486
38
5.2
852
13
6.6
674
17
9.5
438
6
Al2
O3
1.8
002
400
2
1.0
914
054
32
1.3
818
574
83
2.3
672
884
18
7.3
948
605
7
2.1
189
633
99
2.9
744
697
95
2.8
556
334
72
9.9
173
015
01
9.9
864
099
64
12.5
98
084
42
18.0
33
123
47
Fe
0.3
983
51
0.4
558
81
0.6
665
92
0.8
360
67
1.1
877
05
0.9
795
11
1.0
682
72
1.8
150
145
0.6
440
53
1.4
184
4
0.6
020
22
0.4
840
12
Fe2 O3
0.4
426
874
66
0.5
066
205
55
0.7
407
836
9
0.9
291
212
57
1.3
198
965
67
1.0
885
305
74
1.1
871
706
74
2.0
170
256
14
0.7
157
360
99
1.5
763
123
72
0.6
690
270
49
0.5
378
825
36
Ca
27.1
13
387
5
24.2
54
061
15.7
40
911
17.9
59
095
7.4
289
47
9.0
987
15
8.9
518
23
4.8
884
31
0.4
858
83
5.8
109
82
1.4
860
84
0.2
832
95
CaO
3
7.9
37
051
79
33.9
36
282
15
22.0
24
682
67
25.1
28
365
72
10.3
94
582
64
12.7
30
922
03
12.5
25
390
74
6.8
398
926
55
0.6
798
474
94
8.1
307
260
14
2.0
793
287
33
0.3
963
863
64
K
0.9
135
965
0.7
346
2
1.1
574
3
1.3
195
31
1.9
615
64
1.7
600
31
3.1
459
08
2.5
433
125
2.3
669
31
2.4
455
75
2.9
432
97
2.1
686
11
K2
O
1.1
005
183
44
0.8
849
232
52
1.3
942
401
78
1.5
895
070
43
2.3
628
999
94
2.1
201
333
43
3.7
895
607
77
3.0
636
742
38
2.8
512
050
83
2.9
459
396
45
3.5
454
955
66
2.6
123
088
11
P
0.4
156
13
0.4
105
04
0.2
065
85
0.2
693
41
0.1
220
57
0.0
488
053
33
0
0.0
604
505
0.1
494
02
0.0
588
68
0.2
079
71
0.1
651
95
P2
O 5
0.9
523
356
28
0.9
406
288
66
0.4
733
688
69
0.6
171
679
67
0.2
796
814
1
0.1
118
325
41
0
0.1
385
162
76
0.3
423
397
43
0.1
348
901
35
0.4
765
447
49
0.3
785
278
23
Ba
0.0
202
925
0.0
109
86
0.0
189
63
0.0
043
14
0.0
195
84
0.0
140
33
0.0
265
76
0.0
122
145
0.0
234
45
0.0
199
53
0.0
201
43
0.0
411
24
BaO
0
.0226
565
76
0.0
122
658
69
0.0
211
721
9
0.0
048
165
81
0.0
218
655
36
0.0
156
678
45
0.0
296
721
04
0.0
136
374
89
0.0
261
763
43
0.0
222
775
25
0.0
224
896
6
0.0
459
149
46
Bal
56.9
30
956
5
63.9
41
894
66.3
69
538
61.9
58
125
60.6
73
569
65.0
32
204
67
57.5
30
75
63.9
99
541
64.3
65
181
52.1
65
709
54.0
29
6
59.9
42
544
Mo
0.0
003
1
0
0.0
002
98
0.0
003
72
0
0.0
001
406
67
0.0
010
24
0.0
001
585
0.0
007
35
0.0
005
67
0.0
007
93
0.0
006
03
Th
0.0
008
595
0.0
006
28
0.0
007
67
0.0
006
98
0.0
009
66
0.0
010
03
0.0
161
82
0.0
006
575
0.0
010
17
0.0
016
09
0.0
015
58
0.0
024
34
Zr
0.0
112
375
0.0
234
16
0.0
281
82
0.0
192
76
0.0
213
99
0.0
257
856
67
0.0
106
85
0.0
218
35
0.0
312
06
0.0
382
95
0.0
465
71
0.0
448
04
Y
0.0
014
285
0.0
013
2
0.0
012
13
0.0
011
47
0.0
018
14
0.0
008
823
33
0
0.0
007
225
0.0
011
26
0.0
021
52
0.0
013
79
0.0
018
08
Sr
0.0
201
955
0.0
182
1
0.0
136
27
0.0
144
01
0.0
121
37
0.0
088
563
33
0.0
053
85
0.0
084
89
0.0
024
51
0.0
145
67
0.0
050
65
0.0
026
59
Rb
0.0
018
78
0.0
014
09
0.0
021
59
0.0
024
31
0.0
038
17
0.0
040
186
67
0.0
048
71
0.0
043
33
0.0
043
11
0.0
045
37
0.0
049
1
0.0
056
68
Zn
0
0.0
010
57
0.0
007
77
0
0.0
016
77
0.0
004
13
0.1
320
65
0.0
010
175
0.0
012
07
0.0
012
28
0.0
012
73
0.0
023
5
52
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 Sr/Ca
Si Ca Sr 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
53
Phragmocone Rostrum solida
SAMPLE Si Ca Sr Sr/Ca
Si Ca Sr 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
54
Phragmocone Rostrum solida
SAMPLE Si Ca Sr Sr/Ca
Si Ca Sr 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
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